{

PIIHET PEIN LEAVY US ON, Be 3 ce Se oh Ok Reece elo aaa eam

NLILSNI

BRARIES

NOILNLILSNI

NOIL

NOILNLILSNI

NVINOSHLINS SJIyvuaty

RARITIES SMITHSONIAN se

;yYVvudgi

NYINOSHLIWS

INSTITUTION

| | | NOILN

INSTITUTION

OSHLINS S314¥VHN9IT LIBRARI

VINOSHLINS S3INWHSIT LIBRAR ies

VINOSHLINS $3

INSTITUTION NOLLMLILSNI

SMITHSONIAN

NYIN

= LIBRA

S SMITHSONIA

AR1ES” SMITHSONIAN NVINOSHIINS S31NVu GIT

RARIES

UNLILSNI

NOILALILSNI

LIBRARIE

NYINOSHLIWS SMITHSONIAN” INSTITUTION

3 “ily ;

fi. “a a

Ca ; {5 he

“i Pode ais

‘Ye ae tease

INSTITUTION NOILOLILSNI_ NVINOSHLIWS

Pet

INSTITUTION NOILALILSNI

INSTITUTION

SJIYVUd}

INSTITUTION

Saliuvuda

saruvual

YINOSHLIWS

SMITHSONIAN

NYINOSHLI

| NVINOSHLIWS

_NVINOSHLIWS LIBRARIES

)

PALILSNI

NOILNLILS

NOILALILSN

LIBRARIES

NOILALILSN

INSTITUTION NOILOLILSNI

ARIES SMITHSONIAN NVINOSHLINS S31uWUudIT LIBRA

SJIMVUGIT LIBRARIES

INSTITUTION

‘i

S S3INYVYNSII LIBRARIE

PALILSNI saiuvugi ,SMITHSONIAN INSTITUTION NOILMI

fs .

alle

SON]

* Cs

" 4 va

: *,

INSTITUTION NOILNLILSNI

NVINOSHLINS S31Y¥Vudl

NVINOSHLIW

NVINOSHLIWS

: *, “

NVINOSHLIWS S31uVYdI1

S SMITH

RARIES SMITHSONIAN

NOILALILS

(Ag =

LIBRARIES SMITHSONIAN

LIBRARIES

NOILNLILSNI

NOILMLILSNI

LIBRARIE

de f }

ce +t mane REFER ERODED RET mT © eee tr mse cim a ee ae ee Ge ne Oca ae Ce eee ol eam gee? eae OLD é a een ga aed

Contributions

of the

American Entomological Institute

Volume 7, Number 1, 1970

aN HSON

SEP 2 2 1970

L/BRARIES

MOSQUITO STUDIES (Diptera, Culicidae)

XXII. A new subgenus and species of Aedes from Arizona.

By Thomas J. Zavortink

XXIII. Additions and corrections to nes revision of the

Aedes terrens group. By Robert X. Schick

CONTRIBUTIONS

of the

AMERICAN ENTOMOLOGICAL INSTITUTE

The Contributions are for larger papers on insects. - Each paper is a separate number, with

separate pagination and index. Separate numbers aggregating about 500 pages constitute a volume.

Issues appear irregularly, as suitable manuscripts are available.

Copies are sold separately or in subscriptions to complete volumes. Complete volumes are

$12.00. The price of separate numbers varies. Subscribers are billed for each volume with its

beginning number, and receive the parts as issued. Orders for separate numbers that total less

than $8.00 must be accompanied by payment. Address orders or correspondence to the American

Entomological Institute, 5950 Warren Road, Ann Arbor, Michigan 48105, U.S.A.

Parts of Volume 1, with prices

No. 1. Dasch, Clement E., 1964. The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae).

75 pages, 69 figures. Price: $2.25, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1965.

I. Belkin, John N., etal. A project for a systematic study of the mosquitoes of

Middle America. 8

Il. Belkin, John N., et al. Methods for the collection, rearing and preservation of

mosquitoes.

78 pages, 4 figures. Price: $2.25, postpaid.

No. 2a. Same as no. 2, but in Spanish. Price: $2.25, postpaid.

No. 3. Matthews, Robert W., 1965. The biology of Heriades carinata Cresson. (Hymenoptera,

Megachilidae). 33 pages, 23 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1965.

Ill. Ramalingam, Shivaji, and John N. Belkin. Two new Aedes from Tonga and Samoa.

IV. Belkin, John N. The mosquitoes of the Robinson-Peabody Museum of Salem

expedition to the southwest Pacific, 1956.

34 pages, 3 figures. Price: $1.00, postpaid.

No. 5. Mosquito studies (Diptera, Culicidae), 1965.

V. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from Middle America. 95 pages. Price: $2.75 postpaid.

No. 6. Mosquito studies (Diptera, Culicidae), 1966.

VI: Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from North America. 39 pages. Price: $1.00, postpaid.

No. 7. DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae

(Homopotera, Cicadellidae). A synopsis of the genus Ponana.

86 pages, 257 figures. Price: $2.50, postpaid.

No. 8. Contributions to the mosquito fauna of southeast Asia, 1967.

I. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald in

Thailand. 55 pages, 20 figures. Price: $1.75, postpaid.

Parts of Volume 2, with prices

No. 1. Contributions to the mosquito fauna of southeast Asia, 1967.

II. Bram, Ralph A. The genus Culex in Thailand (Diptera: Culicidae).

296 pages, 104 figures. Price: $9.00, postpaid.

No. 2. Schuh, Toby, 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region.

35 pages, 30 figures. Price: $1.25, postpaid.

No. 3. Snider, Richard J., 1967. The chaetotaxy of North American Lepidocyrtus s. str.

(Collembola: Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid.

No. 4. Contributions to the mosquito fauna of southeast Asia, 1968.

II. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald

of southeast Asia. 76 pages, 26 figures. Price: $2.25, postpaid.

No. 5. Contributions to the mosquito fauna of southeast Asia, 1968.

IV. Knight, Kenneth L. Species of the subgroup Chrysolineatus of group D, genus

Aedes, subgenus Finlaya Theobald. 45 pages, 12 figures. Price: $1.50, postpaid.

Parts of Volume 3, with prices

No. 1. Mosquito studies (Diptera, Culicidae), 1968.

VII. Belkin, John N. The Culicidae of New Zealand.

182 pages, 30 figures. Price: $5.50, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1968.

VIII. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia.

221 pages, 41 figures. Price: $6.75, postpaid.

No. 3. Fleming, Richard C., 1968. Head musculature of sphinx moths (Lepidoptera:

Sphingidae). 32 pages, 37 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1968.

IX. Belkin, John N. The type specimens of New World mosquitoes in European

museums. 69 pages. Price: $2.00, postpaid.

(Continued on back cover)

MOSQUITO STUDIES (Diptera, Culicidae)

XXII. A NEW SUBGENUS AND SPECIES OF AEDES

FROM ARIZONA’

Thomas J. Zavortink”

In June 1969, John F. Burger of the University of Arizona sent a single female

of a very unusual mosquito with a mesonotal pattern superficially similar to that

of Aedes aegypti to John N. Belkin at the University of California at Los Angeles.

The specimen had been collected in the Coyote Mountains, Pima County, Ari-

zona, by Martha L. Noller, also of the University of Arizona. Several members

of the ‘Mosquitoes of Middle America” project staff examined the specimen and

concluded that it was an undescribed species of Aedes so distinct from other known

New World forms that it should probably be recognized at the subgeneric level.

From the highly ornamented nature of the adult it was surmised that the imma-

ture stages would be found in treeholes or rockholes. After both Mr. Burger and

Miss Noller kindly provided additional information about Mendoza Canyon, where

the mosquito had been taken, and furnished directions for getting to the area,

I visited the region and collected for a few days during the first week in Septem-.

ber 1969. Although mosquito larvae and pupae were collected from numerous tree-

holes and rockholes and many biting adult mosquitoes were caught, the new form

was not found. In late December 1969, Lewis T. Nielsen of the University of Utah

and I revisited Mendoza Canyon and collected water and debris from a large num-

ber of treeholes. The material which we gathered was divided and portions of it

were reared at the University of Utah and portions at the University of California

at Los Angeles. Although no specimens of the unusual new species were recov-

ered from the material reared in Utah, 2 males and 3 females with associated

larval and pupal skins were reared in Los Angeles from eggs taken from an oak

treehole by Dr. Nielsen. Study of these additional specimens, particularly the im-

mature stages and males, supports the original conjecture that this new species

should be placed into a separate subgenus of Aedes. The new subgenus, Abraedes,

is briefly diagnosed and the species, papago, is fully described below.

The limited material presently available does not permit preparation of slides

of adults and, consequently, no drawings of adult morphology are presented here.

It is hoped that such drawings can be included in a future revision of the New

World treehole and rockhole Aedes.

‘Contribution from project “Mosquitoes of Middle America’”’ supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478.

* Department of Zoology, University of California, Los Angeles, California 90024.

Zz Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

I am grateful to John N. Belkin for his counsel in the preparation of this paper.

I thank William A. Powder for help in the rearing and preparation of specimens,

L. Margaret Kowalczyk for preparation of the preliminary and final illustrations

and Caryle Abrams for typing the preliminary drafts and the final copy for litho-

printing. It is a pleasure to acknowledge the invaluable help of John F. Burger

and Martha L. Noller of the University of Arizona and Lewis T. Nielsen of the

University of Utah; without their cooperation, as noted above, the present study

would not have been possible.

ABRAEDES, new subgenus

TYPE SPECIES: Aedes (Abraedes) papago, n.sp.

FEMALE. Highly ornamented; integument of head and thorax dark brown to

black, strongly contrasting with yellowish to amber integument of abdomen.

Head: Eyes broadly separated above antennae, the space with broad flat sil-

very scales. Decumbent scales of vertex broad and flat. Erect scales of vertex few

in number. Proboscis conspicuously longer than forefemur. Palpus short, 0.26-0.29

length of proboscis; apparently 4-segmented. Antenna subequal to proboscis in

length.

Thorax: Long rows of acrostichal and dorsocentral bristles present. Mesonotum

with narrow scales; dark with conspicuous pattern of white and silvery lines. su-

perficially similar to that of Aedes aegypti. Scutellar lobes with patches of large

broad flat silvery scales. Paratergite moderately broad; with broad flat silvery scales

in lower anterior portion. Pleural bristles absent on lower mep, few on stp. Pleu-

ron with broad flat silvery scales in small dense patches or lines.

Legs: Disproportionately short. Tarsi conspicuously marked with white bands

or rings. Claws simple.

Wing: Plume scales present on dorsum of veins Rs, R,+3, R2, R3, middle 0.60-

0.70 of M and sometimes apical portion of IA.

Abdomen: Distal tergites and sternites with dense lateral patches of outstand-

ing black and iridescent silvery scales; tergites VI and VII with large bristly sub-

median apical scaleless area. Sternite VIII large, exserted, nearly devoid of scales,

but with numerous setae. Genitalia deeply retracted, only dark tips of cerci pro-

jecting from sternite VIII. 3

MALE. Similar to female except for sexual characters. Palpus subequal to pro-

boscis in length; 5-segmented; segments 4 and 5 bent ventrad. Antenna slightly

shorter than proboscis; torus enlarged; flagellum strongly plumose. Larger claw of

foreleg and midleg toothed. Apical abdominal segments and genitalia bent ven-

trad. Tergite VIII short, only 0.70-0.78 length of sternite VIII; largely retracted;

with only a few apical black scales.

MALE GENITALIA. Tergite [IX very poorly developed, short, without lobes;

1 or 2 fine submedian setae on each side. Sidepiece without basal or apical lobes;

sternomesal surface without specialized scales. Claspette filament developed, long,

simple. Aedeagus subparallel-sided in basal half, bulbous in distal half in dorsal

aspect. Proctiger unusually long in dorsal aspect; without the normal dorsal lobe

of the basolateral sclerotization.

PUPA. No cephalothoracic hairs elongate. Trumpet short; tracheoid virtually ab-

sent. Hair 1-II farther from midline than 1-I. Hair 2-I-VII relatively long and strong;

Zavortink: New Subgenus of Aedes 3

usually mesocephalad of hair 1 on segments III-V. Hair 5-[V-VI shorter than cor-

responding tergite. Hair 9-II-VI relatively long and strong, becoming longer on pos-

terior segments; usually cephalad of level of hair 6 on segments II-IV, cephalad

or caudad of level of hair 6 on V, caudad of level of hair 6 on VI. Hair 9-VIII

relatively far mesad of caudolateral angle of segment. Paddle deeply emarginate,

outer margin distad of external buttress and inner margin with long filamentous

spicules. |

LARVA. Length and width of head subequal. Labial plate subquadrate. Hairs

5,7-C at about level of antennal base. Hairs 5-7-P on separate tubercles. Hair 2-

III-V usually mesocephalad of hair 1. Hair 3-VII relatively short, not reaching base

of siphon. Hair 12-I absent. Hair 13-VI relatively long, usually double or triple

(2-4), laterad and usually caudad of 10-VI. Comb scales large; few (4 or 5); con-

sisting of an elongate to oval plate from which 1 or rarely 2 large unfringed spines

and several smaller spines project. Siphon short; without acus. Caudal margin of

anal saddle without strong spines. Ventral brush weakly developed; with 5 pairs

of hairs from irregular basal boss; hair 4a-X short, multiple.

DISCUSSION. See papago below.

Aedes (Abraedes) papago, n.sp.

Figs. 1,2

TYPES: Holotype male (UCLA 550-39) with slides of associated larval and pupal skins and

genitalia, Mendoza Canyon, Coyote Mountains, Pima County, Arizona (31°59' N, 111°30' to

111°31' W), elevation about 1070 m, egg obtained from a rothole in a living evergreen oak tree,

28 Dec 1969, L.T. Nielsen [USNM]. Allotype female (UCLA 550-33) with slide of associated

larval and pupal skins, same data as holotype [USNM] . Paratypes: 1 lpM (550-36), 2 lpF (550-

37,38), same data as holotype [UCLA]; 1 F, E slope Coyote Mts., Pima Co., Arizona, 3500 ft,

13 Aug 1968, M.L. Noller [ARIZ].

FEMALE. Wing: 3.15 mm. Proboscis: 2.06 mm. Forefemur: 1.65 mm. Abdo-

men: about 3.2 mm.

Head: Integument dark brown to black. Frontal bristles absent. Orbital bristles

10-13 pairs. Ocular border of broad flat silvery scales. Ornamentation complex,

consisting of white median longitudinal line bordered on each side by 3 progres-

sively smaller black patches alternating with 3 progressively smaller white to silver-

white patches; posterior portion of submedian black patch usually with some white

scales; occiput with narrow curved white scales. Underside of head usually with

silver-white scales. Erect scales of vertex largely restricted to anterior portion of

submedian dark patch, black, some elongate and narrow, some proclinate; erect

scales of occiput numerous, entirely or largely black, moderately long and broad.

Clypeus moderate in size, bare. Proboscis scales predominantly black with metal-

lic coppery to green reflections; white speckles in middle portion, especially dor-

sally. Palpus black scaled with white patch at base of segment 3; white patch

covering all of segment 4; usually with white speckles or patch on segment 2.

Torus with large dorsomesal patch of broad silvery scales; flagellar segment | with

mesal line of white scales.

Thorax: Integument largely dark brown to black. Mesonotum with numerous

long, strong, dark bristles; humeral, lateral prescutal, posterior fossal, supraalar and

parascutellar bristles present. Scutellum with 4-6 well developed bristles on mid-

4. Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

lobe and 3-7 well developed bristles on lateral lobe. Mesonotal background of small

dark brown to black curved scales which become larger and broader laterally and

posteriorly. Conspicuous pattern of narrow white or silvery lines as follows: (1) a

long para-acrostichal line of narrow white to silver-white scales from anterior pro-

montory to near level of wing root; becoming narrower and slightly converging

towards its mate posteriorly, (2) a more or less continuous line of enlarged sil-

very scales from humeral angle to scutellum via scutal suture and outer posterior

dorsocentral line; scales denser in posterior part of line; line not broadened in

area of scutal angle, (3) a nearly transverse patch of large silvery scales in front

of wing root, (4) a short median acrostichal line of white to silver-white scales

from about level of end of para-acrostichal line to prescutellar bare space, (5) a

tuft of silver-white scales on anterior promontory, (6) an irregular patch of large

silvery scales behind scutal angle, and (7) sometimes enlarged silvery scales in a

patch at anterior end of prescutellar bare space or in a narrow lateral prescutellar

line. Pleuron with bristles on apn, ppn, ppl, psp, stp, pra and upper mep; ssp

bristles absent. Pleural scales forming a long more or less continuous silvery diag-

onal line from upper anterior apn across lower ppn, ssp and upper middle stp to

lower posterior portion of mep; additional silvery patches on upper and lower

pst, ppl, upper portion of pcx, pra, lower posterior stp and upper mep; longer

nearly horizontal silvery line across middle portion of stp; ppn usually with a few

silvery scales, sometimes bare; psp and metameron usually without scales.

Legs: Integument of coxae dark brown to black. Coxal bristles normal. All cox-

ae with patches of broad flat silvery scales, as follows: (1) forecoxa with upper

and lower anterior patches, (2) midcoxa with small upper posterior, middle an-

terior and lower anterior patches, (3) hindcoxa usually with large patch from up-

per portion to middle of anterior surface and small lower anterior and lower pos-

terior patches; forecoxa with black scales between silvery patches. All trochanters

with black and white scales. Femora predominantly black scaled on all surfaces

of all legs; many black scales, particularly on midfemur and hindfemur, with me-

tallic silvery, light green, blue or violet reflections; forefemur with incomplete nar-

row subbasal oblique white band and numerous scattered white scales; midfemur

and hindfemur with usually complete narrow subbasal oblique white ring and few-

er scattered white scales. Knee spots small, usually incomplete on forefemur. Tib-

iae predominantly black scaled, with slight metallic green, blue or violet reflec-

tions; all tibiae with narrow to moderately broad dorsally incomplete basal white

band; foretibia and midtibia with narrow white ring or band 0.25-0.33 distance

from base; hindtibia with broader white ring 0.45-0.50 distance from base. Tarsi

black scaled, with conspicuous white dorsal patches, bands or rings, as follows:

(1) foretarsus with moderately broad patch or band at base of segment | and

smaller patch at base of segment 2 and usually segment 3, (2) midtarsus with

moderately broad basal band or ring on segment 1 and small patch at base of

segment 2 and sometimes segment 3, and (3) hindtarsus with moderately broad

subbasal white band or ring on segment 1, moderately broad basal band or ring

on segments 2 and 3 and moderately broad basal patch or band on segments 4

and 5. Claws of all legs subequal in size, small, simple.

Wing: Dark scaled except for a white line in basal 0.16-0.20 of anterior sur-

face of costal vein.

aa Integument light tan. Stem black scaled distally. Knob black and white

scaled.

Abdomen: Integument predominantly very light, yellowish white to amber. Sil-

Zavortink: New Subgenus of Aedes y

very and, to lesser extent, dark scales with light metallic coppery, green, blue or

violet reflections. Tergite I with numerous dark scales dorsally; laterotergite with

large patch of silver-white scales. Tergites II-VIII predominantly dark scaled, with

lateral basal or subbasal silvery patch followed distally by lateral patch of black

scales; scales of lateral patches becoming progressively denser and more outstand-

ing on distal segments; tergites VI and VII with outstanding black scales middor-

sally; tergite VIII with dense outstanding black scales and long apical setae which

project beyond apices of cercal setae. Sternite I bare. Sternites [I-IV variably scaled,

sometimes largely bare, usually with at least a few dark or black scales apically,

sometimes with small lateral subbasal or median silvery patch. Sternites V-VII us-

usally at least partly bare basally, black scaled apically and with lateral median

silvery patch; scales becoming progressively denser and more outstanding on distal

segments.

MALE. Essentially as in female except for sexual characters.

Head: White speckling of proboscis reduced or absent. Palpal segments 2 and

3 ankylosed, long, making up about 0.67 length of palpus; segment 4 short, about

0.16 length of palpus; segment 5 short, about 0.12 length of palpus; slender ex-

cept for slightly swollen apex of segment 3; apex of segment 3 and all of segments

4 and 5 with bristles; dark scaled except for conspicuous dorsal white patch or

band near base of segment 2 and at base of segments 3-5. Torus with silvery scales

dorsomesally; flagellar segment 1 with small scales.

Legs: Anterior foreclaw and midclaw large, with a single long slender subbasal

tooth; posterior foreclaw and midclaw medium sized, simple; hindclaws small, sim-

ple.

Abdomen: Scaling of sternite VIII similar to tergite VIII of female.

MALE GENITALIA (fig. 1). Unusually colored, segment IX, proctiger, phallo-

some, claspette and clasper largely very weakly pigmented, yellowish; sidepiece

and spiniform more deeply pigmented, light brown; apex of paraproct black.

Segment IX: Tergite without the normal integumentary spicules middorsally.

Sternite well developed, long, with numerous strong setae distally.

Sidepiece: Well developed, relatively long and slender, more or less spindle shap-

ed. Mesal surface membranous from base to apex. Basal tergomesal area not swol-

len, without clumped or enlarged bristles, but with normal bristles of dorsal sur-

face of sidepiece shorter and slightly more numerous. Median sternomesal sclerite

not developed. Distal dorsomesal surface with short setae; dorsal surface, distal

lateral surface and ventral surface with long bristles. Dorsal surface with dark scales

among bristles; basal portion of dorsolateral and lateral surfaces with dense broad

silver-white scales with iridescent coppery, green, blue or violet reflections; distal

portion of lateral surface with dense broad dark scales.

Claspette: Well developed. Stem moderately long, bent mesally in distal portion

in dorsal aspect. Base and all but distal portion of stem spiculose. Mesal and meso-

ventral surface of stem with 3-6 weakly developed setae. Filament more or less

evenly curved dorsally, subterete; without a ridge or retrorse barb on convex side.

Clasper: Simple, moderately long, curved inward distally. Broadest at base, tap-

ering evenly to narrower apex. Dorsal basal portion spiculose. Surface not con-

spicuously wrinkled. Ventral surface with 2 setae near apex. Apical spiniform mod-

erately long, about 0.30-0.35 length of clasper, slightly curved inward.

Phallosome: Aedeagus moderately long, without teeth. Ventral paramere appear-

ing unusually short and broad in dorsal aspect.

Proctiger: Strongly developed. Paraproct with single large heavily sclerotized

6 Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

curved apical spine. Cercal setae fine, short, 2-4.

-PUPA (fig. 1). Abdomen: about 4.0 mm. Trumpet: 0.54 mm. Paddle: 0.81 mm.

Integument very lightly pigmented, uniformly light straw yellow, strongly contrast-

ing with dark trumpet and hairs.

Cephalothorax: All hairs relatively short, stout, darkly pigmented and usually

single or double. Hair 6-C usually as stout as or stouter than 7-C.

Trumpet: Dull brown in color except for slightly lighter apex of pinna. Trach-

eoid represented at extreme base of lateral (ventral) surface only. Index about

3.1-3.3; pinna about 0.22-0.33 of total length.

Metanotum and Abdomen: Integumentary reticulation and spiculation very in-

conspicuous on anterior segments, becoming slightly more pronounced on poster-

ior segments. Most hairs relatively short, stout, darkly pigmented and single or

double. Float hair (1-1) well developed, relatively short, with numerous dendritic

branches; lateralmost branches of float hair usually diverging at an angle of less

than 100 degrees. Hair 1-II-VII relatively short, usually subequally developed and

single or double (1-3) on all segments, but sometimes enlarged and multibranched

(5-7) on segment II. Hair 2 usually laterocephalad of hair 1 on segments II,VI,

VII. Hair 3-I-III subequal in development, moderately long, single (1 or 2); hair

3-IV-VII subequal in development, usually single on V-VII, single or double on

IV. Hair 4-II mesad of 5-I]. Hair 5-II,II] subequally developed, relatively short,

distinctly finer than hair 3 of corresponding segment, usually single (1 or 2); hair

5S-IV-VI subequally developed, subequal to or slightly longer than 3-I-III, about

0.7-0.9 length of corresponding tergite, usually single (1 or 2); hair 5-VII similar

to 5-II,III or slightly longer and/or stronger, usually double (1 or 2). Hairs 6,7-I

much finer and shorter than 3-I. Hair 6-II slightly thickened and elongate, single

(1 or 2); hair 6-III-V usually distinctly shorter and finer than 6-II, usually single

(1-3); hair 6-VI quite stout and elongate, single; 6-VII usually only slightly longer

and stronger than 6-III-V, single. Hair 9-VII long and strong, double or single.

Hair 9-VIII most strongly developed and longest hair of pupa, with 2-4 (2-5) pri-

mary branches. Hair 10 conspicuously more mesad and cephalad of hair 11 on

segment VI than on segment VII. |

Paddle: Shape as figured; length 1.30-1.55 of maximum width. Evenly light straw

yellow in color. Midrib brighter and deeper yellow in color, extending to highly

wrinkled area basad of apical emargination. Hair 1-P relatively long and strong,

single.

LARVA (fig. 2). Head: 0.97 mm. Siphon: 0.72 mm. Anal Saddle: 0.28 mm.

Strongly developed stellate hairs and conspicuous spicules absent.

Head: Integument smooth. Medium straw yellow in color except for slightly

darker anterior portion of ventral surface and much darker collar. Ocular area not

lightened. Mental plate normal, brown, with 8 or 9 (8-10) lateral teeth. Hair 1-C

very stout, widened before middle, smooth. Hairs 4,6-C relatively far cephalad.

Hair 4-C small, with 5-8 branches (4-9), located mesocephalad of hair 6 and mesad

of hair 1. Hair 5-C single, laterad of 6-C. Hair 6-C single, usually slightly widened

beyond base, slightly laterad of 1-C. Hair 7-C usually triple or double (1-3). Hair

11-C weakly developed, short, stellate, with 3-5 branches. Hair 14-C rather stout,

usually triple (2 or 3). Hair 15-C located in anterior 0.25 of labial plate; long, ex-

tending beyond apex of mental plate, with 3 or 4 branches.

Antenna: Relatively short. Integument without spicules, but sometimes with con-

spicuous annular wrinkles. Medium straw yellow in color except for slightly dark-

ened base. Hair 1-A usually single; unusually short, length only 1.5-2.0 diameter

Zavortink: New Subgenus of Aedes 7

of antennal shaft at insertion of hair; inserted on dorsal or dorsomesal surface of

shaft.

Thorax: Living larva white, without conspicuous pigmentation in epidermis or

fat body. Integument without spicules. Hairs moderately pigmented. Hairs 1-3-P

sometimes arising from a weakly sclerotized common plate. Hair 3-P much shorter

and weaker than 1-P. Hair 4-P with 3-5 branches. Hairs 5,7-P long, usually with

4 or 5 branches (2-6), each sometimes with small basal plate. Hair 8-P relatively

short, usually with 4-6 branches (3-7). Hair 11-P,M,T short, weak, usually double

(1-3), much smaller than 9-P. Hair 1-M shorter than 3-M, with 2-4 branches. Hair

5-M double or single. Hair 14-M short, single to triple. Hairs 1,13-T subequal, small,

usually with 2-4 branches (1-4). Hair 4-T usually longer and stronger than 1-T.

Hair 5-T small, single to triple.

Abdomen: Coloration and spiculation as for thorax. Most hairs moderately pig-

mented. Hair 1-I,II subequally developed, moderate, usually with 3 or 4 branches

(2-7); hair 1-III-V subequally developed, slightly longer than 1-I,IJ, with 3 or 4

branches; 1-VI,VII subequally developed, longer than 1-III-V, usually with 3 or 4

branches (3-5). Hair 2-I-VII relatively stout, usually triple (2 or 3) on III, usually

single or double (1-3) on III-V, single on VI,VII; usually located laterocephalad

of hair 1 on I,II,VI and VII. Hair 5-I-VII subequally developed, moderate, similar

to 1-III-V, usually with 3 or 4 branches (2-6). Hairs 6,7-I with separate basal plates.

Hair 6-I-V long, double or triple on I,IJ, double on III-V; hair 6-VI shorter, single

or double. Hair 7-I long, 7-II-VI usually becoming progressively shorter; usually

double (1 or 2) on I, double or triple (1-3) on II-VI. Hair 11-I moderately devel-

oped, usually with 4-7 branches (3-8). Hair 13-I relatively small, usually with 3

or 4 branches (1-4); hair 13-II larger, subequal to 11-I, usually double or triple

(1-3); hair 13-II]-V subequally developed, long, much longer than hair 1 of cor-

responding segment, usually triple (2-4). :

Segment VIII: Integument smooth except for rows of inconspicuous spicules

in area of comb scales. Hair 1-VIII moderately developed, usually with 4 or 5

branches (3-5), frequently arising from a small basal plate. Hair 3-VIII large, usu-

ally with 6 or 7 branches (5-8), arising from basal plate.

Siphon: Short, index apparently about 1.9-2.3 (determined from skins). Light

yellow brown to light brown in color except for darker base, especially dorsally,

and apex. Integument smooth to very weakly and indistinctly imbricate. Basal mar-

gin of siphon irregular and wavy. Pecten teeth moderate in size; usually 12-14

(12-18); brown except for hyaline apex; usually with 2-4 basal denticles. Pecten

in straight, slightly curved or slightly sinuous row in basal 0.32-0.42 of siphon;

often 1 or more of the basal teeth displaced dorsad or ventrad of row; no apical

teeth detached. Hair 1-S arising beyond pecten row, about 0.38-0.52 distance from

base; moderately developed, usually triple or double.

Anal Segment: Saddle moderate in size, extending to or slightly beyond middle

of segment; ventral edge frequently irregular; more or less concolorous with si-

phon; integument smooth to weakly imbricate on body of saddle, slightly spicu-

lose near distal margin dorsolaterally. Hair 1-X arising on saddle near apicoventral

angle; strongly developed, relatively long, usually triple or double (1-4). Hair 2-X

long, usually with 3 or 4 branches (3-7). Hair 3-X very long, single. Hair 4a-X

short, usually with 4 or 5 branches (4-6); hairs 4b,c-X long, double or single; 4d-

X moderately long, single; 4e-X short, double or single. Dorsal and ventral gills

subequal in length, very long, at least 3.0-4.2 length of anal saddle (determined

from skins); sausage-shaped; gills contiguous in living larva until shortly before pu-

8 Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

pation, when they become spreading.

SYSTEMATICS. This species is one of the most remarkable container breeding

Aedes of the New World. The adult characters, in particular, are very striking.

The diagnostic features of all stages are set forth above in the subgeneric description.

This species shows no obvious relationships to any other. It is apparently an-

other one of the many unusual New World container breeding Aedes, most of

which are currently placed, incorrectly in my opinion, in the’subgenus Finlaya.

The resemblance to the Old World subgenus Stegomyia is superficial only.

The living larva is white, rather elongate and is easily distinguished from others

found in treeholes in southeastern Arizona by the apposition of its 4 long anal

gills. The living pupa is macroscopically separated from others of the area by its

weak movements and by the dark trumpets and float hairs which contrast strong-

ly with its very light integument.

The only striking variation observed in any stage of the small series of speci-

mens available is the development of abdominal hair 1-II of the pupa. This hair

is usually short, single or double and developed similarly to 1-III-VII, but on both

sides of 1 specimen and on 1 side of another, the hair is enlarged and has 5 to 7

branches.

BIONOMICS. The original adult female of papago was caught while attempting

to bite. The larvae apparently occur in treeholes. The reared specimens were hatch-

ed from eggs unknowingly collected along with a relatively small amount of water

and organic sludge from a small rothole in a living evergreen oak tree (Quercus).

The nature of the treehole prevented scraping the cavity or collecting large pieces

of debris in a deliberate attempt to obtain aedine eggs. Only a single young instar

Orthopodomyia larva, presumably kummi, was present in the water at the time

of collection. Upon arrival at the laboratory in Los Angeles the water from the

rothole was placed into an enamel pan and diluted with tap water. A few aedine

larvae appeared within the next few days, but all of them and the Orthopodomyia

larva died when oil from an overhead refrigerating unit dripped into the pan. The

pan was moved to a more favorable position in the rearing room and the water

evaporated. After the debris had been dry for 7-10 days the pan was filled with

distilled water. Several eggs hatched within the next 1 or 2 days. Most of the

larvae, an undescribed species in the Aedes kompi group, matured rapidly. When

the unusual attitude of its long gills was noticed macroscopically, the | slow-grow-

ing larva was examined microscopically and determined to be an unknown spec-

ies. This was the first larva of papago. The debris was permitted to dry again and,

after several days, was reflooded with distilled water and subjected to a vacuum

of 20-25 inches of mercury for 30 minutes. Several additional eggs hatched; again,

most larvae were the undescribed species near kompi, but 1 was Aedes (Kompia)

purpureipes. After these larvae were reared, the debris was dried, reflooded with

distilled water and subjected to the vacuum treatment 3 times in close succession

without any additional hatch. Following this series of floodings the debris was

left dry for 6 or 7 weeks. It was then flooded and subjected to the same va-

cuum treatment as before. Quite unexpectedly, numerous Aedes eggs hatched;

among the larvae were more or less equal numbers of purpureipes and the undes-

cribed species near kKompi and fewer of papago. Four of the latter were success-

fully reared. As before, these larvae matured much more slowly than the accom-

panying species. The debris has been subjected to 1 additional drying and reflood-

ing but no additional eggs hatched.

The oak bearing the rothole from which papago was collected was growing in

Zavortink: New Subgenus of Aedes 9

a small lateral canyon formed along a fault in the north-facing wall of Mendoza

Canyon. This oak was at an elevation near that of the main canyon bottom and

was one of the lowest in the area. Since no specimens of papago were obtained

in numerous collections of debris and water made from oaks higher on the canyon

walls, the species may be restricted to the lower edges of the xeric evergreen for-

est. If so, this may explain why it has not been found earlier, for there is a ten-

dency, at least on my part, when looking for mosquitoes, to go to higher eleva-

tions where oaks and other trees are more numerous and more easily accessible.

The relatively long larval life may also restrict the number of treeholes suitable

for breeding of this species.

DISTRIBUTION. At present papago is known from only Mendoza Canyon, Coy-

ote Mountains, Arizona. It undoubtedly occurs in northwestern Mexico and could

be more widespread in southeastern Arizona. where it may have been overlooked

because few attempts have been made to collect treehole mosquitoes at elevations

as low as 1100 m, the approximate elevation of Mendoza Canyon. All material

examined is listed above in the type series.

FIGURES

1. Aedes (Abraedes) papago; male genitalia and pupa

2. Aedes (Abraedes) papago; larva

Fig. |

ABRAEDES

Sf aA a te

i nae

Ne =

y Wie

Pigc

ABRAEDES

papago ¥

UCLA 550

rizona

United States

MOSQUITO STUDIES (Diptera, Culicidae)

XXII. ADDITIONS AND CORRECTIONS TO THE REVISION

OF THE AEDES TERRENS GROUP’

Robert X. Schick”

The present additions and corrections to my recent revision of the Terrens Group

(Schick, 1970a) are based upon collections obtained in Venezuela by a UCLA

team and local collaborators in 1969; in Argentina by Osvaldo Casal, Miguel Garcia

and associates in 1967-1969; in Colombia by Marston Bates in 1941 and 1944;

and from these and other countries in South America by other workers. Described

here are 1 new species, 2 new forms and the hitherto unknown immature stages

and female of braziliensis Gordon & Evans and the male of berlini Schick. Im-

portant new records of previously described species are given, some revisions are

made in the Terrens and Insolitus Subgroups and corrected keys are provided for

the entire group.

The Terrens Group is still poorly known throughout most of South America.

This is especially evident in northern Argentina from where there are several small

collections of adults which do not agree with any of the known species.

The format and methods of this paper (Schick, 1970b) is similar to that of the

earlier one except that (1) complete literature citations are not given for the prev-

iously treated species, (2) in the descriptions of the species only the diagnostic

features are given, the less useful characters being omitted, (3) the length of the

free portion of the midapical comb scales is given to the nearest 0.005 mm rather

than to the nearest 0.001 mm, (4) the L/S to the nearest 0.1 rather than 0.01,

and (5) an anterior view of femur III is included in the illustrations of the species

in which the female is described to indicate the width of the dark bands. The

figure numbers follow the sequence started in Schick (1970a). Figures 1-60 ap-

peared in the latter paper and 61-68 are represented in the present one. The known

geographic distributions are indicated by circumscribing lines on the maps but on-

ly the new records are spotted.

I thank Alan Stone for the loan of material in the U.S. National Museum; T.H.G.

Aitken, Marston Bates and Osvaldo Casal for providing me with supplementary

collection data; Margaret Kowalczyk for the drawings; and Caryle Abrams for the

preparation of the manuscript.

The following are corrections in Schick (1970a):

‘Contribution from project “Mosquitoes of Middle America’ supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478.

* Department of Zoology, University of California, Los Angeles, California 90024.

14 ~ Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

p. 58, line 38. Length of sidepiece should be corrected to 0.27-0.35 mm.

p. 59, line 6. Hair 4-C should read 5-C.

p. 60, line 28. Cundinimarca should read Cundinamarca.

p. 69, line 13. 12 lpd should read 12 Ip.

p. 70, line 21. 352-12 should read 352-13.

Thorntoni Subgroup

2. Aedes (Finlaya) argyrothorax Bonne-Wepster & Bonne

Figs. 2,13,14,62

1920. Aedes argyrothorax Bonne-Wepster and Bonne, 1920:179.

Aedes (Finlaya) argyrothorax of Schick (1970a:36-38).

SYSTEMATICS. Aedes argyrothorax, previously known from the coastal low-

lands of South America, from eastern Venezuela southward into the state of Rio

de Janeiro, Brazil, with 1 possible record from Iquitos, Peru (fig. 2), is now re-

corded from the highlands of Colombia. The Colombian argyrothorax may be dis-

junct from the populations to the east since this species was not taken by the

UCLA team in the northcentral state of Aragua, Venezuela, either along the coast

or in the Cordillera de la Costa.

NEW RECORDS (fig. 62). Material examined: 6 specimens; 2 M, 1 F, | pupa,

2 larvae; 2 individual larval rearings.

COLOMBIA. Meta: Villavicencio se ca 460 m], 1944, M. Bates, 2 IM (42-A, 42-7), 1 pF

(57-AA) [USNM].

Terrens Subgroup

The following taxonomic changes are made in this subgroup: (1) berlini is re-

moved, (2) apollo is synonymized with terrens, and (3) terrens and braziliensis are

redefined. The subgroup consequently comprises 3 distinct species, terrens, bra-

ziliensis and zavortinki. In addition, a new form, the Teresopolis form, is recognized.

With the removal of berlini, 1 variable, the presence or absence of subspiracular

scales, can be eliminated from the subgroup description (Schick, 1970a:39). All

the species have these scales. However, another variable is introduced with the

inclusion of the Teresopolis form. The primary branches of hair 1-I of the pupa

of this form are predominantly single or double, whereas in the other species they

are predominantly multiple.

The character of the color of the broad decumbent scales of the male vertex

can be added to the subgroup description to further distinguish the Terrens and

Insolitus Subgroups. In the former the scales are dark and in the latter they are

silver. This character is of value in determining the species when the mesonotum

is rubbed. These 2 subgroups can be generally separated in the larva by the num-

ber of hairs of the ventral brush. In the Insolitus Subgroup it is 11-13 and in the

Terrens Subgroup, 13-17.

The undescribed form without a basal dark band on femur III mentioned in

Schick: Additions to Aedes terrens Group 15

Schick (1970a:43) is casali, actually a member of the Insolitus Subgroup.

4. Aedes (Finlaya) terrens (Walker)

Figs. 3,7,8,15,16,61

1856. Culex terrens Walker, 1856:429.

1970. Aedes (Finlaya) apollo Schick, 1970a:46-48. NEW SYNONYMY.

Aedes (Finlaya) terrens in part of Schick (1970a:41-43).

Aedes (Finlaya) apollo in part of Schick (1970a:46-48).

FEMALE (figs. 7,15). Vertex with all decumbent scales narrow curved; erect

occipital setae generally pale or dark; complete acrostichal line absent; fossal mac-

ula variously. developed, at fullest extent not interrupted laterally by dark scales

and fairly broad, at least extent reduced to narrow submarginal, longitudinal band

divided into anterior and posterior portions; pra hairs pale or dark; vein C with

silver scales at base.

MALE (figs. 8,15). Vertex with all decumbent scales narrow curved; complete

acrostichal line absent; mesonotal disc not transversely silvered or transversely sil-

vered anteriorly for at most 0.25 length of fossa; fossal macula reaching mesal mar-

gin of fossa or not; pra hairs pale or dark.

MALE GENITALIA (fig. 15). Sidepiece length 0.31-0.38 mm; median sterno-

mesal sclerite of sidepiece weakly developed; prosophallus width 0.11-0.15 mm.

PUPA (fig. 15). Cephalothorax without highly contrasting dark markings on wing

and leg cases; hair 1-I with primary branches predominantly multiple; paddle round-

ed or slightly tapered apically.

LARVA (fig. 16). Head hair 5-C single to triple; 6-C single or double; 11-C

much shorter than 7-C; L/S 2.0-2.4.

SYSTEMATICS. Aedes terrens is redefined here to include those forms of the

Terrens Subgroup in which (1) all the decumbent scales of the vertex are narrow

curved, (2) the mesonotum of the male is not transversely silvered or only very

narrowly so, and (3) a complete acrostichal line is absent. These characters sep-

arate adult terrens from the other species of the subgroup except for the female

of zavortinki. Female terrens can often be distinguished from zavortinki by the

presence of a characteristic lateral dark area on the fossa but this dark area may

or may not be developed in some populations of terrens including those of the

Villavicencio area of Colombia where the 2 species are sympatric. I have identi-

fied female terrens from this region by the generally dark erect scales of the occi-

put, assuming that zavortinki here would show pale erect scales. The closest known

population of terrens shows generally dark scales (Venezuela) and the closest one

of zavortinki, generally pale scales (Panama). Further support for the terrens de-

termination is the occurrence of a male and female terrens in the same lot (57xy).

It should be noted that a female of the zavortinki type with pale erect scales is

unknown from Colombia, but the total number of specimens of the subgroup from

the country is small and male zavortinki is represented by only 2 specimens. The

immature stages of terrens can be separated from those of braziliensis and the

Teresopolis form (see) but apparently not from zavortinki.

A third species of the subgroup, braziliensis, also occurs in the Villavicencio

area. Here both terrens and braziliensis share an unusual character for the Terrens

16 Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

Group, the presence in both sexes of dark rather than pale hairs on the upper

mep. This led me to incorrectly associate the female of terrens and the male of

braziliensis of this region in my previous paper and to describe them as a dis-

tinct species, apollo. I did note, however, that 2 species were possibly involved

(Schick, 1970a:47). Whether the occurrence of these dark hairs in both species

is due to parallelism or hybridization is unknown. The former possibility is sug-

gested by development of this character in the single female of berlini (Insolitus

Subgroup) known from the Villavicencio area but not in the females from the

other parts of its range.

Aedes terrens in my previous revision was recorded from Brazil south of the

Amazon basin southward into northern Argentina. Owing to the generally broad-

ened definition of this species, a northern strip in the Caribbean countries of South

America is now added to this geographic range, but terrens is still unknown from

the intervening Amazon basin.

The widely distributed terrens comprises several distinctive geographic popula-

tions. The majority can be grouped into 2 general types in which there is a con-

cordance of most or all of the following characters.

Type A. Female: Occiput with erect scales generally dark; fossal macula rela-

tively broad, usually not interrupted laterally by dark scales but when so inter-

rupted dark scales forming at most small patch or narrow line; pra hairs dark.

Male: Mesonotum transversely silvered anteriorly; fossal macula coextensive with

fossa; pra hairs dark. Larva: Head capsule with conspicuous fine spicules; hair 5-I

often shorter than 4-I; saddle often extending about 0.5 or less around anal seg-

ment; ventral brush usually with 15,16 hairs (14-17); hair 4a-X with 9-13 branches.

Type B. Female: Occiput with erect scales generally dark; fossal macula inter-

rupted by relatively broad lateral area of dark scales, the macula consequently

forming narrow submarginal longitudinal band; pra hairs usually pale, rarely dark.

Male: Mesonotum not transversely silvered; fossal macula not reaching mesal mar-

gin of fossa; pra hairs pale. Larva: Head capsule without conspicuous fine spic-

ules; hair 5-I subequal in length to or longer than 4-I; saddle extending more than

0.5 around anal segment; ventral brush usually with 13, 14 hairs (13-15; in 1

specimen, 17); hair 4a-X with 7-10 branches.

Type A terrens occurs in northern South America from Colombia eastward in-

to French Guiana and disjunctly in the Salvador area of Brazil and in the General

Enrique Mosconi (Vespucio) area of Argentina and type B in Brazil south of the

Amazon basin southward into northern Argentina (Iguazu) and disjunctly in French

Guiana. Type A would appear to represent the more primitive state since the fos-

sal macula is not reduced and the distribution is more of a relict type. The popu-

lations of type B vary in the branching of hairs 5,6-C in the larva; in some they

are usually single (1 or 2), in others usually double or triple (1-3). Only the form-

er condition is developed in type A larvae.

NEW RECORDS (fig. 61). Material examined: 137 specimens; 20 M, 42 F, 48

pupae, 27 larvae (mounted); 45 individual rearings (25 larval, 20 pupal).

ARGENTINA. Salta: General Enrique Mosconi (Vespucio) [elev. ca 500 m], 18 Feb 1967,

treehole, H. Fernandez, M. Garcia and O. Casal (ARG 601,603), 2 IpM (603-15,17), 3 lpF (601-

10,11; 603-16), 1 pM (603-101), 3 pF (601-102,103; 603-100), 14 L (601-2), 40 L (603-1)

[UCLA]. Tablillas, 7 km from (nearest town General Enrique Mosconi, elev. ca 500 m), 25

Feb 1967, O. Casal and M. Garcia (ARG 618), 1 lpM (618-14), 4 IpF (618-10,12,16,18), 3 pM

(618-104,108,115A), 1 F, 2 p, more than 100 L (618-1); 6 June 1969 (ARG 773), 7 pF (773-

102,103,105,106,110,114,115), 3 L (773-2) [UCLA]. Tucuman: Churqui, 4 Dec 1922, 1 F

(IMR 29) [UCLA].

Schick: Additions to Aedes terrens Group 17

COLOMBIA. Meta: Finca Vanguardia (Schick, 1970a:48). Villavicencio [elev. ca 460 m], 2

June 1941, very low treehole, M. Bates, 1 F (277); June 1942, W. Komp, 1 F (207B-45);

M. Bates, 1 IF (C-57AA), 2 F, 2 p (57-AA); same data, 1 M, 1 F, 21(57xy) [USNM]. Villa-

cencio, river rd to Bosque Ocoa, 1 June 1942, treehole, 1 M (207B-10) [USNM].

FRENCH GUIANA. Guyane: Montabo (nearest town Cayenne), elev. 80 m, 30 Jan 1965,

small treehole, height 1 m, T. Aitken, R. Martinez and A. Guerra (FG 1), 1 IpM (1-10), 1 Ip

(1-11) [UCLA]. Locality not specified: 1944, H. Floch, 2 M, 2 F (207E-1) [USNM].

VENEZUELA. Aragua: El Limon (nearest town Maracay), elev. 600 m, 15 July 1969, small

treehole, height 2 m, J. Pulido and J. Valencia (VZ 211), 2 pM (211-100,101), 2 F (211-1)

[UCLA]. El Ricon Bonita (nearest town Guigue), elev. 500 m, 24 July 1969, small treehole,

height 1 m, J. Pulido and J. Valencia (VZ 266), 1 lpM (266-20), 1 IpF (266-20), 2 Ip (266-

21,23) [UCLA]. Guamitas (8 km S Rancho Grande on rd to Maracay), elev. 700 m, 18 Aug

1969, small treehole, height 2 m, J. Pulido and J. Clavijo, 1 lpM (VZ 373-10) [UCLA]. Ocumare

de la Costa, 2 km N on hwy 8, elev. 100 m, 12 July 1969, small treehole, height 2 m, J.

Pulido and J. Valencia (VZ 178), 1 lpM (178-10), 2 IpF (178-11,12); 28 July 1969, small tree-

hole, height 1 m, J. Pulido and J. Valencia (VZ 272), 1 pM (272-100), 3 pF (272-100,102,120),

1 M, 4 F (272-2) [UCLA]; 15 Aug 1969, large treehole, height 1 m, J. Valencia, 2 lpF (VZ

307-10,50); same data, but small treehole, 2 lpF (307-10,11) [UCLA].

5. Aedes (Finlaya) braziliensis Gordon & Evans

Figs. 3,21,61,63,64

1922. Aedes (Finlaya) oswaldi var. braziliensis Gordon and Evans, 1922:329.

Aedes (Finlaya) braziliensis of Schick (1970a:43-44).

Aedes (Finlaya) apollo in part of Schick (1970a:46-48).

FEMALE (fig. 63). Vertex with an area of broad decumbent scales adjacent

to narrow curved scales of median longitudinal line; occiput with erect scales gen-

erally pale; complete acrostichal line absent; fossal macula well developed, not in-

terrupted by dark scales; pra hairs dark; vein C with silver scales at base.

MALE (fig. 21). Vertex with decumbent scales as in female; complete acro-

stichal line present; mesonotal disc transversely silvered anteriorly for at least 0.5

length of fossa; fossal macula reaching mesal margin of fossa; pra hairs dark.

MALE GENITALIA (fig. 21). Sidepiece length 0.30-0.34 mm; median sterno-

mesal sclerite of sidepiece weakly developed; prosophallus width 0.11-0.13 mm.

PUPA (fig. 63). Wing pad with broad and dark subbasal band; leg cases with or

without dark pigmentation; hair 1-I with primary branches predominantly mult-

iple; paddle slightly tapered apically.

LARVA (fig. 64). Hair 5-C single or double; 6-C single; 11-C much shorter than

We LIS: 25-28.

SYSTEMATICS. Aedes braziliensis, as more broadly interpreted here, comprises

those populations of the Terrens Subgroup in which there is a lateral area of broad

scales on the vertex. The mesonotum of the male is broadly silvered anteriorly as

in zavortinki. The ornately marked cephalothorax of the pupa may separate bra-

ziliensis from the other members of the subgroup but the reliability of this char-

acter cannot be determined without additional material. The larva generally dif-

fers from terrens and zavortinki in the greater L/S. In the Villavicencio area of.

Colombia braziliensis and terrens may be further distinguished by the length of

hair 5-I; in braziliensis it is much shorter than 4-I and in ferrens it is subequal.

18 Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

Aedes zavortinki also occurs in the Villavicencio area but the larva is unknown

from this region.

Aedes braziliensis was previously known only from Brazil and French Guiana

at low elevations (fig. 3). The range is now extended into Colombia at moderately

high elevations. This species, however, was not taken by the UCLA team in an

intervening area, namely the state of Aragua, Venezuela.

NEW RECORDS (fig. 61). Material examined: 31 specimens; 6 M, 5 F, 6 pupae,

13 larvae; 1 individual larval rearing.

COLOMBIA. Meta: Villavicencio [elev. ca 460 m], 1944, M. Bates, 4 1 (24-4), 2 1 (24-78),

1 M, 3 p, 31 (C-24), 1 M, 4 F (24); same data, 1 1 (4268), 2 P (4296, 4298) (C-42-6); Komp, 1 M,

1 F (H-9-10) [USNM]. Villavicencio, Bosque Ocoa, 11 June 1944, 1 M [USNM].

FRENCH GUIANA. Guyane: Cabassou, elev. ca 30 m, 31 Jan 1945, small treehole, height

0.5 m, T. Aitken, R. Martinez and A. Guerra (FG 12,14), 1 IpF (12-14), 1 L (12-1), 1 L (14-1)

[UCLA].

6. Aedes (Finlaya) zavortinki Schick

Figs. 3,8,9,61

1970. Aedes (Finlaya) zavortinki Schick, 1970a:45-46.

FEMALE (fig. 8). Vertex with all decumbent scales narrow curved; occiput with

all or most erect scales pale; complete acrostichal line absent; fossal macula well

developed, not interrupted laterally by dark scales; pra hairs dark; vein C with

silver scales at base.

MALE (fig. 8). Vertex with decumbent scales as in female; mesonotal disc trans-

versely silvered anteriorly for at least 0.5 length of fossa; fossal macula reaching

mesal margin of fossa; pra hairs dark.

MALE GENITALIA (fig. 8). Sidepiece length 0.32-0.35 mm (0.27-0.42 mm);

prosophallus width 0.14-0.15 mm (0.13-0.15 mm).

PUPA. Cephalothorax without highly contrasting dark markings on wing and

leg cases; hair 1-I with primary branches predominantly multiple; paddle rounded

apically.

LARVA (fig. 9). Head hair 5-C usually double (1-3); hair 6-C single; 11-C much

shorter than 7-C; L/S 2.1-2.4.

SYSTEMATICS. The above description is essentially condensed from Schick

(1970a) and the definition of the species remains unchanged.

Aedes zavortinki is very similar morphologically to some forms of terrens and

generally can be separated from that species only by the mesonotal markings of

the male (see terrens). The separation of zavortinki from braziliensis and the Ter-

esopolis form is discussed under those species.

Aedes zavortinki, previously known only from the Canal Zone and eastern Pan-

ama at low elevations, 90 m or less (fig. 3), is now recorded from Colombia at

an elevation of about 460 m.

NEW RECORDS (fig. 61). Material examined: 2 M.

COLOMBIA. Meta: Villavicencio [elev. ca 460 m], 17 May 1939, 1 M; 2 June 1941, very

low treehole, M. Bates, 1 M (277) [USNM].

Schick: Additions to Aedes terrens Group 19

7. Aedes (Finlaya) sp., Teresopolis form

Figs. 61,65,66

Aedes (Finlaya) terrens in part of Schick (1970a:41-43).

FEMALE (fig. 65). Vertex with all decumbent scales narrow curved; occiput

with erect scales pale; complete acrostichal line present; fossal macula with a nar-

row sublateral longitudinal band transversely divided into anterior and posterior

portions; pra hairs pale; vein C without silver scales at base.

MALE (fig. 65). Vertex with all decumbent scales narrow curved; complete ac-

rostichal line present; mesonotal disc not transversely silvered; fossal macula not

reaching mesal margin of fossa; pleural hairs missing.

MALE GENITALIA (fig. 65). Sidepiece length 0.35 mm; median sternomesal

sclerite of sidepiece broad and well developed; prosophallus width 0.14-0.16 mm.

PUPA (fig. 65). Cephalothorax without highly contrasting markings on wing and

leg cases; hair 1-I with primary branches predominantly single or double; paddle

broadly rounded apically.

LARVA (fig. 66). Hair 5-C triple or 4-branched; 6-C with 4-6 branches; 11-C

subequal in length to 7-C; L/S 1.7-2.1.

SYSTEMATICS. The Teresopolis form, apparently most closely related to ter-

rens, differs from that species in the following characters: (1) complete acrostichal

line present in both sexes, (2) vein C of female without silver scales at base, (3)

the median sternomesal sclerite of the sidepiece more strongly developed, (4) hair

l-I of the pupa with fewer secondary branches, (5) hair 6-C of the larva more

highly branched, (6) hair 11-C of the larva much longer (longer than in any of

the other species of the group), and (7) L/S tending to be lower (1.7-2.1 vs 2.0-

2.4). | .

The male from Teresopolis is unusual in that the cercal setae are absent. These

are present on the P 48-1 male (see distribution).

The Teresopolis form is not provided with a formal name at this time because

of insufficient adult material and lack of collections from adjacent localities.

DISTRIBUTION (fig. 61). Brazil. Material examined: 9 specimens; 3 M, | F,

2 pupae, 3 larvae; 2 individual rearings (1 larval, 1 pupal).

BRAZIL. Rio de Janeiro: Fazenda Boa Fe, Teresopolis [elev. 500-1000 m], 13 Nov 1942,

treehole, L. Gomes, 2 L (28773-3,4); same data, but habitat not specified, 1 lpF (28774-12);

12 Dec 1942, 1 pM (29078-1) [UCLA]. Locality not specified: 1 M (P 48-1); 1 M (P 100(1);

lam. P 82/P 81 [genitalia slide missing) [UCLA].

Alboapicus Subgroup

9a. Aedes (Finlaya) sp., Chaco form

Figs. 62,63

FEMALE (fig. 63). Vertex with an area of moderately broad, dark decumbent

scales adjacent to narrow curved scales of median longitudinal line; erect occipital

scales dark; proboscis subequal in length to femur I; acrostichal setae absent; fos-

sal macula reduced to small posterior spot about 0.5 length of fossa; supraalar

20 Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

macula small, very narrowly joined to fossal macula; ssp scales present; pra hairs

dark; femora I and II with well-developed posterior patch of silver scales; femur

I with small knee spot; tarsi 5-I-III silver; femur II with moderately broad knee

spot, a few of the silver scales extending basad of subapical setae; tarsus 1-II with

median dark band 0.75; femur III with broad basal dark band and very broad

subapical dark band (about 0.42); veins C and R with silver lines subequal in length,

neither reaching level of crossvein h.

MALE, PUPA, LARVA. Unknown.

SYSTEMATICS. The Chaco form is provisionally placed in the Alboapicus Subs

group because of the silvered 5th tarsal segments and the broad basal band of fe-

mur III. A less tenuous placement must await the discovery of the other stages.

The Chaco form differs from alboapicus in the completely silvered tarsus 1-I,

the presence of a knee spot on femur I, the greater reduction of the fossal macula,

the presence of well-developed posterior silver patches on femora I and II, the

very broad subapical dark band of femur III and the long silver line on vein R.

The presence of a knee spot on femur I and a silver line on vein R in the female

are unusual characters for the Terrens Group. The former character occurs else-

where only in amabilis and the latter only in argyrothorax.

DISTRIBUTION (fig. 62). Northern Argentina. Material examined: 1 F.

ARGENTINA. Chaco: Saenz Pena (rd to Tres Isletas) [elev. ca 100 m], 6 Apr 1963, in forest,

biting in afternoon, M. Castro, 1 F (Ch 61b) [UCLA].

Insolitus Subgroup

Three species may now be recognized in this subgroup, insolitus, berlini (trans-

ferred from the Terrens Subgroup) and a new species, casali.

The description of the subgroup must be broadened to accommodate the latter

2 species. The additions to the description are indicated in italics in the following

statements: (1) proboscis of female usually longer than femur I, (2) femur II with

knee spot usually present, narrow to broad, (3) femur III with basal dark band ab-

sent, incomplete or complete, usually broad, and (4) sidepiece with median sterno-

mesal sclerite weakly to strongly developed.

Further subgroup characters that are of value in separating this from the Ter-

rens Subgroup are given under the latter.

Aedes casali occurs in northern Argentina, the southern limit of the geographic

range of the Terrens Group, and is broadly disjunct from the other 2 species of

the subgroup. The occurrence of such a relict species at the periphery of the range

of the group supports the hypothesis that the Insolitus Subgroup is of relatively

great antiquity (Schick, 1970a:20).

12. Aedes (Finlaya) insolitus (Coquillett)

Figs. 4,29,30,62

1909. ‘Verrallina insolita Coquillett, 1906:62.

Aedes (Finlaya) insolitus of Schick (1970a:57-61).

SYSTEMATICS. Aedes insolitus was reported from Central America, Colombia

Schick: Additions to Aedes terrens Group 21

and Trinidad, primarily at high elevations, in my previous paper (fig. 4). New rec-

ords show that this species also occurs in northern Venezuela.

NEW RECORDS (fig. 62). Material examined: 82 specimens; 16 M, 21 F, 25

pupae, 20 larvae; 24 individual rearings (14 larval, 9 pupal, | incomplete).

COLOMBIA. Meta: Villavicencio [elev. 460 m], 19 June 1941, treehole, M. Bates, 1 F (278)

[USNM].

VENEZUELA. Aragua: Carretera 2, 8.7 km S Choroni, elev. 300 m, 16 July 1969, cut bam-

boo, T. Zavortink et al, 1 M (VZ 228-2); 15.9 km S Choroni, elev. 650 m, 16 July 1969, cut

bamboo, T. Zavortink et al (VZ 230), 1 pF (230-100), 1 M, 4 F (230-1); Cumbre de Choroni,

elev. 900 m, 26 July 1969, cut bamboo, J. Pulido and J. Valencia (VZ 269), 1 IpM (269-50),

1 IpM (269-50), 1 lp (269-51), 1 M, 1 F, 21(269-5); 20 km N Maracay, elev. 800 m, 6 Aug 1969,

cut bamboo, J. Valencia (VZ 314,315), 1 lpM (314-70), 1 pM (315-104); 4 km N Maracay, elev.

800 m, 6 Aug 1969, small treehole, height 1 m, J. Valencia (VZ 317), 1 IpM (317-10), 1M, 1 F, 1

1 (317-1) [UCLA]. Guamita (8 km S Rancho Grande), elev. 700-800 m, 15 July 1969, cut bam-

boo, T. and J. Zavortink (VZ 203), 1 IpM (203-54), 2 lpF (203-56,58), 1 lp (203-61); 11 Aug

1969, small treehole, height 1 m, J. Valencia (VZ 328), 1 IpF (328-12), 1 pM (328-105); 18

Aug 1969, cut bamboo, J. Pulido and J. Clavijo (VZ 371), 1 lpF (371-10), 2 pF (371-101,103);

18 Aug 1969, small treeholes, height 1-2 m, J. Pulido and J. Clavijo (VZ 372,373,375), 2 lpM

(373-11,12), 3 lpF (373-13-15), 1 pM (372-100), 3 pF (873-100,101; 375-101), 1 IP (373-16),

1 1 (375-1) [UCLA]. Turiamo [elev. near sea level], 11 Sept 1944, 1 M; date unknown, 2 M

[UCLA].

12a. Aedes (Finlaya) casali Schick, n.sp.

Figs. 62,67,68

TYPES: Holotype male (ARG 618-108) with associated pupal skin, ca 7 km from Tablillas

(nearest town General Enrique Mosconi (Vespucio), elev. ca 500 m), 25 Feb 1967, fallen tree,

O. Casal and M. Garcia [USNM]. Allotype female (618-106) with associated pupal skin, same

data as holotype [USNM]. Paratypes: 1 pM (618-103), 1 M (618-113), same data as holotype

[UCLA]. This species is dedicated to Osvaldo Casal of the Instituto Nacional de Microbiologia,

Buenos Aires, Argentina.

FEMALE (fig. 67). Vertex with broad decumbent scales usually forming var-

iously developed dark patch surrounded by silver scales; infrequently all scales sil-

ver; anterior promontory silvered; acrostichal line absent; acrostichal setae present,

most posterior seta at about 0.5 from anterior end; fossal macula moderately well

developed (ARG 773-109) or reduced to small anterior patch (IMR 29); ssp scales

present; pra hairs pale; femur II with knee spot narrow, often longitudinally di-

vided anteriorly by streak of dark scales; tarsus 1-I] with complete median dark

band, usually about 0.4-0.6 (0.4-0.7); tarsus 2-II with complete dark apical band;

femur III usually without basal dark band, incomplete when present.

MALE (fig. 67). Mesonotal disc transversely silvered anteriorly for about 0.5

length or up to prescutellar space; ssp scales present; tarsus 1-II with incomplete

median dark band or band complete, as broad as 0.6; femur III with basal dark

band as in female.

MALE GENITALIA (fig. 67). Sidepiece length 0.32-0.35 mm; median sterno-

mesal sclerite weakly to strongly developed; prosophallus length 0.09-0.10 mm;

filament ratio 0.55-0.80.

PUPA (fig. 67). Cephalothorax without pale inverted V-shaped marking; hair

1-I with primary branches predominantly single-double or double-triple; 2-II lat-

ze Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

erad of 3-II or mesad for as much as 0.4 the distance from 1-II to 3-Il.

LARVA (fig. 68). Hairs 5,6-C single or double; 14-C, bmh usually double and

branching from base (1-3); hair 11-P longer than 0.5 of 14-P; hair 14-P single;

4-M double; 7-II single to triple; 12-VII single; apical comb scale with free por-

tion 0.035-0.040 mm; 2-VIII single; siphon length 0.67-0.92 mm; L/S 2.1-2.3.

SYSTEMATICS. Aedes casali is distinguished from the other species of the sub-

group by the absence of a complete band at the base of femur III and by the col-

oration of the broad scales of the vertex of the female. The pupa is similar to that

of the other species. The larva apparently shows no features that separate it from

insolitus but can be distinguished from berlini by fewer branches in many of the

hairs and by the smaller L/S.

DISTRIBUTION (fig. 62). Northwest Argentina at elevations of about 500 m.

Material examined: 52 specimens; 9 M, 22 F, 11 pupae, 10 larvae; 6 individual

pupal rearings. |

ARGENTINA. Salta: Tablillas, ca 7 km from (nearest town General Enrique Mosconi (Ves-

pucio), elev. ca 500 m), 25 Feb 1967, fallen tree, O. Casal and M. Garcia (ARG 618, type ser-

ies), 2 pM (618-103,108), 1 pF (618-106), 1 M (618-113), 8 L (618-1) [USNM, UCLA]; 6 June

_ 1969, treehole, O. Casal and M. Garcia, 3 pF (ARG 773-109,113,116) [UCLA] . Tucuman: Chur-

qui, 4 Dec 1922 (IMR 29), 1 pF (slide 2217), 8 M, 17 F, 1 M gen (360), 4 p (2216), 2 L (2213)

[UCLA].

(8) 12b. Aedes (Finlaya) berlini Schick

Figs. 3,21,22,62,63

1970. Aedes (Finlaya) berlini Schick, 1970a:48-49.

FEMALE (fig. 21). Vertex with decumbent scales usually all dark, some scat-

tered silver scales sometimes present; anterior promontory silvered or dark; acro-

stichal line absent; acrostichal setae present or absent, most posterior seta at about

0.25 from anterior end or as far caudad as in posterior 0.5; fossal macula relative-

ly well developed; ssp scales absent; pra hairs dark; femur II with broad knee

spot, the silver scales extending well basad of anterior subapical setae; tarsus 1-II

with complete median dark band, about 0.4-0.5; tarsus 2-II with complete dark

apical band; femur III with complete and broad basal dark band.

MALE (fig. 63). Mesonotal disc transversely silvered anteriorly for about 0.5

length or up to prescutellar space; ssp scales absent; tarsus 1-II with incomplete

or complete median dark band, as broad as 0.5; femur III with basal dark band

as in female. |

MALE GENITALIA (fig. 63). Sidepiece length 0.29-0.31 mm; median sterno-

mesal sclerite weakly developed; prosophallus length 0.08 mm; filament ratio 0.90-

1.10.

PUPA (fig. 21). Cephalothorax with or without pale inverted V-shaped mark-

ing; hair 1-I with primary branches usually predominantly double or triple, often

multiple, sometimes single; 2-II mesad of 3-II for 0.2-0.5 the distance from 1-II

to 3-II.

LARVA (fig. 22). Hairs 5,6-C single to triple; 14-C, bmh single or double, when

double not branching from base; 11-P greater or less than 0.5 of 14-P; hair 14-P

single or double; 4-M usually triple or 4-branched (2-4); hair 7-II with 4-7 branches;

12-VII single or double; apical comb scale with free portion 0.030-0.035 mm; 2-

Schick: Additions to Aedes terrens Group 23

VIII single or double; siphon length 0.80-0.92 mm; L/S 2.3-2.6.

SYSTEMATICS. Aedes berlini, previously known from only the type locality,

the island of Tobago (fig. 3), is now recorded from Trinidad, Venezuela and Co-

lombia. The populations fall into 2 types, Island (Trinidad, Tobago) and Main-

land (Venezuela, Colombia). The former populations occur at elevations of up to

120 m and the latter from 700 to 900 m. They differ in the following 3 morph-

ological characters (Island character state cited first): (1) acrostichal setae absent

or when present not extending caudad of anterior 0.25 vs acrostichal setae always

present, most posterior seta at 0.5 or in caudal 0.5, (2) hair 11-P of the larva

less than 0.5 of 14-P vs subequal in length to 0.5, and (3) mesonotum of male

transversely silvered up to prescutellar space vs to about 0.5-0.67 of length.

This broad treatment of berlini is consistent with that of insolitus. The latter

species also occurs at a wide range of elevations and shows the same or a similar

type of geographic variation in the above 3 characters.

Aedes berlini differs from the other species of the subgroup in the absence of

ssp scales, the dark pra hairs of the female, and the single or nonbasally branched

hairs 14-C and bmh of the larva. The fossal macula of the female, relatively well

developed, is more constant in its degree of development than in the other species.

Emphasis was placed on the taxonomic value of the length of hair 7-VII of

the pupa in Schick (1970a:49). Although long in all the new material examined,

this character is variable enough to negate its taxonomic value at least as a species

criterion in the Insolitus Subgroup.

Aedes berlini and insolitus are sympatric in Venezuela and Colombia and have

been taken in the same treehole in the former country.

NEW RECORDS (fig. 62). Material examined: 114 specimens; 21 M, 19 F, 40

pupae, 34 larvae; 38 individual rearings (23 larval, 11 pupal, 4 incomplete).

COLOMBIA. Cundinamarca: Santander (Santandercito), 1 F (207D-6/2), 1 F (207D-8/3) |

[UCLA]. Meta: Forzosa Forest (ca 5 mi from Villavicencio, elev. ca 460 m), 12 June 1944,

treehole 6, M. Bates, 1 IM [USNM]. Villavicencio, 1944, M. Bates, 1 lpF (78) [USNM].

TRINIDAD. Saint George: Grandwood (ca 2 km Chaguaramas), elev. ca 120 m, 11 June

1961, treehole, T.H.G. Aitken, 2 IpM (11-VI-61-8,13), 3 lpF (11-VI-61-11,12,14), 1 pM (11-

VI-61-4), 7 Ip (11-VI-61-5,7,9,10,15,16,18), 3 p (11-VI-61) [UCLA].

VENEZUELA. Aragua: Guamita (8 km S Rancho Grande), elev. 700-800 m, 15 July 1969,

small treehole at ground level, T. and J. Zavortink (VZ 198), 1 IpM (198-10), 4 L (198-2)

[UCLA]; small treehole, height 1 m (VZ 200,201), 3 IpM (200-10,13; 201-11), 2 lpF (200-

11,12), 1 pM (200-100), 1 M, 1 1 (200-1) [USNM, UCLA]; cut bamboo, height ca 1-2 m (VZ

203), 1 IpM (203-50), 4 IpF (203-51,52,60,64), 3 pM (203-100,103,110), 1 lp (203-53), 1

M, 2 L (203-6); 11 Aug 1969, small treehole, height 1 m, J. Valencia (VZ 328), 1 IpF (328-10),

1 pM (328-103), 1 pF (328-101); cut bamboo, height ca 1 m, J. Valencia and J. Clavijo (VZ

331), 1 IpF (331-11), 1 M (331-1); 18 Aug 1969, cut bamboo, height ca 1 m, J. Pulido and J.

Clavijo, 1 pM (VZ 371-100); small treehole, height 1 m, 2 lpM (VZ 372-10,40); small treehole,

height ca 2 m (VZ 374), 1 pF (374-100), 1 F (374-1) [UCLA]. Maracay, 4 km N on rd to

Choroni, elev. 800 m, 6 Aug 1969, small treehole, height ca 1 m, J. Valencia, 1 pM (VZ 317-

100) [UCLA]. Maracay, 20 km N on rd to Choroni, elev. 800 m, 6 Aug 1969, cut bamboo,

height near soil level, J. Valencia, 1 lpF (VZ 314-71) [UCLA].

Podographicus Subgroup

25. Aedes (Finlaya) podographicus Dyar & Knab

Figs. 6,52-57

24 Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

1906. Aedes podographicus Dyar and Knab, 1906:165.

Aedes (Finlaya) podographicus of Schick (1970a:82-86).

SYSTEMATICS. The podographicus of Schick (1970a) comprised coastal pop-

ulations in Central America and Mexico which occurred at elevations of less than

300 m. A widely disjunct but apparently conspecific population, cited simply as

belonging to the podographicus complex, was known from the Maracay area of

Venezuela at higher elevations but was not treated as podographicus proper since

the immature stages were not known. These are now available and show no strik-

ing differences from those of typical populations of Central America and Mexico.

Although podographicus abounds in the coastal areas to the north, it is appar-

ently absent along the coast of South America. In recent UCLA collections in

the state of Aragua, Venezuela, podographicus was taken only in the Maracay area,

although another member of the Terrens Group (terrens) was commonly encoun-

tered on the coast.

NEW RECORDS. Material examined: 292 specimens; 81 M, 142 F, 37 pupae,

32 larvae; 33 individual rearings (24 larval, 9 pupal).

VENEZUELA. Aragua: Guayabita, elev. 560 m, 14 July 1927, 1M (35. III.20a) [USNM]; 30

Aug 1966, automobile tire, E. Russian Vasquez, 1 F (VZ 29) [UCLA]. Hacienda Santa Clara

(nearest town San J oaquin), elev. 400 m, 19 July 1969, large treehole near soil level, J. Pulido

and J. Valencia (VZ 249), 1 IpM (249-21), 1 IpF (249-20) [UCLA]. Macaro (nearest town

Turmero), elev. 500 m, 12 Aug 1969, small treehole, height 2 m, J. Valencia and J. Pulido, 1

IpM (VZ 335). Maracay [elev. 600 m], 10 Jan 1926 (1-10-26), M. Nunez Tovar, 7 F; 24 Aug

1926, M. Nunez Tovar, 1 F; 8 Sept 1926, M. Nunez Tovar, 6 F; 11 Sept 1926 (9-11-26), M.

Nunez Tovar, 1 F; 13 Sept 1926, M. Nunez Tovar, 4 F; 18 Sept 1926, M. Nunez Tovar, 1 F;

1926, M. Nunez Tovar, 5 M, 1 F; 3 Aug 1927, M. Nunez Tovar, 1 M, 1 F [USNM]; 30 M,

12 F (VZR 252); 23 M, 86 F (VZR 257) [UCLA]. Maracay, Hacienda Militar de San Jacinto,

elev. 550-600 m, 17 July 1969, small treehole, height 1 m, J. Pulido and J. Valencia (VZ 233,

234,239,240), 4 lpM (233-12; 234-10; 239-20,21), 3 lpF (233-11; 234-12,13), 3 pM (233-101;

234-100,101), 1 pF (233-100), 1 lp (234-16), 1 1 (233-1), 6 M, 3 F, 3 p, 3 1 (234-1), 2 F

(240-3) [UCLA]. Maracay, Universidad Facultad de Agronomia, elev. 600 m, 15 July 1969,

small treehole, height 1 m, J. Pulido and J. Valencia (VZ 205,206), 3 lpM (205-31; 206-10,11),

7 IpF (205-10,12-16,30), 2 pM (205-100,101), 3 pF (205-102,103; 206-100), 1 lp (205-17), 1 L,

2 1 (205-1), 1 F, 1 p (206-1) [UCLA]. Carabobo: Mariara, 2 km E, elev. 400 m, 19 July 1969,

small treehole, height 1 m, J. Pulido and J. Valencia, 1 IlpM (VZ 245-11) [UCLA]. Punta Pal-

mita (nearest town Mariara), elev. 400 m, 19 July 1969, small treehole, height 1 m, J. Pulido

and J. Valencia (VZ 244), 1 lpM (244-12), 1 lp (244-11), 1 1 (244-1) [UCLA].

KEYS TO SPECIES

FEMALES

(15. impostor unknown)

L. Femur III with complete basal dark band and supraalar macula broadly

reaching scutal suture (figs. 63, braziliensis;65,67)... . . nee

Without the above combination of characters; femur III sometimes with ;

complete basal dark band and supraalar macula sometimes broadly

reaching scutal suture but both conditions not developed simultane-

BUSIV eh a ee a a eed

a1).

3(2).

4(3).

5(4).

6(4).

7(6).

8(6).

9(8).

10(8).

11(10).

12(11).

Schick: Additions to Aedes terrens Group 25

Midtarsi and hindtarsi unmarked except for inconspicuous silver band at

base of tarsi 1-IT,IT]. . . . . . .10. buenaventura

Midtarsi and hindtarsi with prominent silver bands at base and apex of

tarsi (c1L-TUl and-at baseof tats 2-112 ees eee. 8

Tarsi 5-II,III silvered . . . . 9. alboapicus, in part

Tarsus 5-1 usually dark, rarely silverd: 5-Il] always dark . Lo ae

Vertex with decumbent scales along longitudinal midline all broad. . .5

Vertex with decumbent scales along longitudinal midline all narrow curved

. 6

Mesonotal disc transversely silvered: anteriorly Se 9); vein R without

silver scales. 6° . . 1. thorntoni

Mesonotal disc not transversely silvered (fig. 13); vein rR with small basal

patch of.silver scales’ ¢ is a eee ee are yrothorax

Ss states dsentis) 45 Phe oe Oe ee ee

Biponies present... ee ee ae ee

Vertex with all decumbent scales narrow curved, most or all along longi-

tudinal midline dark. . . . . . . 11. metoecopus

Vertex with an area of broad deounbent scales adjacent to narrow curved

scales of median longitudinal line, latter scales silver . (8) 12b. berlini

Vertex with all decumbent scales narrow curved. . . og

Vertex with an area of broad decumbent scales adj scent to narrow earved

acalés Of-inedian longitudinal line 2 arse". Se Re Or ete a TO

Pra hairs dark; fossal macula not strongly reduced, mesal margin sharply

defined, evenly and gently curved or essentially straight (fig. 63)

5. braziliensis

Pra hairs pale: fossal macula usually markedly reduced and mesal margin

indistinct and irregular, infrequently well developed as above or com-

pletely absent (fie. 29) ane 6 eRe eZ, ipsolitus

Basal dark band of femur III narrow, at most about 0.05 (Mexico) .

. . 14. homoeopus, in part

Basal. cee band or fom Tl broad: more than 0.10 (Panama, South

AMeTiCg) do eo poate ese as ee ee Oe ee a

Complete acrostichal line present ... .. . .. 7. Teresopolis form

Complete acrostichal dinéabsent. cas) ip Ci ee a ee ea, a

Fossa without dark scales at lateral margin (fig. 17) (Panama, Colombia) .

6. zavortinki

Fossa usually with at least a : few: dark scales at lateral margin, these often

forming well developed patch or — band sil 15) (Colombia

southward into Argentina) . ..... : . . . 4, terrens

13(1).

14(13).

15(13).

16(15).

17(16).

18(15).

19(18).

20(19).

21(20).

22(19).

23(22).

Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

Mesonotal disc transversely silvered anteriorly. ......... 14

Mesonotal disc not transversely silvered . . .......... 15

Transverse silvered area of mesonotum espera: caudad to about 0.5

(fig. 11);ppnsilverscaled. . ... . . . . 3. bertrami

Transverse silvered area of mesonotum much - narrower, extending caudad

to less than 0.25 (fig. 39); ppn dark scaled . . . . . . . 13. aitkeni

Taras Oi silvers face Sa eats Penge ee.

Tapes OT ars ee Pes Pee ea

Tarsus 1-I with broad apical silver band, about 0.4; midlegs and hindlegs

shaggy . . pee i a aaa

Tarsus 1-I with at most very - narrow ‘apical silver band, much less than

0.4; midlegs and hindlegs not shaggy .... . a ie lara a 17

Fossal macula developed up to anterior margin of fossa (fig. 23); femur I

without knee spot; tarsus 5-I at most only partly silvered

. . 9, alboapicus, in part

Fossal macula a small posterior spot (fig. 63): femur I with small knee

spot; tarsus 5-I entirely silvered . . ... . . . . 9a.Chaco form

Supraalar macula broadly reaching scutal suture (fig. 67) and mesal margin

of fossal macula poorly defined (Argentina). . . . . . . 12a. casali

Supraalar macula rarely broadly reaching scutal suture; mesal margin of

fossal macula usually sharply defined (Mexico, Central America) . 19

Acrostichal setae absent; tarsus 1-I] with median dark band usually in-

complete or complete and narrow, at most about 0.33... . . 20

Acrostichal setae present; tarsus 1-II with median dark band usually com-

piete and Dradd, about O53 of greater: Go ee ee a

Proboscis shorter than or subequal in length to femur I; femur II with

knee spot moderately broad, the silver scales at most just reaching an-

terior subapical setae . . . . . 25. podographicus

Proboscis longer than femur I. femur 1. with knee spot broad, the silver

scales extending basad of anterior SUDADICR Seige 2 eEOS ACE

Vertex with all decumbent scales silver .. . . . . 26, tehuantepec

Vertex with silver and dark decumbent scales, latter forming submedian

Rg eG >, wer ea a ake, uaa ky <5 aid a: See

Femur II with or without knee spot, when present, narrow, a single row

of apical scales; ssp scales absent. . . ; ae

Femur II with broad knee spot, the scales extending basad of anterior sub-

apical setac. Ssp scales Drese@l OF a0Senl 4... 4 ee OE

Fossal macula reduced only mesally (figs. 48,50); supraalar macula reach-

ing scutal suture; femora I,II with well developed posterior patch of sil-

Wr SCAN, ps inc. A oe ae ee Ge oS WL ee OO

24(22).

25(24).

26(25).

27(25).

28(27).

29(28).

Schick: Additions to Aedes terrens Group 27

Fossal macula reduced anteriorly and mesally (fig. 44); supraalar macula

not reaching scutal suture; femora I,I] without posterior patch of sil-

Wer segles i. end dew. Gad Ee eee kd. Salindaly 23. campana

Ssp scale patch absent although | or 2 scales sometimes present .

reid tne ineties tO. vargasi

Ssp scale patch present: Cheek Piri) aber: pane gi REPL en dl tie, pe

Proboscis shorter than or subequal in length to femur I; midlobe of scu-

tellum with silver scales ... Bes ie

Proboscis usually longer than, sometimes subequal it in length to femur I,

when subequal midlobe of scutellum without silver scales . . . . 27

Acrostichal line absent or represented by scattered silver scales (fig. 39);

midlobe of scutellum usually with a mixture of silver and dark scales,

the dark scales usually predominating, infrequently all scales silver .

19. sumidero

Actostichal line present, ‘complete (fig. 35); ‘midlobe of scutellum with

allscales silver 24. x pete ye SEE obo dledg Sincere LA ogabeel

Femur I with narrow knee spot; complete and strong acrostichal and pos-

terior dorsocentral lines present (fig.37). . . . . . . 16. amabilis

Femur I without knee spot; complete and strong acrostichal and posterior

dorsocentral lines usually absent, sometimes complete but weak. . 28

Vertex with an area of broad decumbent scales — to narrow curved

scales of median longitudinal line. . . . . . . . . 18. idanus

Vertex with all decumbent scales narrow curved. a/b Tati ien govencee:

Midlobe of scutellum without silver scales; proboscis subequal in length

to orlongerthanfemurl. . . . . . . 12. heteropus

Midlobe of scutellum usually with silver ‘scales; proboscis longer than

femurt... . kA wy 4k we eS oe pe ocarns, inet

MALES

(3. bertrami, 9a. Chaco form, 13. aitkeni, 16. amabilis,

27. schroederi and 28. diazi unknown)

Vein C with basal line of silver scales reaching crossvein h; mesonotal disc

usually transversely silvered. . . ae

Vein C with small basal patch of silver scales or - line ‘reaching at most to

about 0.5 to crossvein h; mesonotal disc not transversely silvered . 14

Vein R with basal line of silver scales much longer than that of veinC. .3

Vein R with basal line of silver scales much shorter than that of veinC .4

Median sternomesal area of sidepiece with well developed tuft and usually

with well developed convexity and sclerite (fig. 31). . 14. homoeopus

Median sternomesal area with these structures not strongly differentiated

41 BES) Geir Mee Gee eae ety Ma ol ERM EME

4(2).

5(4).

6(4).

7(6).

8(6).

9(8).

10(9).

11(8).

12¢11).

13(11).

14(1).

15(14).

Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

Ssp scale patch absent . Pe ee

Dep stale patch Present a pe te oc ee XO

Midtarsi and hindtarsi unmarked except for inconspicuous silver band at

base of tars: TALI oe 2 | . . . . .10. buenaventura

Midtarsi and hindtarsi with prominent silver bands at base and apex of

tarsi 1-II,IJI and at base of tarsi 2-II,[II; tarsi 5-II,III and sometimes

Sl eiveted ge a ee 8, albeapicus

Vertex with decumbent scales along longitudinal midline all broad. . .7

Vertex with decumbent scales along longitudinal midline all narrow curved

Transverse silvered area of mesonotal disc not emarginate posteriorly (fig.

9); palpus 1 or 2 labellum lengths shorter than proboscis. Claspette

filament not expanded distally (figs.9,11) . . . . . . 1. thorntoni

Transverse silvered area of mesonotal disc emarginate posteriorly (fig. 13);

palpus 4-7 labellum lengths shorter than proboscis. Claspette filament

expanded distally (fig. 13)... s- : 6s es OP 2 areyrothorax

Vertex with an area of broad, silver decumbent scales adjacent to narrow

curved scales of median longitudinal line; mesonotal disc usually trans-

versely silvered anteriorly, the silvered area usually reaching caudad of

fossa (fig. 29) and not emarginate caudally... . a9

Vertex with or without this lateral area of broad decumbent scales, these

dark when present; mesonotal disc transversely silvered or not, the sil-

vered area not reaching caudad of fossa and emarginate caudally (fig.

Pye ee a a Oo ee Ee ie

Femur III without basal dark band or with incomplete band . 12a. casali

Femur dil withecomplete basal: dark Band). G22 0 sw. TO

Dep scales present = yt Pee ee ee a 2, Insolitus

Ssp scales absent 2 Os op ee ore igs. (8) 1 2b; beskian

Mesonotal disc not transversely silvered © 2 so 4 we. Se eh

Mesonotal disc transversely silvered anteriorly. . ........ 13

Complete acrostichal line present (fig.65) . . . . .7.Teresopolis form

Complete acrostichal line absent (fig. 15, Rio de Janeiro) . . .4. terrens

Vertex with an area of broad decumbent scales en to narrow curved

scales of median longitudinal line. . . . . . . . 5. braziliensis

Vertex with all decumbent scales narrow caaed . . . . 6. zavortinki

MCrORte tal-se(ae aseht 4 oko SS ak eee ee ed Sw DD

PCIOSICUA Seta PCtet a ee a a OR

Occiput with erect scales dark. . ..... . . . . 11. meteocopus

Occiput with erectecalen-pale: ont cose sice luedeeerwe ts cere. ar 16

16(15).

17(14).

18(17).

19(18).

20(17).

21(20).

22(20).

pol 22),

Schick: Additions to Aedes terrens Group 29

Palpal segment 3 without prominent tuft of setae, the setae at ventro-

lateral apex shorter than segments 4 and 5 combined. Prosophallus

with lateral portion of mesal lobe usually moderately inclined, between

15° and 30° from horizontal; stems usually bowed and convergent

(fins: 52.54, 56)u0%. cuits . . . . . 25, podographicus

Palpal segment 3 with prominent tuft. of setae as long as segments 4 and

5 combined. Prosophallus with lateral portion of mesal lobe slightly

inclined, about 15° or less from horizontal; stems not bowed, diver-

gent or essentially parallel (fig. 58) . . .. . . . . 26. tehuantepec

Femur II without or with narrow knee spot, the silver scales a single row

at apex of segment; ssp scale patch absent . . . Hie oie, le

Femur II with broad knee spot, the silver scales extending basad of an-

terior subapical setae; ssp scale patch present . ........ . 20

Femora I,II with well developed posterior patch of silver scales. Sidepiece

with median sternomesal tuft poorly differentiated and the setae not

wavy (figs.48,50) .... yoann 6 ee. Corel

Femora I,II without posterior patch of silver ‘scales. Sidepiece with med-

ian sternomesal tuft well differentiated and the setae wavy (figs. 44,46)

Prosophallus with median lobe projecting farther cephalad than lateral

lobe (hie, 44) 2 se . . . .22, galindoi

Prosophallus with median lobe projecting to about same level as lateral

1008 (1G AO) ek ok Kh ie he he te 1 OR a ea ee eae

Median sternomesal area of sidepiece with sclerite and tuft well developed;

hook of filament strongly angulate (figs. 35,37) . . ...... 2!

Median sternomesal area of sidepiece with sclerite and tuft absent or

poorly developed; hook of filament not ig Suda hee 39,40,

Oh rN tae SEEM RR et ot he Sele

Palpus subequal in length to or slightly longer than proboscis. Basal tergo-

‘mesal area of sidepiece without dense patch of long setae (fig. 35) .

17. gabriel

Palpus about 7 labellum lengths shorter than ‘proboscis. Basal tergomesal

area of sidepiece with dense patch of long setae (fig. 37). . 18. idanus

Ssp scale patch absent; complete acrostichal or posterior dorsocentral

lines absent (fig. 40); tarsus 1-II with median dark band incomplete

ee . 20. vargasi

Ssp scale patch present; “complete acrostichal « or posterior dorsocentral

lines present; tarsus 1-II with median dark band complete, about 0.33-

Acrostichal line absent; posterior dorsocentral line complete (fig. 39);

segment 3 of palpus with apical ventrolateral tuft not as long as seg-

ments4and5 combined. .. . . . 19. sumidero

Acrostichal line present, complete, sometimes weakly developed: posterior

2(1).

3(2).

4(3).

5(4).

6(5).

7(S).

8(7).

92).

10(9).

Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

dorsocentral line incomplete (fig. 42); segment 3 of palpus with apical

ventrolateral tuft as long as segments 4 and 5 combined . 21. heteropus

LARVAE

(3. bertrami, 9a. Chaco form, 16. amabilis

and 19. sumidero unknown)

Hair 11-C subequalin length to7-C. . .. . . . .7.Teresopolis form

Plate ti tec chor tet (a ee ee ae

Hair 5-C usually with 4 or more branches (when fewer branches, only on 1

side) and/or 14-P branched . . . ee

Hair 5-C usually single or double, sometimes triple (when more ‘branches,

only on | side) and 14-P usually single (when rarely branched, only on

Fe ee ee ee eee ot a aca ian oe eee

Hair 5-VII cephalad of 4-VIJ_ . ...... . . . .10. buenaventura

Death Oe Ue NT en a a a le ee ee

Hairs 4-VII and 3-VI branched. . . . . . 9, alboapicus

Hair 4-VII single and 3-VI ie one single (when rarely branched, nae on

emi ge ee Cae

Hair 2-II well mesad of 4-Il (fig. 42); hair 14-P single . . oO

Hair 2-II mesad of 4-II for about only 1 alveolus width, often laterad of

li (tig 44) nan tee branched... Re Sk cee oe eee

Hair 11-P less than half length of 14-P; hair 1-VIII shorter than 2-VIII;

hair 6-C single or double; bmh single . . . . 18. idanus

Hair 11-P about half length of 14-P; hair 1-VIII usually at least subequal

in length to 2-VIII; hair 6-C usually with more than 2 branches; bmh

usually branched but often single ....... . . 21. heteropus

Hair 14-C usually with 3 or more branches; 1-A usually branched but of-

ten single; free portion of apical scales ligulate, awl shaped or spatulate

. . 24, daryi

Hair 14-C ‘usually with fewer ‘than 3 ‘branches; eA usually single (when

branched, only on | side); free portion of apical scales spatulate. . .8

Comb scales 35-47, in 4 rows; free portion of midapical scale longer than |

sessile portion. . . . . . 22, galindoi

Comb scales 23-32, in 3r rOWS; free portion of midapical scale shorter than

or subequal in length to sessile portion. . . ... . . 23. campana

Hair 7-C short, less than half length of 6-C; hair 11-C short, less than

length of mentum . . Pe. ae

Hair 7-C more than half length of 6-C: hair 1-C longer than mentum. 11

Hair 8-S single; 2-A about 2.0 distal portion of 6-A; anal saddle extending

less than halfway around segment (fig. 14) . . . . . 2. argyrothorax

Schick: Additions to Aedes terrens Group 31

Hair 8-S multiple; 2-A about 3.0 or more length of distal portion of 6-A;

anal saddle extending more than halfway around segment (figs. 10,12)

1. thorntoni

11(9). Hairs 14-C and bmh usually basally branched, rarely single; 11-P usually

atleastO0.50f14P .... . . 12. insolitus; 12a. casali

Hairs 14-C and bmh usually single, when double usually branching about

0.25 or more from base; 11-P usually less than0.5 of 14-P.... 12

12(11). Hair 4-VII at least double; 4-M and 3-III usually at least triple; hairs 10,

12-VII often branched. . Lie 13

Hair 4-VII usually single, rarely double: 4-M and 3- Il with less than 3

branches; :10,12-Vilsingles! oo. oe eee AS

13(12). Ventral brush usually with 12 hairs (11-13); hair 4a-X usually 6-8 branch-

ed (5-9) (northern South America, Trinidad, Tobago). (8) 12b. berlini

Ventral brush usually with 14-16 hairs (13-17); hair 4a-X eae 10-12

branched (9-15) where sympatric with berlini. . . . See

PB OLS 2, S42 Be Et a ees Pa a Ee . . 5. braziliensis

Bis 22-04 6 fet eg ae ee ee 4. tertens: 6. zavortinki

15(12). Saddle extending around segment to at most moderate distance beyond

horizontal midline, submarginal slit absent, ventral margin either with

broad rounded incision or irregular in outline (fig. 52) ges

11. metoecopus; 14. homoeopus

Agee 16. impostor: 1a: gabriel; 20. vargasi; 26. podographicus

Saddle extending around segment far beyond horizontal midline, with

ventral submarginal or marginal slit (figs. 59,60). . ..... . 16

16(15). Comb scales 33-54, in 4 rows, narrow (fig. 59) . . . . 27. tehuantepec

Comb scales 20-29, in 2-3 rows, stout (fig.60) . . . . .28. schroederi

REFERENCES CITED

Bonne-Wepster, Jean and C. Bonne

1920. Diagnoses of new mosquitoes from Surinam, with a note on synonymy. In-

secutor Inscitiae Mens. 7:165-180.

Coquillett, Daniel W.

1906. Five new Culicidae from the West Indies. Can. Entomol. 38:60-62.

Dyar, Harrison G. and F. Knab

1906. Notes on some American mosquitoes with description of new species.

Biol. Soc. Wash., Proc. 19:159-172.

Gordon, Rupert M. and A.M. Evans

1922. Mosquitoes collected in the Manaos region of the Amazon. Ann. Trop.

Med. Parasitol. 16:315-338.

Schick, Robert X.

1970a. Mosquito Studies (Diptera, Culicidae). XX. The Terrens Group of Aedes

(Finlaya). Amer. Entomol. Inst., Contrib. 5(3). 158 p.

32 Contrib. Amer. Ent. Inst., vol. 7, no. 1, 1970

1970b. Mosquito Studies (Diptera, Culicidae). XXIII. Additions and corrections

to the revision of the Aedes terrens group. Amer. Entomol. Inst., Contrib.

7(1):13-40.

Walker, Francis

1856. Insecta Saundersiana. Vol. 1. Diptera. London, van Voorst. p. 415-474.

FIGURES

61. Distribution of the species of the Terrens Subgroup

62. Distribution of the species of the Thorntoni, Alboapicus and Insolitus Sub-

groups

63. Aedes (F.) braziliensis; female mesonotum, femur III and pupa. Aedes (F-.) sp.,

Chaco form; female mesonotum and femur III. Aedes (F.) berlini; male

mesonotum and genitalia .

64. Aedes (F.) braziliensis; larva

65. Aedes (F.) sp., Teresopolis form; female and male mesonotum, femur III of

female, male genitalia and pupa

66. Aedes (F.) sp., Teresopolis form; larva

67. Aedes (F.) casali; female and male mesonotum, femur III of female, male

genitalia and pupa

68. Aedes (F.) casali; larva

INDEX TO SCIENTIFIC NAMES

Names mentioned in the keys only are not included.

alboapicus, 20 insolitus, 20, 20-21, 22, 23; 62f

Alboapicus Subgroup, 19, 20; 62f Insolitus Subgroup, 13, 14, 15, 16, 20, 23;

amabilis, 20; 62f , 62f 3

apollo, 14, 15, 16,17 podographicus, 23-24

argyrothorax, 14, 20 Podographicus Subgroup, 23

berlini, 13, 14, 16, 20, 22, 22-23; 62f, 63f Teresopolis form, 14, 18, 19; 61f, 65f, 66f

braziliensis, 13, 14, 15, 16, 17-18, 18; 61f terrens, 14, 15-17, 17, 18, 19, 24; 61f

63f, 64f Terrens Subgroup, 13, 14-15, 15, 17, 20; 61f

casali, 15, 20, 21-22; 62f, 67f, 68f Thorntoni Subgroup, 14; 62f

Chaco form, 19-20; 62f, 63f zavortinki, 14, 15, 17, 18, 18; 61f

SaTIW Se aa

OOT Os

‘SUALAWOTIN

VORIaINY ICAI

| eype< | ( Naniteagiliaeeso |

SISUAI|IZDIG ©

IYUL}JOADZ

suads9}@) = SNSAYYSL

—

3 a

; 3

CEatSe - Wy

: of ct

a * *)

‘ al * >

02 x

vier

09 7 C

‘ Wed

i A

~ 4

ene

2-S961-1N3-A90100Z-V19N VOIUIWY 3J1GGIW JO S3011NDSOW

SaTIW BL EYES:

000t ool Os 0

00st oot 0010s 0

SYaLAWOTIA

VOLUN SIGN

een: xD40y4}04A610 ©

INOLNYOHL

—<-

~f See

; ek

amis u

SNLITOSNI--

xpsousosABun ce

INOLNYOHL

29 ire vine

& Ba Cees

FINLAYA

VZ 203-103

basal dark band {mm

VZ 372-40

braziliensis

FG 12-14

FINLAYA

A \ y

SS 7 Y Q

Ss : . *sp

—— ‘3

Sunaina reel

8 7 y

braziliensis

FG 12-14

FINLAYA

3 ll 40

MSR TYE

XK) AWN h

By i} )

} h basal dark band

VI 8

Teresopolis form

Rio de Janeiro

Brazil

AAA

fait

Nyt

High

« FINLAYA

! hy ha

Mi hip

\ mies Wy

ee 9

ANU WN eM

aA Mig

ARNT)

WP mi"

UR Minh {

WWE UN aN H NY

Weed inv ty I DV IAM

AE hn

may aie Miia

oi) nba 4

ef thi Vii at

rth

HAT: Hy

iret at

dss Ata Nt

i) }

‘th hy

UTA

ARG 773-109

fee

No e ARG 618-103

)

ae:

Argentina

casali

ARG 618-108

1.0

ARG 618-108

No.

(Continued from inside front cover)

Parts of Volume 4, with prices

Mosquito studies (Diptera, Culicidae), 1968.

X. Peters, T. Michael. Dixinae originally described from North America. 7 pages.

XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes

originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay.

29 pages. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XII. Berlin, O. G. W. A revision of the Neotropical subgenus Howardina of Aedes.

Carlson, Robert W. and Fred B. Knight, 1969.

Biology, taxonomy, and evolution of four sympatric Agrilus beetles (Galeoutera.

Buprestidae). 104 pages, 47figures. Price: $3.25, postpaid.

. Mosquito studies (Diptera, Culicidae), 1969.

XII. Barr, Ralph A. and Sylvia Barr. Pupae of the genus Psorophora in America

north of Mexico. 19 pages, 11 figures.

XIV. Berlin, O. G. W. Redescription of Toxorhynchites (T.) gigantulus from the

Philippines. 7 pages, 2 figures.

XV. Zavortink, Thomas J. A new species of treehole breeding Anopheles from

the southwestern United States. 12 pages, 4 figures.

Price: $1.25, postpaid.

Matthews, Robert W., 1970. A revision of the genus Spathius in America north of Mexico

(Hymenoptera, Braconidae). 86 pages, 31 figures. Price: $3.50, postpaid.

postpaid.

Parts of Volume 5, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XVI. Zavortink, Thomas J. A new species of treehole breeding Aedes (Ochlerotatus)

from southern California. 6 pages, 2 figures.

XVII. Adames, Abdiel J. and Charles L. Hogue. Two new species of Deinocerites from

Costa Rica. 6 pages, 6 figures.

XVII. Berlin, O. G. W. The subgenus Micraedes of Culex. 30 pages, 12 figures, 1 map.

Price: $2.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XIX. Zavortink, Thomas J. The treehole Anopheles of the New World. 35 pages, 12

figures. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XX. Schick, Robert X. The Terrens Group of Aedes (Finlaya). 158 pages, 60 figures.

Price: $5.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

V. Reinert, John F. Genus Aedes, subgenus Diceromyia Theobald in southeast Asia.

43 pages, 15 figures. Price: $1. 50, postpaid.

Howden, Anne T. The Tanymecini of the West Indies (Coleoptera: Curculionidae). 174 pages,

76 figures. Price: $2.75, postpaid.

Kethley, John B., 1970. A revision of the family Syringophilidae eae Acarina).

76 pages, 38 figures. Price: $2.75, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

VI. Mattingly, P. F. The genus Heinzmannia Ludlow in southeast Asia. 104 pages,

dl figures. Price: $4.00, postpaid.

Parts of Volume 6, with prices

. Mosquito studies (Diptera, Culicidae), 1970.

XXI. Belkin, John N., Sandra J. Heinemann, and William A. Page. The Culicidae of

Jamaica. 458 pages, 126 figures. Price: $15. 00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

Vil. Tyson, W. H. Genus Aedeomyia Theobald in southeast Asia. 27 pages, 6 figures.

Vil. Tyson, W. H. Genus Aedes, subgenus Mucidus Theobald in southeast Asia. 54 pages,

21 figures. Price: $3.50, postpaid.

Parts of Volume 7, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXII. Zavortink, Thomas J. A new subgenus and species of Aedes from Arizona.

11 pages, 2 figures.

XXIII. Schick, Robert X. Additions and corrections to the revision of the Aedes

terrens group. 27 pages, 8 figures. Price: $1.50, postpaid.

Contributions

of the

American Entomological Institute

Volume 7, Number 2, 1971

MOSQUITO STUDIES (Diptera, Culicidae)

XXIV. A revision of the crabhole mosquitoes of the

genus Deinocerites. By Abdiel J. Adames.

AGM dah ee

( | \

MAY 24 1971

S/BRARIED

CONTRIBUTIONS

of the

AMERICAN ENTOMOLOGICAL INSTITUTE

The Contributions are for larger papers on insects. Each paper is a separate number, with separate

pagination and index. Separate numbers aggregating about 500 pages constitute a volume. Issues appear

irregularly, as suitable manuscripts are available.

Copies are sold separately or in subscriptions to complete volumes. Complete volumes are $12.00.

The price of separate numbers varies. Subscribers are billed for each volume with its beginning number,

and receive the parts as issued. Orders for separate numbers that total less than $8.00 must be accom-

panied by payment. Address orders or correspondence to the American Entomological Institute, 5950 —

Warren Road, Ann Arbor, Michigan 48105, U.S.A. |

Parts of Volume 1, with prices

No. 1. Dasch, Clement E., 1964. The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae).

75 pages, 69 figures. Price: $2.25, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1965.

I. Belkin, John N., etal. A project for a systematic study of the mosquitoes of Middle America

Il. Belkin, John N., et al. Methods for the collection, rearing and preservation of mosquitoes. |

78 pages, 4 figures. Price: $2.25, postpaid. |

No. 2a. Sameasno. 2, but in Spanish. Price: $2.25, postpaid.

No. 3. Matthews, Robert W., 1965. The biology of Heriades carinata Cresson. (Hymenoptera,

Megachilidae). 33 pages, 23 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1965.

Ill. Ramalingam, Shivaji, and John N. Belkin. Two new Aedes from Tonga and Samoa.

IV. Belkin, John N. The mosquitoes of the Robinson-Peabody Museum of Salem expedition to

the southwest Pacific, 1956.

34 pages, 3 figures. Price: $1.00, postpaid.

No. 5. Mosquito studies (Diptera, Culicidae), 1965.

V. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from Middle America. 95 pages. Price: $2.75 postpaid.

No. 6. Mosquito studies (Diptera, Culicidae), 1966.

VI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from North America. 39 pages. Price: $1.00, postpaid.

No. 7. DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae (Homopotera,

Cicadellidae). A synopsis of the genus Ponana.

86 pages, 257 figures. Price: $2.50, postpaid.

No. 8. Contributions to the mosquito fauna of southeast Asia, 1967.

I. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald in Thailand.

55 pages, 20 figures. Price: $1.75, postpaid.

Parts of Volume 2, with prices

No. 1. Contributions to the mosquito fauna of southeast Asia, 1967.

Il, Bram, Ralph A. The genus Culex in Thailand (Diptera: Culicidae).

296 pages, 104 figures. Price: $9.00, postpaid.

No. 2. Schuh, Toby, 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region.

35 pages, 30 figures. Price: $1.25, postpaid. |

No. 3. Snider, Richard J., 1967. The chaetotaxy of North American Lepidocyrtus s. str. (Collembola:

Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid. :

No. 4. Contributions to the mosquito fauna of southeast Asia, 1968. ;

Ill. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald of southeast

Asia. 76 pages, 26figures. Price: $2.25, postpaid.

No. 5. Contributions to the mosquito fauna of southeast Asia, 1968.

IV. Knight, Kenneth L. Species of the subgroup Chrysolineatus of group D, genus Aedes,

subgenus Finlaya Theobald. 45 pages, 12 figures. Price: $1.50, postpaid.

Parts of Volume 3, with pfices \

No. 1. Mosquito studies (Diptera, Culicidae), 1968.

VIl. Belkin, John N. The Culicidae of New Zealand.

182 pages, 30 figures. Price: $5.50, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1968.

VIN. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia.

221 pages, 41 figures. Price: $6.75, postpaid.

No. 3. Fleming, Richard C., 1968. Head musculature of sphinx moths (Lepidoptera: Sphingidae).

32 pages, 37 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1968.

IX. Belkin, John N. The type specimens of New World mosquitoes in European museums,

69 pages. Price: $2.00, postpaid.

(Continued on back cover)

MOSQUITO STUDIES (Diptera, Culicidae)

XXIV. A REVISION OF THE CRABHOLE MOSQUITOES

OF THE GENUS DEINOCERITES

Abdiel J. Adames”

CONTENTS

ENTRODUCTION .. .. 2

MATERIAL AND METHODS . a

TAXONOMIC CHARACTERS 3

SYSTEMATICS 5

BIONOMICS . . Yate ha, Lee, Reet aaa, Slats Ge gk te Renae. Eames Does

MEDICAL AND PEST IMPORTANCE pith See Sta, gh (AEA SS gap RehiNy, MPR ae BRT te, A

TAXONOMIC TREATMENT . ie ae Rn achat Oe |

Genus Deinocerites . i

Keys to Groups and Species. CM rata ue ys) aa! 2 are acho vices tol es a Re

Spanius Group. . . OTR eA Rn nue. Ak NRIGT IN AOE. SEI E aap ng: hake 2S

1. Deinocerites spanius wh iA MON i ik se a i | RN Ck a a ae

BUC COMICS GION TOUS (on 45 PARE 8 Wiis aeg CAN Seb a? oe

SIO COMICS THOT CSOT ry) eae REDS ES Tenge ETN TED EE oF ae

eM OCCIIIES DCTRIME 8 ir alle ORT re Sek LE RR Te AP aaa

DyariGroup . . . ee Tete Med 8 ORday Rea a AAG TARE) Mk TANT, OPS RANE, AMAR Mtl a Ne Seay

5. Deinocerites aarti i HIE Oye ke, eh aN ad 6 EGE Raat), ATL EMP IR geletig cit EE

CMP CTACETITEG DEITCH MOA vito! PU ole AMA hk CO RN Med, ap hl ot a)

7. Deinocerites fe aieed ee Seg not ae Aa UN Mas es es Ne ee nhs ine TL 2)

Mancer Group 30°. PARA 5: POR ANC EA | Bees oy AEA RIAA, | a EAR a

Mis CEPILER COMICON tk for wae ME, gk OM A ONG LG tani? imran ew aehe iia

y 1 einocerites MIClQnHODAVIGIN: (2) ON CR CE oe SOR AQ

MIE CLNOCETIICS THGGRES Vinci 8b Spee my emer ee ee oe a na

‘Contribution from project “Mosquitoes of Middle America” supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478. Based on PhD dissertation submitted to the University

of California, Los Angeles.

* Department of Zoology, University of California, Los Angeles, California 90024. Present

address: Gorgas Memorial Laboratory, Apartado 6991, Panama 5, R. de P.

2 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Epitedeus Group. . . ee ie ey a nn

11. Deinocerites epitedeus Oe ee ee a ee

12 Deporeiies Panes we ee

WSs Deo eeries POO 8 a a a OG

PA Dei CerIeNUsIor ices a a a a a

Po. Depp oe a ee eee

Pseudes Group. . . ee ee a aoa ee

16. Deinocerites pseudes pO ee ee Oe ee

17) Deimoeees WOW a sa ee es ee

1S. Democeriies econ ea ee ea ee

REPERENGC ES CIE oo a a a ee

FIGURES. . oo a a

INDEX TO SCIENTIFIC NAMES Pee ee ee a

INTRODUCTION

In the 12 years since the appearance of Belkin and Hogue’s review (1959) of

the genus Deinocerites a great deal of material has accumulated and considerable

information has been obtained on the bionomics and behavior of some of the

species. Recent studies have shown Deinocerites pseudes to be naturally infected

with the viruses of St. Louis encephalitis and of Venezuelan equine encephalo- —

myelitis (Tempelis and Galindo, 1970:175). It was therefore considered appropri-

ate to thoroughly revise this biologically interesting and potentially medically im- ©

portant genus. While Belkin and Hogue recognized 11 species (1 not named), in ©

only 3 of these were both the larvae and pupae known and in 1 only was there ©

definite association of all stages. In the present revision, I recognize 18 species —

and in 16 of these all the stages are known and associated through individual —

rearings. |

I am indebted: to John N. Belkin for guidance in the study and the prepara-

tion of the manuscript; to present and past members of the project “Mosquitoes —

of Middle America’”’, particularly Thomas J. Zavortink, O.G.W. Berlin, Robert X. —

Schick, also to my fellow students, Michael Nelson and Jose Valencia, for much —

assistance and stimulating discussions; and to William A. Powder and Sandra J.

Heinemann for the preparation of material. For the loan of material I thank: Alan

Stone, U.S. National Museum; Charles L. Hogue, Los Angeles County Museum —

of Natural History; Donald B. Bright, California State College, Fullerton; Bruce —

Eldridge, Walter Reed Army Institute of Research; Alfonso Diaz Najera, Instituto —

de Salubridad y Enfermedades Tropicales, Mexico City; Pablo Barreto, Universi- —

dad del Valle, Cali, Colombia; and Martin D. Young and Pedro Galindo, Gorgas —

Memorial Laboratory, Panama, who also provided laboratory facilities in the field. —

I am also grateful to the Organization of American States for a fellowship that —

allowed me to pursue the initial part of this study and to the National Science —

Foundation which supported some of the field work through grant GB 3871. For —

the preparation of the final illustrations I am indebted to Nobuko Kitamura and ©

for the text copy for lithoprinting to Caryle Abrams.

MATERIAL AND METHODS

MATERIAL. Some 24,366 specimens were examined for this study, 3,005 males,

Adames: Genus Deinocerites 3

4.679 females, 13,298 larvae and 3,384 pupae; included were 2,009 individual rear-

ings (1112 larval, 680 pupal, 217 incomplete) of the 16 of the 18 known species.

The bulk of this material was collected for the project “Mosquitoes of Middle

America” (Belkin, Schick et al, 1965, 1967) and is deposited at the University

of California at Los Angeles [UCLA]. Other collections used were those of the

U.S. National Museum [USNM] and of the project on tropical land crabs and

their associates [LCBA] being conducted by Charles L. Hogue and Donald B. Bright.

Smaller loans were also made available from the Instituto de Salubridad y Enfer-

medades Tropicales, Mexico [ISET], Universidad del Valle, Cali, Colombia, and

Gorgas Memorial Laboratory in Panama. I collected additional material and made

field observations specifically for this project in Honduras, Nicaragua, Panama and

Colombia in August and September 1967.

TAXONOMIC PROCEDURE. The taxonomic methods used here are the clas-

sical comparative morphological ones. Species were first recognized on the basis

of a series of constant correlated morphological characters in all stages. They were

then compared minutely and those showing correlated features in common were

assembled into primary phyletic lines called groups. In some instances smaller units,

species complexes, were recognized. In the final interpretation of evolution in the

group, distributional data were also taken into account.

DESCRIPTIONS. The method of presentation, terminology and abbreviations

used in the description of the taxa in general follow Belkin (1962). A few spec-

ial terms introduced in this revision are self-explanatory and are labelled on the

_ figures.

ILLUSTRATIONS. For every species all the available stages are illustrated. The

full chaetotaxy of the immature stages is figured, with a modal value of the hair

branching based on 5-10 topotypic specimens (exact number specified in the des-

cription of each species). The illustrations of the adults show pertinent details

only: male and female heads, male and female genitalia, female forefemur, and

claws of foreleg and midleg of female. In addition, the female of cancer, the type

Species of the genus, is illustrated completely.

DISTRIBUTION. Only synoptic distributional data are given here: locality, col-

lector, collection number (when available) and depository. Full data for every spec-

imen examined have been entered in a permanent ledger and will be available on

demand to interested parties on microfilm or other desired reproduction method

from the Department of Zoology, University of California, Los Angeles.

TAXONOMIC CHARACTERS

The following morphological features of the different stages were found to be

particularly useful as diagnostic specific characters as well as taxonomic or group

characters. I have indicated whenever possible the primitive or derived states of

the various characters. There is no absolute criterion for determining the nature

of the character state without paleontological evidence but the derived state is

usually indicated by the trend within a taxon and the primitive state by com-

parison with related taxa, in this case the subgenus Culex and the genus Galindo-

myia (see under affinities in the section on systematics).

GENERAL ADULT CHARACTERS. Coloration. — The color of the integument

of the pleural sclerites as compared with that of the mesonotal integument is a

useful diagnostic specific character but is of no value as a group character.

A. Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Antenna. — The elongation of the basal flagellar segments and the consequent

elongation of the entire antenna are the most important taxonomic characters in

the general external morphology of the adults. The trend within the genus is from

an antenna similar in the 2 sexes and with only the first flagellar segment elon-

gate to dissimilar and with additional flagellar segments elongated. The length of

the antenna is expressed in terms of its extension beyond the proboscis with ref- —

erence to specific flagellar segments. This was determined in slide preparations.

Proboscis. — The length of the proboscis, from the basal ventral bristles to the

tip of the labella, was determined in slide preparations. As noted above, the pro-

portional length of the antenna and proboscis is the most useful taxonomic char-

acter in the adults.

Mesepimeron. — The development of a large patch of scales on the mesepimeron

is interpreted as a derived state in Deinocerites. It appears to have arisen inde-

pendently in the Pseudes and Epitedeus Groups. 7

Forefemur. — The replacement of the simple bristles of the anteroventral and

posterodorsal rows by short spiniforms is a derived condition. It appears to have

arisen independently in the Pseudes and Epitedeus Groups.

FEMALE SEXUAL CHARACTERS. The female genitalia were studied in part

only. The following characters showed taxonomic as well as diagnostic value.

Sternite VII. — Excellent taxonomic characters are (1) presence (primitive) or

absence of scales on the distal sclerotized part and (2) caudal margin without (prim-

itive) or with strongly differentiated bristles.

Postgenital plate. — Primarily of diagnostic specific value. Only the distal part

was studied, in lateral as well as ventral aspect.

Cercus. — The general shape as well.as the differentiation (derived) of apical

and subapical setae are excellent taxonomic as well as diagnostic specific characters.

MALE SEXUAL CHARACTERS. These characters are primarily diagnostic at

the specific level but are of some value in classification except for the claws.

Claws. — The length, shape and dentition of the foreclaws and midclaws of

nearly every species is uniquely developed.

Tergite IX. — The articulated lobe of tergite IX shows a characteristic devel-

opment in most species. It is of value as a group character primarily in the Can-

cer Group where it is markedly flattened (derived).

Sidepiece. — The presence (primitive) or absence of scales on the sidepiece is —

correlated with a similar development of sternite VIII of the female and is an

excellent group character.

Aedeagus. — The intromittent organ is basically similar throughout the genus

except-for the absence (primitive) of a dorsal basal articular process for the dorsal

paramere in the Spanius Group.

Dorsal paramere. — As with the claws and IX tergite lobe, the dorsal paramere

is characteristically developed in most species. It is differentiated into 2 parts (prim-

itive) in the northern complex of the Spanius Group but is compact in all other —

species.

PUPA. The following pupal group characters are usually concordant with the ©

group characters of the adults: (1) on the cephalothorax, the increase (derived)

in the length of and in the distance between hairs 2 and 3, the reduction (derived)

in the length of hair 5, (2) on the metanotum, the reduction (derived) in the

length of hair 10, and (3) on the abdomen, increase in length and reduction in

the number of branches in hair 1-II (both derived), the displacement of 5-II me-

sad (derived), and the reduction (derived) in the length of 1-VII. Other important

Adames: Genus Deinocerites 3 5

group characters are found in cephalothoracic hairs 5,7,8-C and abdominal hairs

1-II and 3-V. There are numerous diagnostic specific hair characters as indicated

in the pupal Key.

FOURTH INSTAR LARVA. There are numerous characters in the fourth in-

star larva that show congruent development with adult and pupal characters at

the group level and are very useful in classification. Among these are (1) on the

head, reduction and lateral displacement (both derived) of hair 2-C, and reduc-

tion in number of branches and elongation (both derived) of 6-C and to lesser

extent 5-C, (2) on the thorax, branching (derived) of hair 9-P, (3) on the abdo-

men, increase (derived) in the number of branches in hairs 6-II and 6-VI, and

decrease (derived) in the length of 1-VII, (4) on the siphon, decrease (derived)

in the number of branches of hair 1-S, and (5) on the anal segment, independent

reduction (derived) in several lines of the number of hairs in the ventral brush

from. 7 to 6 pairs. There are also numerous specific differences in the branching

of other hairs as indicated in the larval key.

SYSTEMATICS

TAXONOMY. The 18 species recognized in this revision fall into 5 well-defined

groups on the basis of morphological similarities in 1 or more of the different

stages. Two of these groups, Spanius and Cancer, are essentially as defined by

Belkin and Hogue (1959); the Dyari Group of these authors is broadened to in-

clude 2 other species; and the Pseudes Group is split off from the Epitedeus Group.

Each of the groups is marked by 1 or more unique features or unique com-

binations of features in 1 or more stages. In all but the Spanius Group these fea-

tures represent derived states. They are listed below in the diagnosis of each group

and are only briefly reviewed here.

In the Spanius Group unique features are present in the adults (extensive scal-

ing of the genitalia and short similar antennae in both sexes) and the larvae (ab-

dominal hair 6-II single); all of these appear to be primitive. In the Dyari Group

the best diagnostic features are found in the immature stages (reduced length of

pupal hair 10-C and larval hair 1-VII). The Cancer Group is differentiated from

the others most clearly by the genitalia in both sexes (flattening of IX tergite lobe

in the male and spatulate specialized setae of the cercus in the female). The Epi-

tedeus Group (in the present restricted sense) is the most derived of all and is

clearly marked by (1) the absence of a strongly developed lower mesepimeral bris-

tle in the adults and the great elongation of flagellar segments 1-4 of the antenna

of the female, (2) the reduced length of hair 5-C and the long double or triple

hair 1-II in the pupa, and (3) hair 9-P. double or triple in the larva. Finally, the

Pseudes Group shares with the Epitedeus Group the development of a patch of

mesepimeral scales in the adults (absent in the other groups) but possesses the

differentiated mesepimeral bristle; its immature stages however are more similar

to those of the Cancer Group from which they are not always easily differentiated.

The Spanius Group appears to be the most ancient segregate of the Deinocerites

stock because it is characterized largely by primitive and not derived features and

because of its widely disjunct distribution. The Dyari Group has also retained many

primitive features and in some respects appears to be transitional between the Span-

jus Group and the others with predominantly derived features, but as indicated

above it too possesses unique derived features in the immature stages.

6 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

DISTRIBUTION. The genus is largely restricted to Middle America in the sense

of Belkin, Schick et al (1965:2), i.e. Central America, the West Indies and adja-

cent portions of North America and South America. The only known extension

from this area is on the Atlantic seaboard of South America down to the state

of Maranhao, Brazil, south of the Amazon River.

All the species within a given group are allopatric except for the Pseudes Group

where all 3 species have been recorded from a single locality. Only 1 species,

pseudes, is known with certainty to be present both in the Atlantic and Pacific

basins, all others are confined to either one or the other ocean basin.

Each of the groups has a characteristic distribution but 2 or more groups occur

in 1 locality. The Cancer Group is confined to the Atlantic basin, centered around

the Caribbean Sea but extending into the Bahamas and Florida in the north and

into eastern South America south of the Amazon. The Dyari Group is known

only from the Pacific shores from Nicaragua to central Colombia. The other 3

groups have been reported from both the Atlantic and Pacific coasts. The Pseudes

Group is largely confined to the Pacific coast from the Gulf of California to the

Gulf of Guayaquil with only pseudes itself invading the Atlantic basin northward

from the Isthmus of Tehuantepec, Mexico, into southern Texas. The Epitedeus

Group is equally represented on the Atlantic and Pacific sides of Central America

and Colombia. Whereas the distributions of all these groups are probably contin-

uous within their range, that of the Spanius Group is widely disjunct, with a pair ©

of geminate Atlantic and Pacific species north of Tehuantepec and a similar pair

south of Costa Rica.

All of the 5 groups are known from Panama, 4 on the Pacific side (Spanius,

Dyari, Epitedeus and Pseudes) and 3 on the Atlantic (Spanius, Cancer and Epi-

tedeus). Elsewhere at most 2 groups occur sympatrically in the Atlantic basin (Span-

ius and Pseudes, north of Tehuantepec; Cancer and Epitedeus, Central America;

Spanius and Cancer, Colombia and Venezuela) and at most 3 in the Pacific basin

(Spanius and Pseudes, north of Tehuantepec; apparently only Pseudes in Guate-

mala and Salvador and Pseudes and Dyari in Nicaragua; Pseudes, Epitedeus and

Dyari in Costa Rica and Colombia; and only Pseudes in Ecuador and northern

Peru). In Florida, Bahamas, Greater and Lesser Antilles, Trinidad and Tobago, and

northeastern South America only the Cancer Group is represented.

AFFINITIES. The genus Deinocerites is undoubtedly a member of the tribe

Culicini in the sense of Belkin (1962:117). Only 2 other genera are currently rec-

ognized in this tribe, the dominant worldwide Culex Linneaeus, 1758 and the mon-

otypic Galindomyia Stone & Barreto, 1969 from Colombia.

Deinocerites and Galindomyia share the following features not exhibited by Cu-

lex: (1) flagellar segment 1 elongate in both sexes, (2) dorsal paramere developed

in the male, and (3) general development of the subapical lobe of the sidepiece

and of the clasper in the male. Galindomyia leei Stone & Barreto, 1969 resembles

members of the most primitive group (Spanius) of Deinocerites in the small size,

relatively short antenna and the type of articulation of the dorsal paramere with

the aedeagus. Therefore, I consider it likely that Deinocerites and Galindomyia

were derived from a common ancestral stock which separated very early from the

stock that gave rise to the dominant genus Culex. However, the relationship be-

tween Deinocerites and Galindomyia cannot be determined with any degree of

certainty without evidence from the immature stages. Unfortunately the latter are

unknown for leei; it seems likely that they will be found in crabholes to which

the adults appear to be confined as in the case of Deinocerites. For the present,

Adames: Genus Deinocerites 7

Galindomyia is retained as a genus distinct from Deinocerites because of the fol-

lowing features not found in any of the groups of the latter: (1) absence of artic-

ulation between tergite and sternite IX, poor development of tergite lobe IX, and

unique development of terminal flagellar segment of the antenna and of the fore-

claw in the male, and (2) vertex of head with decumbent scales all broad and erect

scales restricted to occiput in both sexes. In some respects Galindomyia appears

to be annectent between Deinocerites and Culex, notably in the position and de-

velopment of the subapical lobe of the sidepiece.

Dyar’s view (1928:261,337) that Deinocerites was derived from Culex (Tino-

lestes) latisquama (Coquillett, 1906) was apparently based entirely on the mis-

taken homology of a process from the basolateral sclerotization of the proctiger

with the IX tergite lobe of Deinocerites and the presence of a short palpus in

the male, and the similarity in breeding sites. Actually there is little in the morph-

ology of the adults of latisquama to suggest close relationship to Deinocerites and

its immature stages are completely different and indicate affinity with the Melano-

conion line of the genus Culex.

Of the major lines of Culex, the subgenus Culex appears to share the greatest

number of similarities with Deinocerites. Among these are: (1) in the adults, de-

cumbent scales of vertex all narrow, erect scales numerous, (2) in the genitalia,

sidepiece of male and sternite VIII of female with scales, and (3) in the larva,

abdominal hairs 6-I-VI subequal, with 6-I,II not sharply differentiated, and siphon

with only 1 lateral hair out of line.

SPECIATION AND PHYLOGENY. Although the interrelations of the various

species are now better understood because of the knowledge of all the stages of

16 of the 18 species, the other data needed to determine the speciation and phy-

logeny of the genus are still fragmentary and little can be added to the specula-

tions of Belkin and Hogue (1959:420-423) except at the species and group level

(see).

Many areas are still not surveyed for Deinocerites and it is very probable that

some of the existing gaps in distribution will eventually be filled. Accurate deter-

mination of distributions may be important to evaluate the places of origin and

the relationship to former sea portals in the area for it appears that even dom-

inant species of Deinocerites are incapable of crossing relatively small land and

sea barriers and that dispersal occurs only through contiguous areas suitable for

breeding. This is rather clearly seen in Panama where all the species are restricted

either to the Atlantic or to the Pacific basin although there is less than an 80-

kilometer gap between the basins and a connection by way of the canal. The

few specimens in this area reported from the wrong basin are probably strays as

there is no modern record of any species breeding outside of its normal basin.

The present allopatric pattern of species within groups of Deinocerites is high-

ly suggestive that speciation in this genus has taken place primarily in conjunc-

tion with geographical isolation. Only in the Pseudes Group do the included spec-

ies occur together and in case of howardi (see), as pointed out by Belkin and

Hogue (1959:442), it is possible that speciation took place through hybridization

or introgression (between mcdonaldi and pseudes stocks).

If the above pattern of allopatric speciation has occurred in Deinocerites in the

past, then the present partially overlapping pattern of the distribution of the 5

groups represents subsequent dispersal of the original stocks over more or less con-

tiguous areas of suitable breeding sites. Thus, it may be possible to get some sug-

gestions as to barriers (sea portals) from the distributions of species within a group

8 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

and both barriers and connections (or contiguities) from those of groups of Dein-

ocerites. It is tempting to date these by correlating them with geological events.

Unfortunately the historical geology of the area occupied by the genus is poorly

known and subject to different interpretations, sometimes in part based circularly

on the distribution of plants or animals. Up to now, no fossil material of Deino-

cerites is known and it is therefore impossible to date directly any of the events

in the phylogeny of the genus.

However, the general sequence of events in the evolution of Deinocerites may

have been as proposed by Belkin and Hogue (1959:423) with slight modifications

as follows:

1. The origin of the Spanius Group stock on an island in the center of the

Middle American area.

2. Subsequent connection of this island with both North and South America

and the invasion of these areas by the Spanius Group.

3. Fragmentation of the central area into islands and the formation of 4 sep-

‘arate lines from the Spanius Group populations, each on a different island and

giving rise in the southwest to the Dyari Group, the northeast to the Cancer Group,

the northwest to the Pseudes Group and the southeast to the Epitedeus Group.

4. Subsequent connection of the islands containing the different stocks and fur-

ther species differentiation in the.groups by later fragmentation of portions of

the various connected areas. This was probably a very complex series of events

and extended over a long period of time. The latest fragmentation appears to have

taken place in the region of Panama and northwestern South America, but the

presence of an undifferentiated population of the dominantly Pacific pseudes in

the Gulf of Mexico suggests that even more recently there was a gap (mangroves?)

through the isthmus of Tehuantepec which allowed this species to cross over into

the Atlantic basin.

BIONOMICS

All species of the genus Deinocerites normally utilize as breeding sites, as well

as adult resting sites, burrows of land crabs of the families Gecarcinidae and Ocypo-

didae. In Cuba, cancer has been reported also from burrows of species of stream

inhabitating crabs of the genus Epilobocera of the family Pseudothelphusidae (Mont-

chadsky and Garcia, 1966:47). No conclusive evidence has yet been obtained as

to the specificity of association of any given mosquito with a particular species

of crab and relatively few positive identifications of the burrow makers have been

reported. However, it appears that members of the Spanius Group are normally

found in small burrows which are usually made by fiddler crabs of the genus Uca

which contains small species of the family Ocypodidae; only 2 species have been

identified, both associated with mathesoni: Uca pugilator (Bosc, 1802) and Uca

subcylindrica (Stimpson, 1859). Two species of the Dyari Group (dyari and ni-

coyae) have been found associated with the Wide Red Land Crab, Ucides occi-

dentalis (Ortmann, 1897), a medium-sized species of the family Ocypodidae. Spec-

ies of the Cancer, Epitedeus and Pseudes Groups are usually found in large or

medium-sized burrows made by members of the family Gecarcinidae, primarily

the large Cardisoma crassum Smith, 1870 (Mouthless Crab; Pacific) or Cardisoma

guanhumi Latreille, 1825 (Great Land Crab; Atlantic) and less frequently the med-

Adames: Genus Deinocerites 9

ium-sized Atlantic Black Land Crab, Gecarcinus lateralis (Freminville, 1835).

The immature stages of species of Deinocerites have occasionally been reported

from a variety of abnormal habitats such as postholes (Jennings in Howard, Dyar

and Knab, 1915:209), rockholes (Wirth, 1945; Belkin, Heinemann and Page, 1970:

49), a beach pool (van der Kuyp, 1948a:748), treeholes (Peyton, Reinert and Pe-

terson, 1964:454; Porter, 1964:222) and various artificial containers such as tin

cans, a wooden bucket and an abandoned septic tank (Dyar, 1928:265; Porter,

1964:222). They have also been found in ground pools formed by the flooding

of depressions containing crabholes.

It is not unusual to find the immature stages of more than | species of Deino-

cerites in the same crabhole but these always belong to different species groups

except possibly in the case of pseudes, howardi and mcdonaldi which are partially

sympatric members of 1 group. Frequently associated with species of Deinocerites

are the immature stages of the other normally obligate crabhole mosquitoes. To

date the following species have been recorded in such associations: Culex (C.) jan-

itor Theobald, 1903; members of Culex (C.) inflictus Theobald, 1901 complex;

Culex (Melanoconion) carcinophilus Dyar & Knab, 1906; Culex (Mel.) undescribed

spp.; Culex (Tinolestes) latisquama (Coquillett, 1906). In exposed shallow crab-

holes, in flooded crabhole areas and other ground water situations, the following

Species have been found associated with various species of Deinocerites: Aedes

(O.) taeniorhynchus (Wiedemann, 1821); Aedes (O.) scapularis (Rondani, 1848);

Anopheles (A.) eiseni Coquillett, 1902; Anopheles (A.) grabhamii Theobald, 1901;

_ Anopheles (Nys.) albimanus Wiedemann, 1821; Culex (C.) bahamensis Dyar & Knab,

1906; Culex (C.) habilitator Dyar & Knab, 1906; Culex (C.) nigripalpus Theobald,

1901; Culex (Mel.) elevator Dyar & Knab, 1906 complex; Culex (Mel.) opisthopus

Komp, 1926; Psorophora (G.) jamaicensis Theobald, 1901. In treeholes or various

artificial containers, 2 species of Deinocerites (cancer and costaricensis) have been

reported to be associated once each with Aedes (F.) triseriatus (Say, 1823) com-

plex; Culex (C.) nigripalpus; Culex (Anoed.) conservator Dyar & Knab, 1906; Hae-

magogus chalcospilans Dyar, 1921; Orthopodomyia fascipes (Coquillett, 1905); and

Corethrella (C.) sp.

Immature stages of Deinocerites occur in crabholes near the seashore as well

as in those at a considerable distance from the coast. The water in the burrows

has been reported to vary from completely fresh to distinctly salty but the only

published records of water analyses are for mathesoni and pseudes and show a

range of chloride content from 1,115 to 8,430 ppm and pH from 7.2-7.6 (Fisk,

1941:547; Peyton, Reinert and Peterson, 1964:452).

The natural food of the larvae is unknown but may be, as speculated by How-

ard, Dyar and Knab (1915:200), “‘matter in suspension in the water, of which

the excrement of the crab not improbably forms an important part’’. As pointed

out by Belkin and Hogue (1959:415) the great development of the subantennal

pouch, the peculiar basal and apical lobes of the mandible and the rudimentary

mental plate suggest an unusual type of food and a peculiar method of feeding.

In the laboratory, larvae have been reared through from eggs on Purina chow (Ga-

lindo, 1967) and half and half mixture of ground dog food and porcine liver pow-

der (Gentry, Gerberg and Hopkins, 1970).

The length of the aquatic cycle under natural conditions is not known for any

species of Deinocerites. Under laboratory conditions it has been determined only

for mathesoni, cancer and pseudes. The reported length of the larval cycle varies

from only 2 weeks to more than 8 weeks; while the pupal stage is stated to last

10 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

from about 3-8 days. In the laboratory, eggs are laid singly in containers above

the water level.

All available data indicate that adults of Deinocerites are crepuscular or noc-

turnal in activity and that during daylight hours they rest in the upper portions

of crab burrows. If the adults are disturbed or driven out of the holes, they fly

out a short distance only and return rapidly to the burrows. Dyar (1928:126)

suggested that “‘the remarkably elongate antennae perhaps serve as special sensory

organs to warn them of ... danger ... when the crustacean host enters’’. As the an-

tennae are elongate in both sexes it is likely that they serve to monitor all the

activities of the adults within the confines of the dark crabholes. As shown by

Downes (1966) and Provost and Haeger (1967) the antennae of the males are in-

volved in the mating behavior of cancer.

It appears that the adults have a relatively short flight range and that their

activities are restricted to the immediate vicinity of their breeding sites in the

crabholes. However, mathesoni (Fisk, 1941:544) and cancer (Branch, Logan et al,

1958:161) have been reported in light traps presumably several miles away from

crabholes. |

Earlier workers generally assumed that the majority of the species of Deino-

cerites do not take blood or at least not from warm blooded vertebrates (Howard,

Dyar and Knab, 1915:200; Fisk, 1941:544). It is now known that a number of

species are attracted to a variety of hosts in animal-baited traps and some will

readily feed on man in the field. Recent studies on the blood meals of wild-caught

females of 5 species of Deinocerites in Panama (Gorgas Memorial Institute, 1970:

20; Tempelis and Galindo, 1970) have revealed a wider spectrum of hosts for some

of these species than has been known for any other group of mosquitoes prev-

iously studied, ranging from man and several domestic and wild mammals through

birds of several orders, lizards, frogs and toads. Specific differences in host pref-

erence were also noted in these studies and also the unusually small blood meals

taken by species of Deinocerites.

Early reports on the mating behavior of species of Deinocerites were based on

field observations and indicated that swarming and mating take place above the

crabholes (Busck, 1908:61; Howard, Dyar and Knab, 1915:207). Recently, ob-

servations have been made in laboratory colonies of cancer by Downes (1966) and

Provost and Haeger (1967) and pseudes by Galindo (1967). In cancer 2 types of

mating behavior were seen. In the first type, males exhibited ‘“‘pupal attendance”’

on the water surface which is described here in the discussion of the bionomics

of cancer. In the second type, males flew to the walls of the container and cop-

ulated with virgin females without pupal attendance. Galindo did not report either

swarming or pupal attendance in the laboratory colony of pseudes but observed

males performing nuptial flights involving hovering over females and contact be-

tween midtarsi of the males and the hindlegs of the females prior to copulation

end to end. It remains to be seen if all the different types of behavior described

occur in a single species in different situations in the field or if different species

have developed different mating behavior patterns.

MEDICAL AND PEST IMPORTANCE

Recent data suggesting the potential importance of species of Deinocerites as

vectors of arboviruses have been reviewed by Tempelis and Galindo (1970: 177-

Adames: Genus Deinocerites 11

178). As noted in the section on bionomics, some species of the genus readily

feed in nature on a variety of cold-blooded and warm-blooded vertebrates as well

as man. That such “‘mosquitoes [are] promising suspects as vectors of viruses which

have been isolated from both warm- and cold-blooded animals” (Gorgas Memorial

Institute, 1970:20) has been demonstrated by the isolation from wild-caught pseu-

des of the virus of St. Louis encephalitis (SLE) and of Venezuelan equine enceph-

alomyelitis (VEE) and by the transmission of the latter in the laboratory by the

same species (Grayson, Sirihongse and Galindo, 1967; Gorgas Memorial Institute,

1970:19; Tempelis and Galindo, 1970:175). However, the importance of species

of Deinocerites in the epidemiology of either arboviral infection has not been dem-

onstrated as yet. Although some species are known to bite man rather freely, their

activity appears to be confined largely to the vicinity of areas with crabholes.

Therefore their importance both as pest species and as direct vectors of patho-

gens to man is probably limited.

In other recent studies in Panama (Gorgas Memorial Institute, 1970:20), epi-

tedeus was found to be naturally infected with trypanosomes (60 specimens) and

plasmodium-like sporozoites (2 specimens). The vertebrate hosts of these parasites

have not been determined.

TAXONOMIC TREATMENT

Genus DEINOCERITES Theobald

1901. Deinocerites Theobald, 1901a:235 (15 July), no included species; 1901b:215 (23 Nov.).

TYPE SPECIES: Deinocerites cancer Theobald, 1901, Jamaica; by subsequent monotypy.

1901. Brachiosoma Theobald, 1901a:235, no included species. TYPE SPECIES: Deinocerites

cancer Theobald, 1901, Jamaica; designation of Coquillett (1910:515). Synonymy with

Deinocerites by Blanchard (1905:414).

1901. Brachiomyia Theobald, 1901b:343-344. TYPE SPECIES: Brachiomyia magna Theobald,

1901, St. Lucia; by monotypy. Synonymy with Deinocerites by Theobald (1903:275).

1902. Deinokerides Giles, 1902:335,472. TYPE SPECIES: Deinocerites cancer Theobald, 1901,

Jamaica. Unjustified emendation.

1905. Dinocerites Blanchard, 1905:413-414. TYPE SPECIES: Deinocerites cancer Theobald,

1901, Jamaica. Unjustified emendation.

1907. Dinomimetes Knab, 1907:120. TYPE SPECIES: Dinomimetes epitedeus Knab, 1907,

On Limon, Costa Rica; by monotypy. Synonymy with Deinocerites by Dyar (1918:

101).

1909. Dinanamesus Dyar and Knab, 1909:259. TYPE SPECIES: Dinanamesus spanius Dyar

& Knab, 1909, Corozal, Panama, Canal Zone; by monotypy. Synonymy with Deino-

cerites by Dyar (1918:101).

Deinocerites of Theobald (1903:275-276; 1905b:37); Dyar (1905a:45-49; 1928:261); Felt (1905:

459,491); Dyar and Knab (1906:178,188; 1907a:48); Knab (1907b:121-123); Mitchell

(1907b: 33): Busck (1908:54); Coquillett (1910:531); Surcouf and Gonzalez-Rincones

(1911:91); Howard, Dyar and Knab (1915:191); Dyar and Shannon (1924:477,485); Bonne

and Bonne-Wepster (1925:168); Shannon (1931:4-5); Gerry (1932:43); King, Bradley and

McNeel (1944:61,65,66,75,77; 1946b:5,7); Carpenter, Middlekauff and Chamberlain (1946:

25,34,274); Knight and Chamberlain (1948:7); Bates (1949:80,165,171,270,322); Lane

(1953: 535); Blanton, Keenan and Peyton (1955:41); Horsfall (1955: 599-600); Perez Vi-

gueras (1956:476); Peyton and Hill (1957:295); Belkin and Hogue (1959:41 1-451); Gibson

and Carrillo (1959:166); Stone, Knight and Starcke (1959:284-285); Scott (1961:244);

12 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Bachmann and Casal (1963:21); Dodge (1962:366; 1963:798,807; 1966:342-343,344-345);

Evans (1962:255); Ross (1962:192; 1964:103,104); Forattini (1965:107,108); Diaz Najera

(1965:57); Downes (1966:1173-1176); Provost and Haeger (1967:565,570-573); Lee and

Barreto (1969:389).

Brachiomyia of Giles (1902:473-474).

FEMALES. Small to medium-sized species; mesonotum usually dark brown, some-

times contrasting with conspicuously lighter portions of pleural integument; with-

out distinct ornamentation on legs or abdomen. Head: Eyes contiguous above an-

tennal bases. Decumbent scales on disc of vertex all narrow, a small patch of small

broad scales laterally; a slight indication of a frontal tuft of elongate scales or

scalelike bristles; orbital line without scales. Erect scales numerous. One pair of

strongly developed interorbital (frontal) bristles and a continuous row of orbitals.

Clypeus prominent, bare. Proboscis usually distinctly shorter than abdomen. Lab-

ium uniform in diameter; weakened beyond middle and often curved caudad; dark

scaled and with numerous short bristles; labella small, elongate, with short bris-

tles and usually a few scales. Palpus short, about 0.2 of proboscis; 3-segmented

and with an indistinct palpifer; segment 3 elongate, dark scaled; all 3 segments

with bristles but only a few well developed in distal part of segments 1 and 2.

Antenna always distinctly longer than proboscis, but varied, exceeding proboscis

from basal two-thirds of segment 5 to base of segment 11; scape distinct, visible

under torus; torus small, with a few short hairs on anterior mesal surface and

sometimes with | to several small broad scales; flagellar segment 1 without dis-

tinct whorl of bristles and always with numerous small broad appressed scales;

segments 2-13 without scales except sometimes a few at base of segment 2, flag-

ellar whorls short, composed usually of 8 bristles, dense vestiture of short hairs;

flagellar segment 1 conspicuously elongated; segments 2-12 of varied length, either

segments 2-12 subequal in length but progressively slightly shortened distad, or

segments 2-4 markedly longer than segments 5-12 which are subequal in length

but progressively slightly shortened distad; segment 13 longer than 12. Thorax:

Mesonotum and scutellum with narrow curved decumbent dark scales, some scales

broader, particularly on lateral scutellar lobes. Acrostichal bristles absent on disc,

sometimes slightly developed on anterior promontory; dorsocentrals, supraalars and

marginal scutellars present and strongly developed; posterior fossals usually 1-3.

Postnotals present or absent. Paratergite bare. Apn with strong bristles and with

a group of weaker bristles posteriorly, usually arranged into a more or less trans-

verse row; ppn with several bristles in dorsocaudal angle and with narrow curved

decumbent scales on upper part followed sometimes by some broad scales caudo-

ventrad, lower part usually with a few hairs, scales absent; ppl with several bris-

tles and sometimes with 1 to several small translucent scales; psp bare; stp with

numerous bristles arranged in a continuous curved row along dorsal and caudal

margins and with a varied, usually extensive, patch of broad appressed translu-

cent scales, sometimes with scattered hairs in addition; pra with several bristles;

mep with or without a variable patch of broad translucent scales, with or with-

out a few to numerous microsetae in addition, Jmep with or without 1 very strong

bristle, umep with a few to numerous bristles; meron and metapleuron simple;

metameron either simple or with a few microsetae; usually a few microsetae near

base of haltere. Legs: Coxae with scales and bristles; anterior surface of forecoxa

largely covered by scales and bristles, basolateral area with a few scales; midcoxa

with scales anterolaterally and with 2 groups or rows of bristles laterally, the pos-

terior stronger, upper lateroposterior surface with or without scales; hindcoxa with

Adames: Genus Deinocerites 13

anterolateral surface with scales, lower lateral with bristles, upper lateroposterior

with or without a few scales, posterior surface with bristles. Femora subequal in

length. and not markedly different in length from proboscis, frequently with var-

iable lighter portions. Forefemur with anteroventral margin with several bristles,

a complete row of bristles or a row of short spiniforms; posterodorsal margin with

several strong bristles or a complete row of weaker bristles, latter sometimes re-

placed by spiniforms near base. Midfemur slightly swollen. Tibiae and tarsi dark

scaled. Foretibia about equal to forefemur, midfemur and hindfemur slightly long-

er, subequal. Basal tarsal segments shorter than respective tibiae. Claws simple on

all legs; empodium well developed, padlike but short and with setiform spicules;

pulvilli small and with setiform spicules. Wing: Membrane with microtrichia pres-

ent. Veins entirely dark scaled. Base of vein R with a few (remigial) bristles; cell

R, longer than its stem; vein 1A ending well distad of furcation of Cu, base with-

out scales on ventral surface; fringe normal, dark except near base; alula with long

narrow marginal scales and broader dorsal scales; upper calypter with a long mar-

ginal row of bristles. Haltere: Stem light, dorsoapical part with several dark scales;

knob dark scaled. Abdomen: Tergite I extensively scaled. Tergites completely dark

scaled. Sternites II-VII usually lighter than tergites, light scaling variable. Bucco-

pharyngeal Armature: A single row of blunt teeth on pharyngeal bar; a larger

roughly hemispherical, strongly reticulate sclerite over point of articulation of ci-

barium and pharynx.

FEMALE GENITALIA. Segment VIII partly retracted into segment VII. Ster-

nite VIII longer than tergite; with strongly sclerotized caudal band; scales present

or absent; deeply emarginate midventrally on caudal margin, side of emargination

sometimes produced into distinct angle with strongly differentiated marginal bris-

tles. Tergite IX a distinct sclerite with or without setae. Tergite X variously de-

veloped, sometimes as a broad strong sclerite at base of cercus. Cercus long, ex-

tending beyond postgenital plate; more or less conical; usually completely retract-

ed, sometimes with apex visible; with numerous setae but without scales on body,

apex with more or less strongly and variously differentiated setae. Postgenital plate

with poorly sclerotized cup-shaped basal part; distal part strongly sclerotized with

median caudal emargination or notch and with more or less strongly differentiated

lateral setose lobes; in lateral aspect lobe rounded or elongate and with variously

differentiated dorsal subapical (1) and apical (1) setae. Cowl represented by a broad

sclerotized arch articulating with tergite IX laterally, deeply emarginate on anter-

lor margin midventrally where its caudal margin is more or less distinctly joined

to base of postgenital plate; sigma poorly sclerotized, very narrow and indistinctly

joined to cowl; insula poorly sclerotized, with several microsetae; atrial plate nar-

row, strongly sclerotized (cowl, sigma, insula and atrial plate studied in cancer

only). Spermathecae 3, one larger than the others.

MALES. Essentially similar to females; labium with a more or less distinct joint

distad of middle; palpal segment 3 longer. Antenna: Distinctly longer but simi-

lar to that of female; varied in length, exceeding proboscis from apex of segment

10 to middle of flagellar segment 3; flagellar whorls essentially as in female but

_ Slightly longer, flagellar segments 1, 1 and 2, 1-3 or 1-4 with scales, segments

2-6 or 7 with variable number of shaft bristles of approximately same length as

those in whorls; flagellar segment 1 distinctly longer, the others of variable length,

either segments 2-12 subequal but progressively slightly shortened distad, or seg-

Ments 2-7 markedly elongated, but progressively shortened distad and segments

7-12 subequal but progressively shortened distad; segment 13 either shorter, equal

14 Contrib. Amer. Ent. Inst., vol. 7, no.2,1971 ~

to slightly longer, but always broader than 12. Legs: Claws of foreleg and midleg

enlarged, either similar in shape or anterior member of a pair longer, with or with-

out teeth; claws of hindleg always small and simple as in females; empodium of

legs with enlarged claws very strongly developed and with branched processes.

MALE GENITALIA. Rather uniform throughout genus; short and thick. Seg-

ment VIIT: Well developed but normally retracted about half or more of its length

into segment VII; without any special features except for broad median emargi-

nation of caudal border of tergite and development of numerous strong bristles

on and near this border, particularly on lateral tergal area. Segment IX: Strongly

developed. Tergite represented by a pair of greatly developed lateral lobes articu-

lated and movable on articulation with a lateral dorsal process of sternite; also

articulating with a dorsomesal process of sidepiece and the basal piece; median

part of tergite largely membranous; lobes variously developed and with numer-

ous setae basomesally. Sternite long and with a median longitudinal apodeme, a

caudolateral winglike lobe, and an anterolateral narrow sclerotized bar which ex-

tends dorsad on each side and articulates at its dorsal end with tergite. Sidepiece:

Conical, very thick, scales present or absent, tergal surface with variable number

of strong bristles. Subapical lobe (median mesal lobe of Belkin and Hogue, 1959)

about median in position, projecting dorsad; with 3 differentiated setae, 2 of which

are always spiniform (a more mesal, and Bb) and the other (c) bristlelike or seti-

form; usually a thumb at dorsomesal angle; always a patch of short setae on base

of lobe on dorsal surface; 3 well differentiated specialized setae on ventromesal

surface, the 3 setae usually bristlelike and attenuated apically, except in 1 case

(nicoyae) in which the 2 distal are spiniform, rather heavy and without apical

attenuation; also a variable number of small hairs on ventromesal surface of lobe.

A differentiated seta laterad of subapical lobe on tergal surface of sidepiece; a

membranous area basad of subapical lobe; usually a distinct apicosternal bristly

lobe, dorsad of which the clasper folds at rest against body of sidepiece, this area

at least partly membranous; apicosternal lobe with or without differentiated long

apical seta. Clasper: Short, irregular in shape, with a dorsal inner postmedian ang-

ular shelflike process, a ventral inner apical tooth, and a dorsal outer apical pro-

cess; inserted between the 2 apical projections is a heavy forked spiniform; dor-

sal surface with a dense vestiture of long hairlike spicules. Phallosome: Complex,

consisting of a ventral aedeagus'and a pair each of ventral and dorsal parameres.

Aedeagus more or less conical or cylindrical in tergal aspect with a pair of lateral

sclerotized plates variously developed and connected by a basal sternal sclerotized

bar and apparently by a sternal membrane not always detectable. Dorsal para-

mere (according to Belkin, 1968a:9) apparently homologous to the opisthophallus

of some Dixinae, articulated ventrally with aedeagus and ventrolaterally with ven-

tral paramere (ventral paramere articulating with aedeagus ventrally and basal piece

laterally); dorsal parameres consisting of a pair of lateral toothed plates with an

apical dorsal spine and ventrolateral teeth; the 2 sclerotized plates in tergal as-

pect are either closely approximated or widely separated and are often more or

less connected by an incomplete dorsal sclerotized bridge. Proctiger: Strongly de-

veloped, with basolateral and paraproct sclerotizations and with a slight dorso-

lateral process; without basal sternal process. Crown of paraproct with a single

row of short, heavy, blunt and curved teeth. Cercal setae numerous (details not

studied).

PUPAE. Cephalothorax: All hairs present, variously developed; hairs 2,3-C ap-

proximated or widely spaced; 5-C usually the longest hair on cephalothorax, ex-

Adames: Genus Deinocerites 15

cept in Epitedeus Group in which it is usually subequal to 8-C; hair 6-C always

shorter than 7-C; hairs 6,7-C very close together; 8,9-C well caudad of trumpet

and moderately close together. Trumpet: Not placed on distinct tubercle; inserted

about halfway between middorsal line and wing base; short to moderate in length;

tracheoid for about 0.3-0.5 of trumpet length; pinna usually short. Metanotum:

Hair 10-C variously developed, from very short and weaker than 11-C to very

strong and longer than 11-C; single to multibranched. Abdomen: Hair 1-1 with

about 4-23 primary branches, fringed, barbed or secondarily dendritic; 1-II either

very similar to or markedly different in appearance from 1-I, but always closer

to midline, with 1 to many primary branches, simple, barbed or markedly second-

darily dendritic; 2-II laterad of 3-II; hair 5-II laterad of 3-II except in Epitedeus

Group in which it is mesad, usually single (1-4); hair 7-I] ventral in position; 2-

III-VII mesad of hair 1; hairs 5-III-VI long, similarly developed, 5-III usually sin-

gle (1-3, except usually triple in mathesoni), 5-IV usually single (1 or 2), hairs

5-V,VI always single; 6-I-VII long, single, as well developed on VII as on other

segments; 9-VII short, branched and caudad of hair 6; hair 9-VIII very long and

single, arising ventrally at caudolateral angle. Terminal Segments: Hair 1-IX not

developed. Median caudal lobe well developed, in general with its posterior mar-

gin more or less rounded. Cercal lobe of female strongly developed. Male genital

lobe projecting well beyond median caudal lobe; IX tergite lobe of adult devel-

oped inside the paddle. Paddle: Midrib very strongly developed; external buttress

_ Slightly developed; margins without spicules. Hair 1-P subapical, long, subequal to

or longer than paddle; 2-P absent.

FOURTH INSTAR LARVAE. Head: Head capsule about as wide as or wider

than long, widest at level of base of antenna owing to development of conspic-

uous lateral expansion on each side caudad of antenna and the absence of a dis-

tinct ocular bulge. A broad deep pouch open cephalad within this subantennal

expansion, pouch with a membranous eversible inner wall into which projects a

conspicuous fingerlike lobe of mandible with long hairlike spicules, wall withdrawn

when lobe of mandible is abducted and expanded out and inflated when lobe

' of mandible is adducted eliminating pouch; mandible (Howard, Dyar and Knab,

1912, fig. 524) with curved fingerlike lobe densely covered with long hairlike spic-

ules arising from anterolateral basal angle and projecting into lateral subantennal

pouch, and with apical:mesal angle below teeth developed into a long curved horn;

maxilla (Howard, Dyar and Knab, 1912, fig. 479) large, normal, palpus short. La-

brum not well differentiated dorsally, rounded on anterior margin; mental plate

rudimentary, poorly sclerotized and with variously shaped marginal spicules; labial

plate very long and narrow; maxillary suture complete and extended dorsolater-

ally caudad of posterior tentorial pit; collar not developed. Hair 1-C long, thin

to moderately thick, widely spaced, arising on dorsal surface of labrum, not api-

cally; 2-C poorly to well developed, usually distinctly mesad of level, or some-

times slightly mesad, in line or laterad of 1-C; hair 3-C when present usually rep-

resented by a spicule; 4-6-C in a group caudad of level of 7-C; hair 4-C minute,

multiple; 5,6-C strongly developed; 10-C quite close to 9-C; hair 11-C dorsal, me-

sad of pouch; 12,13-C close together; 14-C removed from margin; 15-C in anter-

ior half of labial plate; 16,17-C not developed. Antenna: From about one-third

to about half as long as head capsule, unsegmented, slender, gradually tapered

apically; shaft either simple or with few to numerous spicules in basal part. Tho-

rax: Without any apparent outstanding features; hairs 1-3-P on poorly sclerotized

connected basal tubercles or with poorly developed and unsclerotized common

16 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

tubercle; 1,2-P long, single; 3-P shorter than 1,2-P, single to triple; 4-P well de-

veloped, branched; 5,6-P long, single; 7-P long, branched; 9-P long, single to trip-

le; 10-P moderate; 12-P long, single; 13-P absent; 14-P single; 1,2-M short, 3,4-M

moderate; 5-7-M long, single, 6-M heaviest; 8-M long, multiple, with heavy tub-

ercle; 9-M long, multiple; 10,12-M long, single; 14-M short, dendritic; 1-6-T mi-

nute to small; 1,2-T placed far mesad; 1,5-IT always minute; 6-T variously devel-

oped; 7-T large, multiple, with heavy basal tubercle; 9-T long, multiple; 10-T long,

single; 13-T short, multiple. Abdomen: Hairs 6-I-VI long; 7-I short to long, 7-II-

VI always short; 1-I,II distinctly mesad of others, minute; 1-III moderately to —

strongly developed; 2-III-V mesad of 1; hairs 5-I-VII small, branched; 9-II-VI mi- —

nute, single; 10-II-VI moderately to well developed, 10-II usually single (usually

triple in belkini, 1-4), hair 10-III usually single or double (1-3), hair 10-IV usually

single or double (1-3), hairs 10-V,VI usually single (1 or 2); hairs 13-III-V strong;

11,12-II-V laterad of 13. Segment VIII: Comb of numerous scales in large triang-

ular patch; individual scale long, parallel-sided, with expanded, fringed, spatulate

apex; hair 1-VIII mesad of 2; hairs 2,4-VIII strongly developed, single. Siphon:

Elongate, without marked distal tapering. Index varied from 3.2-6.5; acus pres-

ent, attached, with ventral process; valves moderate, normal; trachea well devel-

oped. Pecten not extending to middle of siphon; individual pecten tooth usually

unequally bifid or trifid. Siphonal hairs always 3 lateral pairs, 1 large (1-S) near

middle and 2 small (lad-S dorsal and lav-S ventral). Anal Segment: Saddle usu-

ally represented by poorly defined, unconnected dorsal and ventral sclerotizations;

spicules short, in a patch; acus absent. Gills a single pair, short to moderately

long, either round and slightly emarginated on apex or more or less cone-shaped

and slightly emarginated on ventral margin; without trachea. Hair 1-X small, not

on saddle; 2-X moderately long, multiple; 3-X very long, single; 4-X all on grid,

6 or 7 pairs, well developed, multiple.

EGGS. Very brief descriptions have been made for the eggs of 3 species. Fisk

(1941:547) describes the egg of mathesoni (as spanius) as “ovoid ... about 0.4 mm

in length and resembling hens’ eggs in shape’’; Haeger and Phinizee (1959:35) found

that of cancer to be “‘bullet-shaped, with the head in the blunt end. The grayish

brown chorion and exochorion are very transparent ...”’; and Galindo (1967:68)

states that the egg of pseudes is “elliptical in shape and black in color, being

very similar in structure to Culex ova’.

DISCUSSION. For discussions of the systematics, bionomics and medical and

pest importance, see the separate sections above dealing with these subjects.

The brief taxonomic history of Deinocerites has been reviewed by Belkin and

Hogue (1959:416-417) and will not be repeated here. It is presented in synoptic

form in the synonymy and taxonomic references given above for the genus.

KEYS TO GROUPS AND SPECIES

FEMALES

T. Mesepimeron with a patch of translucent scales © © 2 7... TO. 2

NiceeOUNCTON WittOUL Scales 60. we yl a ee a ee

ZT). Lower mesepimeral bristle present; antenna with only flagellar segment

l markedly longer than others (Pseudes Group)... . 0...

3(2).

4(3).

B(2).

6(5).

7(1).

8(7).

9(8).

Adames: Genus Deinocerites 17

Lower mesepimeral bristle absent; antenna with flagellar segments 1-4

markedly longer than others (Epitedeus rn et Se i

Pseudes Group

Forefemur with anteroventral and te margins with row of bris-

tles'only-7. 08 . . «18. mcedonaldi

Forefemur with atiteroventral” margin “with row of short spiniforms and

posterodorsal margin with row of short spiniforms at base followed by

OEISCLOR Liste ee is yn clear LU a malt eta a ek te a7 eae

Antenna exceeding proboscis at most from base of flagellar segment 8;

tergite IX usually without setae, rarely 1 present. . . . .16. pseudes

Antenna longer, exceeding proboscis at least from basal two-thirds of

Wee ieee 4; Psat IX with 2 or more setae (2-7) on each side

arc as! vie .17. howardi

Epitedeus Group

Forefemur with an anteroventral row of short spiniforms (Atlantic com-

pie 5 i . . . 11. epitedeus; 12. panamensis; 13. colombianus

Forefemur with an pin ferent row of bristles (Pacific complex). . .6

Cercus (in lateral aspect) with dorsal margin of sclerotized part distinctly

longer than ventral; its basal width less than 0.5 of length of sclero-

tized ventral margin; 5 or 6 apical and subapical distinctly thickened

setae with twisted apex ... . . . 14, costaricensis

Cercus (in lateral aspect) with dorsal and ventral margins of sclerotized

part subequal in length; its basal width over 0.6 of length of sclerotized

ventral margin; 5 or 6 apical or subapical setae slightly to moderately

thickened and with apex simple or slightly twisted . . . . 15. curiche

Cercus with 2 long spatulate apical or subapical specialized setae (Cancer

GEOUD i ou. oe eminem oe

Cercus without long spatulate specialized setae a ltt BN SSIK RRA 10

Cancer Group

Specialized setae of cercus not inserted side by side, one apical and short-

er, the other subapical; lateral lobe of postgenital plate elongate and

with long apical bristle. . . . 5) a daineien.« AO, panamnaees

Specialized setae of cercus inserted side by side on apex aaa subequal;

lateral lobe of postgenital lobe short and rounded, its apical bristle

S90Gs Ab icnaiee Mar aehime cceh ay was ved nid wy webct ete ea de ae

Distal part of postgenital plate (in ventral aspect) with width not more

than 1.8 of its length, usually with a distinct median caudal notch as

deep as wide and with lateral lobes well differentiated. . . .8. cancer

Distal part of postgenital plate (in ventral aspect) with width always more

than 2.0 of its length, usually with only a broad shallow emargination

and with lateral lobes appearing as small rounded protuberances .

9. melanophylum

10(7).

11(10).

12(10).

13(12).

21),

3(2).

4(2).

Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Postnotum without setae; antenna relatively short, exceeding proboscis at

most from base of flagellar segment 9; sternite VIII with distal sclero-

tized band uniformly broad and largely covered with scales (Spanius

Group). . ec ora

Postnotum usually with a | few ‘setae in lower part: antenna relatively long-

er, exceeding proboscis from base of flagellar segment 8; sternite VIII

with distal sclerotized band narrow, tapering ventrad and with only a

few Scales CDVarit WOM) 6 604. fis Ge eee ee teed we. TZ

Spanius Group

Antenna relatively short, exceeding proboscis from base of flagellar seg-

ment 11; cercus (in lateral aspect) abruptly narrowed in distal half

(comflicus complex). . . . . . «1. spanius (Pacific); 2. atlanticus

Antenna relatively longer, exceeding proboscis from apex of flagellar seg-

ment 9; cercus (in lateral aspect) evenly narrowed from base (northern

complex). . .. . . .. 3.mathesoni (Atlantic); 4. belkini (Pacific)

Dyari Group

Pleural integument almost uniformly dark brown, not markedly contrast-

ing with dark mesonotal integument; distal part of postgenital plate

(in ventral aspect) with margins of median caudal notch parallel

. 6. barretoi

Pleural integument with whitish: portions strongly contrasting with dark

mesonotal integument; distal part of postgenital plate (in ventral as-

pect) with margins of median caudal notch diverging distad . .. 13

Dersite 1% usually witli bsetain of. ov aa ico. 7. afeopae

Tersite TX usually with 4 setae. cs Sis ew ee hw Be awl

MALES

Mesepimeron without scales . . Ce re ee ee

Mesepimeron with patch of translucent scales . urea ies we gna ae a hen i

Only flagellar segment 1 markedly elongate; antenna exceeding proboscis

at most from apex of flagellar segment 9 (Spanius Group) . ... .3

Flagellar segments 1-5 or 1-6 markedly longer than others; antenna ex-

ceeding proboscis from at least middle of flagellarsegment5 . . . .4

Spanius Group

Antenna relatively short, exceeding proboscis from base of flagellar seg-

ment 11 (southern complex) . . . .1.spanius (Pacific); 2. atlanticus

Antenna relatively longer, exceeding proboscis from apex of flagellar seg-

ment 9 (northern complex). ae at Sa Ets case an ic a nee RE HL

3. mathesoni (Atlantic); 4. belkini (Pacific)

Postnotum usually with a few bristles toward lower part; Pacific distribu-

tion (Dyari Group) . . es

Postnotum without bristles: Atlantic distribution (Cancer Group) . ee |

5(4).

6(5).

7(4).

8(7).

9(1).

10(9).

11(10).

12(9).

13(12).

Adames: Genus Deinocerites 19

Dyari Group

Pleuron almost uniformly dark brown, not contrasting markedly with dark

brown mesonotum; flagellar segments 1-5 markedly elongated

. 6. barretoi

Pleuton with whitish portions eae contrasting with dark brown meso-

notum; flagellar segments 1-6 markedly elongated . . ..... .6

Anterior claw of foreleg with a long, very slender tooth; anterior claw

of midleg with a shorter, heavier tooth . . a. Sayan

Anterior claws of foreleg and midleg both with a heavy tooth . 7. nicoyae

Cancer Group

Pleuron usually whitish and distinctly contrasting with dark brown meso-

notum; tooth of foreclaw and midclaw nearly basal; antenna exceed-

ing proboscis from base of flagellarsegment5. . . . . . 10. magnus

Pleuron darker, only slightly or not at all contrasting with dark brown

mesonotum; tooth of foreclaw and midclaw distinctly subbasal; anten-

na exceeding proboscis from distal third of flagellarsegment4 . . .8

Anterior claws of foreleg and midleg dissimilar, that of foreleg with a

shorter subbasal tooth; flagellar segment 13 slightly longer than 12

9. melanophylum

Anterior claws of foreleg and midleg similar, with large subbasal tooth;

flagellar segment 13 subequalto12...... ... .. .. .8.cancer

Lower mesepimeron without differentiated bristle (Epitedeus Group). 10

Lower mesepimeron with | very strong bristle (Pseudes Group). . . 12

Epitedeus Group

Forefemur with anteroventral row of bristles (Pacific complex) .

. . 14. costaricensis; 15. cache

Forefemiut with aiteroventeal’ row of short spiniforms (Atlantic complex)

Anterior claw of midleg simple .. . . . 13. colombianus

Anterior claw of midleg with a short tooth . ve epitedeus: 12. panamensis

Pseudes Group

Forefemur with anteroventral and 4 git deer rows of bristles only. . .

. 18. mcdonaldi

Forefemur with anteroventral row ‘of short spiniforms and with bristles

of posterodorsal row replaced by short spiniforms near base . . . 13

Antenna exceeding proboscis from at least distal third of flagellar seg-

ment 3; claws of foreleg very slender and with a distinct very slender

subbasaltooth. .. . . 17. howardi

Antenna exceeding proboscis ‘at most from. basal third of segment 4;

claws of foreleg very heavy and usually with a minute submedian pro-

SOO ey ke gt ee ae pepe a es Gul cs: i ee a

2(1).

3(2).

4(2).

5(1).

6(5).

7(6).

Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

MALE GENITALIA

Sidepiece with numerous scales; apicosternal lobe of sidepiece short, more

or less rounded and without a long heavy, differentiated bristle, a short

subapical bristle may be present (Spanius Group) . . e2

Sidepiece usually without scales, at most with 1 or 2 scales; ‘apicosternal

lobe of sidepiece prominent and with a long heavy, differentiated bristle

Spanius Group

Dorsal paramere in form of a small compact, uniformly and heavily sclero-

tized plate, its apical spine short to moderately long, slightly curved

and gradually pointed (southern complex) . . . a:

Dorsal paramere with 2 sclerotizations, tergal and sternal, connected by

a narrow bridge very poorly sclerotized on the margins and membran-

ous centrally, tergal sclerotization bearing the long, slender, strongly

hooked apical spine (northern complex)... .. <0... +42. 2 + si 4

Dorsal paramere with apical spine moderately long and curved, ventral

teeth represented by 2 or 3 strong, moderately long spines in caudal

area; ninth tergite lobe with outer margin more or less straight except

for a bent and angled apex (Pacific). . . . . L. spanius

Dorsal paramere with apical spine short and slightly curved, ventral teeth

represented by 3 or 4 small denticles at caudosternal angle; ninth ter-

gite lobe with outer margin curved and directed mesad, apex rounded

(A aRUG) ree ee ee et ce ees ee OE ES

At least 1 of the dorsal parameres with 2 teeth (2 or 3) on sternal sclero-

tization; dorsal paramere wider than long (Atlantic) . . 3. mathesoni

Dorsal parameres always with 3 or 4 teeth on sternal sclerotization; dor-

sal paramere as wide aslong (Pacific) . . ..... .. . 4. belkini

Ninth tergite lobe strongly flattened, not markedly angled laterad at base

and dorsal in position (Cancer Group). . . : .6

Ninth tergite lobe cylindrical, markedly angled laterad at base and 1 more

PARSE DOS OR i a ks A cs Puedes oe

Cancer Group

Ninth tergite lobe short, not reaching base of subapical lobe of sidepiece;

apical spine of dorsal paramere thick, not strongly differentiated from

body of paramere. . . . . 10. magnus

Ninth tergite lobe long, oe distad of base of subapical lobe of

sidepiece; apical spine of dorsal paramere slender, strongly different-

iated from body of paramere 1 ae cee

Dorsal paramere with ventral teeth outstanding, arising from distinct dis-

tal process or lobe; its dorsal bridge broad and short; its apical spine

rather short and only slightly curved ... . . . 8. cancer

Dorsal paramere with ventral teeth appressed; its dorsal bridge narrow

and long; its apical spine long and strongly curved . 9.melanophylum

8(5).

9(8).

10(9).

11(8).

P2ChI):

aCE2).

14(13).

15(14).

Adames: Genus Deinocerites ZI

Dyari, Epitedeus and Pseudes Groups

Ninth tergite lobe short, not reaching base of subapical lobe of sidepiece;

its distal part more or less conical . . i

Ninth tergite lobe longer, reaching at least base of aabapica! lobe of side-

piece, distal part variously shaped but not conical . . .... . ‘wi

Apical spine of dorsal paramere heavy and strongly curved, in situ reach-

ing or crossing its mate; ninth tergite lobe with distal part distinctly

attenuated or more or less pointed; apex of aedeagus rounded. 5. dyari

Apical spine of dorsal paramere slender and not strongly curved, in situ

widely separated from its mate; ninth tergite lobe with distal part not

strongly attenuated, blunt; apex of aedeagus emarginate or angled . 10

Dorsal paramere with ventral teeth usually about 18, short to moderately

long, and arising from an evenly convex ventrolateral border .

14. costaricensis

Dorsal pare oa witly about 25 lone: moderately heavy ventral

teeth arising from a distinct humplike expansion of ventrolateral border

| . 15. curiche

Dorsal paramere with a strong dentate process from ventrolateral border,

in situ extending almost as far caudad as apical spine . . . 16. pseudes

Dorsal paramere with ventral teeth arising from convex ventrolateral bor-

der, Ot Troma isting! BrGceGs 6 i a es a

Ninth tergite lobe extending well beyond base of subapical lobe, strongly

bent near the middle and with this slender distal part more or less

paralleling sidepiece. . . . 12. panamensis

Ninth tergite lobe not extending beyond base of subapical lobe’. . 3. 43

Ventromesal surface of subapical lobe with the 2 distal setae rather heavy

and spiniform and without apical attenuation; dorsal paramere with

- distal part expanded laterad as a broad hemispherical ledge. 7. nicoyae

Ventromesal surface of subapical lobe with all 3 setae bristlelike and at-

tenuated apically; dorsal paramere with its distal part not expanded

as a broad hemmppnercal ietee Jes ae re a eI

Dorsal paramere with apical spine long and slender, but not touching its

mate in situ; at least the more distal ventral teeth of dorsal paramere

as seen in situ long and conspicuously oriented in the same direction

asthe apical spine’ . =. . On og) eae

Dorsal paramere with apical spine moderately long to long and heavy, us-

usally intercrossing with its mate in situ; ventral teeth of dorsal para-

mere as seen in situ short and not oriented in the direction of the api-

CALGON ia 6 oe a ee ee ee

Ninth tergite lobe with distal part broad and directed laterad; body of

lobe broad in basal two-thirds, cae attenuated in distal third

Se Oe a ee Ce ee . 13. colombianus

ae Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Ninth tergite lobe with distal part slender and directed mesad by distinct

curvatures of inner and outer margins; body of lobe broad basally,

constricted near middle and slender in distal half . . . 11. epitedeus

16(14). Ninth tergite lobe with apex simple; aedeagus in tergal aspect almost uni-

form in width except for a subbasal expansion, distally the 2 plates

come together forming a rounded expansion; seta c of subapical lobe

of sidepiece rather heavy and bent apically. . . . . . . 6. barretoi

Ninth tergite lobe slightly hooked apically; aedeagus in tergal aspect var-

iable in width because of highly sinuous outer margins, with a sub-

apical constriction or sinuosity and narrowed and slightly emarginate

apex; seta c of subapical lobe of sidepiece moderately heavy, attenu-

ated but not bent apically . . . . . . .18.mecdonaldi; 17. howardi

PUPAE

(1. spanius and 6. barretoi unknown)

Be Hair 1-II with 1-3 simple or barbed branches (either subequal to or longer

than tergite III) .. ae

Hair 1-Il multibranched* or, if with less than 4 primary branches, ‘then

markedly secondarily dendritic and 10-C eee multibranched (us-

ually 1-II shorter than tergite III) ...... eae.

2(1). Hair 10-C strongly developed, distinctly longer than 11-C; hair 5-II al-

ways mesad and cephalad of 3-II; hair 5-C single, moderately long, at

most equal in length to distance from its alveolus to base of tiumpet

and to length of trumpet (Epitedeus Group) . . A.

Hair 10-C poorly developed, always shorter than 11-C (0. Be 0, 5 of 11-C);

hair 5-II always laterad of 3-II; hair 5-C single or double, strongly de-

veloped, longer than distance from its alveolus to base of trumpet, at

least 1.5 of trumpet length (Dyari Group)

Epitedeus Group

3(2). Hair 10-C usually double; 1-III-VI branched soem . 13. colombianus

Hair 10-C usually single; 1-II]-VI forked . . . OS ee 2 ae

4(3). Hair 7+C-sinele (Atlantic): coe G) i os vera smokes web epitedens

Hae aialiy Memeo SS SS OE ee

5(4). Hair 3-V usually double (Atlantic) . . . ... . . . 12. panamensis

Hag 3-V usally eines (aciich oe Pees ees ee

6(5). Hair 5-l with 4-6 branches . ........... . . 14. coStaricensis

Pais 5-1 double or triple oo Oe Sai sw a. eee 1S, eurtete

Dyari Group

FEZ). Hair 5-C single; 1-II single . . Dik hos eee

Hair 5-C double; 1-II triple, the middle branch longest . . . 7, nicoyae

8(1).

9(8).

10(9).

11(8).

P20 11).

13(12).

14(11).

15(14).

2(1).

Adames: Genus Deinocerites 23

Hair 1-VII longer than tergite VIII, at least extending to middle of 4-VIII;

hair 7-C usually 4 or 5 branched (Spanius Group) =. He

Hair 1-VII shorter than or sometimes equal to tergite VII, never exceed-

ing the basal third of 4-VIII; hair 7-C usually double or triple . tae 2

Spanius Group

Hair 6-C single; 4-VI apr double or triple; 10-C ls 3 or 4 branched

. . 2. atlanticus

Hair 6C usually ‘double. or triple; AVI usually 4 4 or 5 branched; 10-C

single ordouble ... . Be etc gs ae

Hair 8-C single; 10-C usually single; 3-VI single or double ha Caan :

a ies belkini

Hair 8-C double < or triple 10-C double; 3-VI with 3 or 4 branches (At-

Pate) AS ee SOE OSA a ie enon

Hair 8-C usually 2-4 branched; 3-V usually double (Cancer Group). . 12

Hair 8-C eines se 3-V more often can than double (Pseudes Group)

Cancer Group

Tergal area between hairs 41-I darkly pigmented . . . 9.melanophylum

Tergal atea between fais I-) amber in color: 0. er eee OTS

Hair 3-II usually single; 7-C usually double; 10-C most often double

. 8. cancer

Hair 3 I usually double c or tiple; 1 usually 4. or 4 branched; 10-C usu-

ally 3-5 branched. 3. Ce als 5 IO. magnus

Pseudes Group

Hair 10-C distinctly longer than 11-C, markedly multibranched, resem-

bling float hair 1-I; hair 5-VII usually reaching to about middle of ter-

gite VIII; hair 4-C most often 4-6 branched. . . .. . . 16. pseudes

Hair 10-C subequal to 11-C, usually 2-4 branched, not resembling float

hair 1-1; hair 5-VII not exceeding apex of His he VII; hair 4-C most

often double Or tipie Ye vii i a Sue he aa ae aa

Hair 1-V usually double or triple; 8-III usually 4-6 branched

pues: mcdonaldi

Hair 1-V usually 4. 6 branched; 8 I] usually double « or triple .17. howardi

FOURTH INSTAR LARVAE

(1. spanius and 6. barretoi unknown)

Hair 6-II single; 1-S usually 4-6 branched (Spanius EMP) nea omer et

Hair 6-II double; 1-S usually double or triple . .. . a

Spanius Group

Hair 6-C single or double; 6-T usuaily single . . . 2. atlanticus

Hair 6-C usually 3 or 4 branched; 6-T always 2-4 branched pr Ae a

3(2).

4(1).

5(4).

6(4).

7(6).

8(7).

9(8).

10(8).

11(6).

12(11).

Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Siphon short, index less than 4.3; at least 1 of the branches of siphonal

hair 1-S-reaching alveolus of lay-S: small ventral saddle absent; 3-IV

with 2-4 branches (Atlantic) .. . . . 3. mathesoni

Siphon longer, index more than 5.0; none > of the branches of siphonal

hair 1-S reaching alveolus of lav-S; small ventral saddle present; 3-IV

With 5-7 branehes (Pacific). 2 os i ee 5 & Del

Hair 1-VII poorly developed, shorter than or subequal to 3-VII, not reach-

ing the base of segment VIII; hair 6-VI usually double; 2-C either

slightly mesad, in line with or laterad of level of 1-C (Dyari Group). .5

Hair 1-VII moderately to strongly developed, longer than 3-VII, always

exceeding base of segment VIII and sometimes the base of siphon; 6-

VI single; 2-C always markedly mesad of 1-C .

Dyari Group

Ventral brush (4-X) with 7 pairs of hairs; 6-III with 3 or 4 branches; 6-IV

usually triple . . . . 9. dyari

Ventral brush ie with 6 pains a oe S iI oble: & IV usually double

: ee 7. nicoyae

Hair 9-P double or triple; 2-C inconspicuous, about 0.25-0.33 of 1-C;

hair 5-C usually double or triple (Epitedeus Group) . . Sa

Hair 9-P single; 2-C at least 0.5 of 1-C; hair 5-C usually 4 Boncied |

Epitedeus Group

Hair 1-VII double; 1-V double; 13-V with 4 or 5 branches (Atlantic)

. 13. colombianus

Hair 1-VII single : 1-V single; 13-V single to triple ee es Se ee

Ventral brush (4-4) with © baics OF Mats... sk i ee a

Ventralbrish (4-X).with 7.p4ifs Of hairs. gi, ea 2

Hair 8-P double; 3-IV with 3-5 branches; 2-T ‘bees 4 branched (Atlantic)

oe er er . 12. panamensis

Hair 8-P single; 3-I[V double; 27 T usually d double or * triple (Paeitic). . c-s

re Aig ; . 45. curiche

Hair 7-II usually single to triple; ventral margin of dorsal saddle reaching

alveolus of 1-X (Atlantic). . . . . 11. epitedeus

Hair 7-I] with 4-6 branches; ventral margin of dorsal saddle not reaching

alveolus of 1-X (Pacific) . . .. . ...5. . . . « 14. costaricensis

Pseudes Group

Hiedd Baro. oul Or tipe 2 2. Se Sees xe Shs ke

bead fait O aiwaye sure ee ee el ap ey Ie

Ventral brush (4-X) with 6 pairs of hairs (some peripheral northern and

southern populations) . . . aes Ua Oe owes (in small part)

Ventral brush (4-X) with 7 pairs of hairs . Ue Ras ek gd pees

Adames: Genus Deinocerites 25

13(12). At least 1 of the branches of hair 1-S usually reaching alveolus of lav-S;

hair 3-VIII usually 5 or 6 branched; 1-VII often double or triple; 6-C

usually double, very rarely triple on one side . . . .18. mcdonaldi

Usually none of the branches of hair 1-S reaching alveolus of lav-S; hair

3-VIII usually 3 or 4 branched; 1-VIl one pees 6-C often triple

on at Ieast-one Bide, ue 257k . . .17. howardi

14(11). Hair 1-VIII usually 5-7 branched; 1-VII usually long, seni reaching

the base of siphon (central populations) . . 16. pseudes (in large part)

Cancer Group

Hair 1-VIII usually 3 or 4 branched; 1-VII shorter, never reaching base

of sipliom sine piie: bach estind. eee ee een oe

15(14). Ventral brush (4-X) with 7 pairs of hairs; 2-C usually longer than 1-C;

hair 8-P usually double. . . . . . 10. magnus

Ventral brush (4-X) with 6 pairs me nates 2 cnt deci than 1-C;

hair SP usually single 6s, Se Ree ee ae tice 2 a6

16(15). Sum total of branches of hairs 3,4-P on both sides usually equal to or less

than 10 or 4-P usually triple and 3-P usually single . . . . .8. cancer

Sum total of branches of hairs 3,4-P on both sides usually more than 10

or 4-P usually 4 branched and 3-P usually double. . 9. melanophylum

SPANIUS GROUP

FEMALES. Small species, wing 2.5-2.7 mm. Mesonotum dark, always strongly

contrasting with very distinct whitish portions of pleural integument. Antenna:

Flagellar segment 1 with scales, shorter than combined length of segments 2 and

3; segments 2-12 subequal in length but progressively slightly shortened distad;

relatively short, at most exceeding proboscis from apex of flagellar segment 9.

Thorax: Postnotum without setae. Apn usually whitish; ppn usually whitish, scales

all narrow; ppl whitish; psp, ssp, stp and pra usually slightly darker than rest of

pleuron; paratergite whitish or very light brown; mep whitish and without trans-

lucent scales, lmep with 1 very strong bristle; meron, metameron and metapleuron

whitish. Legs: Coxal integument whitish. All of posteroventral surface of forefe-

mur and midfemur and at least basal two-thirds of hindfemur whitish; anteroven-

tral margin of forefemur without spiniforms, but with 4-6 bristles in distal two-

thirds, posterodorsal margin with about 6-9 bristles. Abdomen: Sternites II-VI and

usually part of VII with pale scales.

FEMALE GENITALIA. Sternite VIII with distal sclerotized band broad and

largely covered by scales; caudal margin not produced into a submedian ventral

angle, this area without strongly differentiated marginal bristles. Tergite IX usu-

ally with at least 1 small seta on each side. Tergite X poorly differentiated dor-

sally and with a very small indistinct lateral sclerotization. Cercus with a few mod-

erately strong apical or subapical normal bristles.

MALES. Extremely similar to females. Antenna: Exceeding proboscis at most

from apex of flagellar segment 9; flagellar segments essentially as in the females,

only slightly longer. Legs: Claws of foreleg and midleg enlarged, anterior member

of a pair always larger and with a heavy subbasal tooth, posterior simple.

26 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

MALE GENITALIA. Segment IX: Tergite lobe cylindrical; not markedly angled

at base; basal one-third broad, distal two-thirds slender; directed inwards distally

by a subapical curvature; apex exceeding the subapical lobe. Sidepiece: Scales num-

erous. Apicosternal lobe short, more or less rounded, without differentiated long

apical seta (short and subapical when present). Phallosome: Dorsal parameres wide-

ly separated in tergal aspect, without a differentiated sclerotized dorsal bridge;

articulated basoventrally directly with dorsal basal part of aedeagus.

PUPAE. Cephalothorax: Hairs 2,3-C closely approximated, 2-C poorly develop-

ed, always weaker and shorter than 3-C; hair 5-C usually double (2 or 3), strongly

developed, longer than distance from its alveolus to base of trumpet, 1.35-1.50

of trumpet length; 7-C usually with 4 or 5 branches (2-6); hair 8-C usually single

or double (1-3). Metanotum: Hair 10-C moderately to strongly developed, sub-

equal to or longer than 11-C, branching varied (1-6). Abdomen: Hair 1-II not reach-

ing apex of tergite III, multibranched (4-13), usually dendritic; 1-III-VI branched;

5-II always cephalad and sometimes slightly mesad of 3-II; hair 3-V usually double

or triple (2-4); hair 1-VII longer than tergite VIII, at least reaching middle of 4-

VIII. Paddle: Hair 1-P subequal to or slightly longer than paddle.

FOURTH INSTAR LARVAE. Head: Hair 2-C about 0.5 of 1-C and always dis-

tinctly mesad of 1-C; hair 5-C with 4 or more barbed branches (4-7); hair 6-C

variously branched (1-5), barbed, either equal to or slightly longer than 5-C. Tho-

rax: Prothoracic hair 9-P single. Abdomen: Hair 6-II single; dorsal sensillum of

segment V laterad of 4-V; hair 6-VI single; 1-VII moderately developed, longer

than 3-VII, exceeding base of segment VIII but not reaching base of siphon; 1-

VIII usually with 3 or 4 branches (2-4). Siphon: Hair 1-S usually with 4-6 branches

(3-7). Anal Segment: Ventral brush (4-X) with 6 pairs of hairs. Gill slightly emar-

ginate on apex; short, about one-half to two-thirds of dorsal saddle length. Dorsal

saddle narrow, its ventral margin far from hair 1-X.

DISCUSSION. This is undoubtedly the most primitive group of the genus as

it shows the least number of derived features and resembles more than any other

sroup the other 2 genera of the tribe, Culex and Galindomyia. The adults have

the shortest antennae in Deinocerites. In the male genitalia, the dorsal paramere

articulates directly with the aedeagus whereas in other groups this articulation is

with a process from the aedeagus. The genitalia of both sexes (sternite VIII in

female and sidepiece in male) are densely covered with scales in the Spanius Group.

These scales are lacking in the males of the other groups and are reduced to only

a few or are completely absent in their females. In the larvae, head hairs 5 and 6

are relatively short, subequal and usually branched while in the other groups 6-C

tends to become elongated and single.

The group has a unique distribution pattern and species composition with wide-

ly disjunct northern and southern complexes, each with 2 geminate Atlantic and

Pacific species. The northern complex has been reported only from the states of

Nayarit and Jalisco on the west coast of Mexico (belkini) and from southern Texas

and possibly down to Veracruz, Mexico (mathesoni). The southern complex is —

known only from Aguadulce (Cocle) to Santa Fe (Darien) on the Pacific coast of

Panama (spanius) and from a wider area on the Atlantic slope from the Canal

Zone, Colombia and Venezuela (atlanticus). It is possible that atlanticus may con-

sist of 2 or more species.

The 2 complexes are rather strongly differentiated in the structure of the dor-

sal paramere of the male genitalia, this being composed of 2 separate sclerotiza-

tions connected by a poorly sclerotized bridge in the northern complex, while in

Adames: Genus Deinocerites 27

the southern complex it is formed as a compact uniformly sclerotized plate. I

consider that the former condition is the primitive one. There are also good cor-

related morphological differences in the genitalia of the females, antennae of both

sexes, larvae and pupae as indicated in the Keys.

Although the adults of a geminate pair cannot be separated on external morph-

ological features there are good male genitalic characters to distinguish them in

both complexes. In the case of the northern complex there are also reliable fea-

tures distinguishing belkini from mathesoni in the pupal and larval stages. Since

the immature stages of spanius are still unknown it cannot be determined if a

similar situation prevails in the southern complex.

The species of the group appear to be rare or at least are seldom collected

as adults. The latter may be partially due to the superficial resemblance of both

males and females to the females of the numerous unornamented species of the

of the subgenus Melanoconion of Culex that at present are impossible to identify

to species.

Little is known about the blood feeding habits of members of the group but

apparently atlanticus may attack man in Panama. The immature stages of the Span-

ius Group have been collected only a few times, most frequently in small burrows

made by fiddler crabs of the genus Uca and occasionally in large holes of Gecar-

cinus lateralis.

1. Deinocerites spanius (Dyar & Knab)

Figs. 3,8

1909. Dinanamesus spanius Dyar and Knab, 1909:259. TYPE: Lectotype female, Corozal, Pan-

ama, Canal Zone, 11 Dec 1907, crabhole, A.H. Jennings, 69 [USNM, 12052; designation

of Stone and Knight, 1957:197].

Deinocerites spanius of Stone and Knight (1957:197).

Deinocerites spanius in part of Dyar (1925:154-155); Carpenter and LaCasse (1955:327-329);

Belkin and Hogue (1959:427-428); Stone, Knight and Starcke (1959:285); Ross (1964:104).

Dinanamesus spanius in part of Howard, Dyar and Knab (1915:213-215).

FEMALE (fig. 8). Wing 2.70 mm. Proboscis 1.45 mm. Forefemur 1.27 mm.

Abdomen about 2.65 mm. Apparently indistinguishable from atlanticus.

FEMALE GENITALIA (fig. 8). Tergite IX with or without 1 or 2 setae on

each side. Cercus (in lateral aspect) very similar to that of atlanticus except for

somewhat longer broadened basal part. Distal part of postgenital plate (in ventral

aspect) deeply notched; lateral lobe elongate (in lateral aspect), its apical bristle

convergent distally with its mate (in ventral aspect).

MALE (fig. 8). Wing 2.60 mm. Proboscis 1.58 mm. Forefemur 1.30 mm. Ab-

domen (not including genitalia) about 1.78. Essentially similar to female.

MALE GENITALIA (fig. 8). Segment IX: Tergite lobe slender; middle part al-

most uniform in width, constricted subapically, then strongly expanded and with

angled borders apically; outer margin more or less straight except for the very

distal part which is bent mesad; ventral surface with a distinct premedian pro-

jection or expansion, visible in toto only in lateral aspect. Sidepiece: Similar to

atlanticus. Phallosome: Dorsal paramere a compact uniformly sclerotized plate; api-

cal spine moderately long, curved and gradually pointed, ventral teeth represented

28 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

by 2 or 3 strong, moderately long spines in caudal area. Aedeagus in general simi-

lar to atlanticus.

PUPA and LARVA. Unknown.

SYSTEMATICS. This species is indistinguishable from atlanticus in external fea-

tures of the adults but the male genitalia readily separate these 2 geminate species.

Deinocerites spanius, the Pacific representative of the southern complex of the

group, is the rarest of the 4 species and is still unknown in the immature stages.

It is unfortunate that Stone and Knight (1957:197) in designating the lectotype

of spanius chose the female syntype from ,Corozal, thus applying this name to

the Pacific population in the Canal Zone and Panama. The much more common

Atlantic population in this area is now provisionally assigned to atlanticus but

this population too is still unknown in the immature stages.

BIONOMICS. The lectotype was said to have been reared from a larva collected

in a crabhole. I have no information on the recent collections of this species other

than the localities as noted below under distribution. Deinocerites spanius appears

to be much less common than its sibling of the Atlantic coast, atlanticus.

DISTRIBUTION (fig. 3). Pacific coast of Panama from Cocle Province to Dar-

ien Province. Material examined: 14 specimens; 5 males, 9 females.

PANAMA AND CANAL ZONE. Canal Zone: Corozal, A.H. Jennings. Ft. Clayton, C.S. Lud-

low [USNM]. Cocle: Aguadulce, GML [UCLA]. Darien: Santa Fe, GML [UCLA]. Panama:

Juan Diaz, GML. Nueva Gorgona, GML [UCLA].

2. Deinocerites atlanticus Adames, n.sp.

Figs. 3,9-11

TYPES: Holotype male with associated pupal skin and genitalia slide (VZ 170-107), La Boca,

Ocumare de la Costa, Aragua, Venezuela, 10 July 1969, T. Zavortink, J. Valencia and J. Pulido

[USNM]. Allotype female with associated larval and pupal skins (VZ 170-30), same data as

holotype [USNM]. Paratypes: 9 lpM (VZ 170-11-14,17,19,29,36,44), 6 IpF (170-16,18,37,40,

45,46), 3 pM (170-108-110), 4 pF (170-100,102,104,114), 2 IP (170-10,15), 2 lp (170-32,38),

1 1 (170-28), 14 M, 3 F, 38 L (170), same data as holotype [BM, UCLA, USNM].

Deinocerites spanius of Dyar (1923a:179; 1928:265,537); Bonne and Bonne-Wepster (1925:176);

Edwards (1932:222); Lane (1953:559); Horsfall (1955:599); Forattini (1965:108); Sutil and

Pulido (1969:119-125); Barreto and Lee (1969:435).

Deinocerites spanius in part of Dyar (1925:154-155); Belkin and Hogue (1959:427-428); Stone,

Knight and Starcke (1959:285); Ross (1964:104).

Dinanamesus spanius in part of Howard, Dyar and Knab (1915:213-215).

Culex (Deinocerites) spanius of Dyar (1918:101).

FEMALE (fig. 9). Wing 2.66 mm. Proboscis 1.73 mm. Forefemur 1.48 mm.

Abdomen about 2.71 mm. Head: Narrow decumbent scales of vertex creamy; erect

scales yellowish or pale brown; lateral patch of broad decumbent scales whitish.

Antenna: Torus occasionally with 1 scale; proboscis reaching apex of flagellar seg-

ment 10 or base of 11. Thorax: Middle and lower areas of ppn usually with 1

or more small bristles; ppl with a few bristles and sometimes a few scales; mep

with a few scales in upper part, microsetae absent; metameron occasionally with

1 or 2 microsetae. Legs: Anterior surface of forecoxa covered with bristles and

translucent scales, basolateral area with a few bristles; midcoxa with scales antero-

laterally and with 2 rows of bristles laterally, the posterior bristles stronger, pos-

Adames: Genus Deinocerites 29

terolateral surface simple; hindcoxa with scales and several bristles laterally, pos-

terior surface with bristles. |

FEMALE GENITALIA (fig. 9). Tergite IX usually with at least 1 seta on each

side, usually 1(0-2) in Venezuelan population and 4(2-5) in Panamanian. Cercus

(in lateral aspect) broadened in basal half, abruptly narrowed in distal; sternal mar-

gin more or less convex, tergal margin distinctly concave in distal half; apex di-

rected dorsad, usually with 4-6 apical and subapical bristles. Distal part of post-

genital plate (in ventral aspect) usually deeply notched except in some Panaman-

ian specimens; lateral lobe usually elongate (in lateral aspect) except in above men-

tioned Panamanian specimens in which it is rounded, its apical bristle convergent

distally with its mate (in ventral aspect).

MALE (fig. 9). Wing 2.62 mm. Proboscis 1.86 mm. Forefemur 1.56 mm. Ab-

domen (not including genitalia) about 2.37 mm. Essentially similar to female.

MALE GENITALIA (fig. 10). Segment IX: Tergite lobe very slender, slightly

widened near middle, then constricted subapically and slightly expanded into a

rounded. apex; outer margin directed mesad by a well-defined distal curvature;

ventral surface with a premedian projection similar to that of spanius. Sidepiece:

Subapical lobe with thumb; seta c a moderately heavy bristle attenuated apically.

Phallosome: Dorsal paramere a small compact uniformly sclerotized plate; apical

spine short, broad at base, slightly curved and gradually pointed; ventral teeth

represented by 2-4 small denticles at caudoventral angle. Aedeagus (in tergal view)

broad at base, then constricted, lateral plates parallel and approaching each other

distally.

PUPA (fig. 10). Abdomen 2.60 mm. Trumpet 0.58 mm. Paddle 0.66 mm. Di-

agnostic characters as in the key; general chaetotaxy based on 6 reared specimens.

Hair 10-C about equal in length to 11-C; hair 1-VII extending to about middle

of 4-VIII. Cephalothorax: Integument light brown. Hair 1(2,2-3), 2(3,3-4), 3(2,2-3),

4(4,3-5), 5(2,2-3), 6(1), 7(4,3-5), 8(2,2-3), 9(2,1-3), 10(4,3-2), 11(2,2-3), 12(3,2-

5). Trumpet: Short; slightly broadening apically; index about 4.92-6.60. Integu-

ment brown distad, blackish on tracheoid; moderately contrasting with cephalo-

thoracic integument. Tracheoid about 0.4 of trumpet length; border of pinna with-

out distinct emargination. Abdomen: Integument light brown. Segment I: hair 1

(11,8-13 primary branches), 2(2), 3(2,1-3), 4(5,4-6), 5(3,3-5), 6(1), 7(2,1-4), 9(1).

Segment II: hair 0(1), 1(7,6-9), 2(1), 3(2,1-3), 4(4,2-4), 5(1,1-2), 6(1), 7(2,1-3),

9(1). Segment III: hair 0(1), 1(4,2-4), 2(1), 3(2,1-3), 4(2,1-2), 5(1), 6(1), 7(2,1-

4), 8(4,3-6), 9(1), 10(2,1-2), 11(1), 14(1). Segment IV: hair 0(1), 1(2,1-3), 2(1),

3(3,2-3), 4(1), 5(1), 6(1), 7(2,1-2), 8(2,2-3), 9(1), 10(2,1-3), 11€1), 14(1). Seg-

ment V: hair 0(1), 1(2,1-3), 2(1), 3(2), 4(3,2-4), 5(1), 6(1), 7(4,3-4), 8(2), 9(1),

10(1), 11(€1), 14(€1). Segment VI: hair 0(1), 1(2,1-2), 2(1), 3(2,1-2), 4(2,2-3), 5(1),

6(1), 7(1,1-2), 8(2,2-4), 9(1), 10(2,1-2), 11(1,1-2), 14(1). Segment VII: hair 0(1),

1(1), 2(1), 3(2,1-2), 4(2,1-2), 5(1,1-2), 6(1), 7(1,1-2), 8(2,2-3), 9(4,2-5), 10(1,1-

2), 11(1,1-2), 14(1). Segment VIII: hair 0(1), 4(2,1-2), 9(1,1-2), 14(1). Paddle:

Width about 0.75 of length.

FOURTH INSTAR LARVA (fig. 11). Head 0.95 mm. Siphon 0.83 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 6 reared specimens.

Head: Integument yellowish. Mental plate slightly wider than long, triangular, with

a median terminal spicule, basal and lateral spicules with denticulate apex, termi-

nal simple. Hair 0(1), 1(1), 2(1), 3(sometimes developed as a minute spicule), 4(4,

3-6), 5(4,4-5), 6(1-2), 7(11,9-11), 8(2,2-3), 9(4,3-5), 10(2,2-3), 11(2,2-6), 12(2),

13(3,2-4), 14(1), 15(2). Antenna: Length about 0.38 of head; shaft simple, oc-

30 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

casionally with 1 spicule on proximal part; hair 1(3,3-4). Thorax: Prothorax: hair

0(4,3-6), 1(1), 2(1), 3(1,1-2), 4(3,2-4), 5(1), 6(1), 7(3,2-3), 8(2,1-4), 9(1), 10(1,

1-2), 11(2,1-3), 12(1), 14(1). Mesothorax: hair 1(1), 2(1,1-2), 3(1), 4(2,2-3), 5(1),

6(1), 7(1), 8(6,6-8), 9(8,7-8), 10(1), 11(€2,1-3), 12(1), 13(6,5-8), 14(5-6). Meta-

thorax: hair 1(1), 2(3,2-4), 3(4,4-5), 4(2,1-2), 5(1), 6(1,1-2), 7(8,7-9), 8(4,4-6),

9(8,6-9), 10(1), 11(2,1-2), 12(1), 13(4,3-4). Abdomen: Segment I: hair 1(1), 2(1),

3(2,2-3), 4(8,6-8), 5(2,2-4), 6(2), F2,143), 9(2,1-3), T0(3 32-3), 11(2), 12(2,2-3),

13(1). Segment II: hair 0(1), 1(1), 2(1), 3(2,2-3), 4(4,3-6), 5(2,1-2), 6(1), 7(3,2-

4), 8(2,1-2), 9(1), 10(1), 11(2,2-3), 12(2,2-3), 13(2). Segment III: hair O(1), 1

(2,1-2), 2(1), 3(2,2-3), 4(1), 5(1), 6(2), 7(5,4-6), 8(1,1-2), 9(1), 10(2,1-2), 111,

1-2), 12(1), 13(1,1-3), 14(1). Segment IV: hair 0(1), 1(1,1-2), 2(1), 3(4,3-4), 4

(1), 5(1), 6(2), 7(5,4-6), 8(1), 9(1), 10(1,1-2), 11(2), 12(1,1-2), 13(2,2-3), 14(1).

Segment V: hair 0(1), 1(2), 2(1), 3(1), 4(5,4-6), 5(1,1-2), 6(2), 7(4,3-5), 8(1,1-

2), 9(1), 10(1), 11(2), 12(€1), 13(2), 14(1). Segment VI: hair 0(1), 1(3,2-3), 2

(1), 3(1,1-2), 4(4,2-4), 5(1), 6(1), 7(2,2-3), 8(2,1-3), 9(1), 10(1), 11(2), 12(1),

13(?), 14(1). Segment VII: hair 0(1), 1(€1), 2(1), 3(3,2-4), 4(1), 5(2,1-3), 6(7,5-

9), 7(2,1-2), 8(6,6-9), 9(1), 10(1), 11¢€1), 12(1), 13(3,2-3), 14(1). Segment VII:

Comb scales very slender in basal three-quarters, apex fringed. Hair 0(1), 1(3,2-

4), 2(1), 3(5,4-6), 4(1), 5(4,3-5), 14(1). Siphon: Integument yellowish. Index about

3.95-4.35. Pecten teeth usually 5(4-7), usually bifid, sometimes trifid, basal tooth

shorter. Hair 1(6,4-6), hair lad(2,2-3), hair lav(3,2-3); hairs 2,6,7,8 and 9 all sin-

gle. Anal Segment: Small ventral saddle present. Gill about 0.5 of dorsal saddle

length. Hair 1-X shorter than gill. Hair 1(2,1-3), 2(1), 3(6,4-7), 4a(10,9-13), 4b

(9,8-10), 4c(8,8-10), 4d(10,8-11), 4e(11,8-13), 4f(10,7-10).

SYSTEMATICS. I am assigning provisionally to atlanticus all the Atlantic pop-—

ulations of the southern complex of the Spanius Group on the basis of similarity

in the genitalia of all the males I have seen from the area. This species is indis-

tinguishable from its geminate Pacific representative of the complex, spanius, in

external features of the adults but is easily differentiated from it in the male

genitalia as noted in the key and descriptions. The immature stages of atlanticus

are known only from the type locality of Ocumare de la Costa and adjacent areas

on the northcentral coast of Venezuela; those of spanius are completely unknown.

Deinocerites atlanticus, as currently interpreted, has a much wider distribution

than any other species of the group, having been found in Panama, Canal Zone,

Colombia, the mainland of Venezuela and its outlying island of Orchila. At least

the Panamanian (including Canal Zone) population shows some differences in the

female genitalia from the topotypic Venezuelan population, having more setae on

the lobe of tergite IX, usually 4 (2-5) instead of 1 (0-2), and the postgenital plate

less deeply notched and its lateral lobe slender. The status of this population,

which was formerly confused with spanius, cannot be determined with certainty

until its immature stages are found and compared thoroughly with those of topo-

typic atlanticus; but the fact that the single female I have seen from Colombia

is intermediate in its genitalic characters between the Venezuelan and the Pana-

manian populations suggests that the differences noted may be clinal only and

that only 1 rather variable species may be involved over the entire range of at-

lanticus.

BIONOMICS. The immature stages of atlanticus are definitely known only from

the coast of Venezuela where they have been collected principally in small crab-

holes along water courses in coconut groves in areas of scrub and thorn forest.

The records of spanius immatures from Panama in Dyar (1928:265) may pertain

| Adames: Genus Deinocerites 31

to atlanticus but there are no specimens to substantiate them and both larvae

and pupae are still unknown from this area.

In Venezuela, adults have been collected together with the immature stages in

the situations noted above. In Colombia, they were collected once in a house in

Manaure in a thorn woodland. They have been collected on numerous occasions

in the Canal Zone in the coastal rainforest but without any record of the type

of crabhole. The only report of the biting habits of atlanticus is that of Dyar

(1925:154, as spanius): ‘“‘the species has lately been found to bite [man] freely”’.

This record may refer to the females collected by D. Baker from October 1924

to May 1925.

This species, as currently interpreted, appears to be quite common in Panama.

In Venezuela, its breeding associates are melanophylum and a member of the Cu-

lex inflictus complex.

DISTRIBUTION (fig. 3). Atlantic coast from the Canal Zone in Panama to the

state of Aragua and the island of Orchila in Venezuela. Material examined: 648

specimens; 123 males, 160 females, 245 larvae, 120 pupae; 88 individual rearings

(SO larval, 29 pupal, 9 incomplete).

COLOMBIA. Guajira: Uribia, Manaure, P. Barreto [USNM].

PANAMA AND CANAL ZONE. Canal Zone: Arenal River, C.S. Ludlow [USNM]. Cativa,

J.B. Shropshire [USNM]. Ft. Davis, D. Baker [UCLA, USNM]. Ft. Randolph, J. Zetek [UCLA];

D. Baker [USNM]; W.P. Murdoch [UCLA]. Ft. Sherman, J. Zetek [UCLA, USNM], D. Baker

[USNM]; W.H.W. Komp [UCLA]. France Field, C.S. Ludlow [USNM]. Locality not specified,

J.B. Shropshire [USNM]. Margarita, J.B. Shropshire [USNM]. Toro Pt., J.B. Shropshire [USNM].

VENEZUELA. Aragua: Cuyagua (VZ 415) [UCLA]. Ocumare del la Costa (VZ 170,175,

346 ,363-365,385,386) [UCLA, USNM]. Puerto Colombia (VZ 216) [UCLA]. Turiamo, W.H.W.

Komp [UCLA, USNM]. Carabobo: Moron (VZ 256-258) [UCLA]. La Orchila, Isla: F.M. Root

(LAR 63) [UCLA, USNM].

3. Deinocerites mathesoni Belkin & Hogue

Figs. 3,12-14

1959. Deinocerites mathesoni Belkin and Hogue, 1959:426-427. TYPE: Holotype male, Browns-

ville, Texas, 8-9 Feb 1940, reared, F.W. Fisk [USNM, 64261].

Deinocerites mathesoni of Stone, Knight and Starcke (1959:285); Dodge (1963:803,811; 1966:

376); Peyton, Reinert and Peterson (1964:449-458); Ross (1964:104).

Deinocerites spanius of Fisk (1941:543-550); Fisk and LeVan (1941:945); McGregor and Eads

(1943:939); Matheson (1944:251-252); Randolph and Neill (1944:81); Knight and Chamber-

lain (1948:10); Eads, Menzies and Ogden (1951:42,44); Breland (1956:95); Pratt (1956:8);

Evans (1962:255).

Deinocerites spanius in part of Carpenter and LaCasse (1955:327-329).

?Deinocerites spanius of Vargas (1956:30).

FEMALE (fig. 12). Wing 2.50 mm. Proboscis 1.23 mm. Forefemur 1.22 mm.

Abdomen about 2.54 mm. Antenna: Torus without scales; tip of proboscis reach-

ing from apex of flagellar segment 9 to base of 10. Thorax: Ppl sometimes with

1 or more scales; metameron without microsetae. Legs: Scaling and bristles as des-

cribed for atlanticus.

FEMALE GENITALIA (fig. 12). Tergite IX with 2-5 setae on each side. Cercus

in lateral aspect broadened at base and: evenly narrowed distally, sternal margin

32 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

convex, tergal margin almost straight throughout except for a slight concavity in

distal half; apex slightly upturned, with 3-5 apical or subapical bristles. Distal part

of postgenital plate (in ventral aspect) distinctly notched; lateral lobe elongate (in

lateral aspect), its apical bristle convergent distally with its mate (in ventral aspect).

MALE (fig. 12). Wing 2.39 mm. Proboscis 1.41 mm. Forefemur 1.27 mm. Ab-

domen (not including genitalia) about 1.83 mm. Essentially similar to female.

MALE GENITALIA (fig. 13). Segment IX: Tergite lobe slender, flattened be-

yond basal part, constricted subapically and then expanded, sometimes widened

near middle; ventral surface with a basal projection. Sidepiece: Similar to atlan-

ticus. Phallosome: Dorsal paramere with 2 sclerotizations, tergal and sternal, con-

nected by a narrow bridge very poorly sclerotized on the margins and membran-

ous centrally; tergal sclerotization bearing the long slender apical spine which is

strongly hooked apically; sternal sclerotization usually with 2 or sometimes 3 out-

standing teeth. Width of dorsal paramere about 0.6 of the length of tergal scler-

otization. Aedeagus in general similar to atlanticus.

PUPA (fig. 13). Abdomen 2.56 mm. Trumpet 0.38 mm. Paddle 0.53 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 6 reared specimens.

Hair 10-C about twice the length of 11-C; hair 1-VII extending beyond apex of

4-VIII. Cephalothorax: Integument yellowish. Hair 1(2,2-4), 2(3,3-4), 3(3,3-4), 4

(6,4-7), 5(2), 6(2,1-3), 7(5,4-6), 8(2,2-3), 9(3,3-4), 10(2), 11(4,3-4), 12(4,3-5).

Trumpet: Short, almost uniform in width throughout except for slight broaden-

ing apically; index about 5.76-6.16. Integument yellowish distad, medium brown

on tracheoid, not markedly contrasting with cephalothoracic integument. Trach-

eoid about 0.4 of the trumpet length; border of pinna with a distinct emargina-

tion. Abdomen: Integument yellowish. Segment I: hair 1(8-15, primary branches),

2(1,1-2), 3(2), 4(4,3-4), 5(5,4-7), 6(1), 7(4,4-6), 9(1). Segment II: hair 0(1), 1

(11,7-11), 2(2,2-3), 3(?), 4(3-4), 5(1), 6(1), 7(3,2-4), 9(1). Segment III: hair 0(1),

1(6,5-6), 2(1), 3(2,2-3), 4(1,1-2), 5(3,1-3), 6(1), 7(2,1-3), 8(6,4-6), 9(1), 10(3-4,

3-6), 11(1), 14(1). Segment IV: hair 0(1), 1(4,3-6), 2(1), 3(4,3-5), 4(1,1-2), 5(1,

1-2), 6(1), 7(2,1-2), 8(3,1-4), 9(1), 10(3,2-5), 11(1,1-2), 14(1). Segment V: hair

OD), MAS), 201), -363;1-3), 44-6), 901), 6(1,1-2); 7(5,3-6), -8(2,2-3), 904),

10(2,1-3), 11(1), 14(1). Segment VI: hair 0(1), 1(4,3-5), 2(1), 3(3,3-4), 4(5,4-5),.

S(1), 6(1), 7(2,1-2), 8(2,2-3), 9(1), 10(3,2-3), 11(1), 14(1). Segment VII: hair

O(1), 1€1,1-2), 2(1), 3(2,2-3), 4(3,2-4), 5(3,1-3), 6(1,1-2), 7(2,1-2), 8(3,2-3), 9(5,

4-5), 10(2,1-2), 11(2,1-2), 14(1). Segment VIII: hair 0(1), 4(3), 9(1), 14(1). Pad-

dle: Width about 0.67 of length.

FOURTH INSTAR LARVA (fig. 14). Head 0.72 mm. Siphon 0.71 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 4 reared specimens.

Hair 6-III always single as opposed to atlanticus and belkini in which it is either

single or double. Head: Integument yellowish. Mental plate distinctly wider than

long, triangular, with or without a median terminal spicule; basal spicule short-

er; all spicules with denticulate apex. Hair 0(1), 1(1), 2(1), 3(sometimes developed

as a spicule), 4(7,6-7), 5(6,5-6), 6(4,3-4), 7(10,8-11), 8(8,2-3), 9(3,2-7), 10(2,2-3),

11(4,3-6), 12(2,1-3), 13(3,3-4), 14(1), 15(2,2-3). Antenna: Length about 0.46 of

head; shaft with a few spicules on proximal part; hair 1(4,5-7). Thorax: Protho-

rax: hair 0(7,4-7), 1(1), 2(1), 3(2,1-2), 4(4,3-5), 5(1), 6(1), 7(4,3-5), 8(2,2-3),

91), 10(1), 11(4), 12(1), 14(1). Mesothorax: hair 1(1), 2(1), 3(1,1-2), 4(3,1-4),

S(1). 6(1), 7(1), 8(9,6-11), 9(8,8-14), 10(1), 11(2), 12(1), 13(?). Metathorax:

hair 1(1), 2(2), 3(4,3-4), 4(1,1-2), 5(1), 6(3,2-3), 7(9-14), 8(6), 9(10,7-11), 10

(1), 11(2), 12(1), 13(4,3-5). Abdomen: Segment I: hair 1(1), 2(1), 3(3,2-3), 4

Adames: Genus Deinocerites 33

(5,5-9), 5(1,1-3), 6(2,2-3), 7(3,2-3), 9(2,1-3), 10(2,1-2), 11(2,2-5), 12(3,2-3), 13

(2,1-2). Segment II: hair 0(1), 1(1), 2(1), 3(4,3-5), 4(3), 5(1,1-3), 6(1), 7(4,3-4),

8(3,2-5), 9(1), 10(1,1-4), 11(3,1-3), 12(2), 13(2). Segment III: hair 0(1), 1(3),

2(1),.. 3(3,2-3), 4(2), 51), 6(1), 706), 8C1,1-2), 9(1), 10(2,1-2), 11(@2), 12(2,1-2),

13(1), 14€1). Segment IV: hair 0(1), 1€1), 3(4,2-4), 4€1), 5(1), 6(2,1-2), 7(4,3-

7), 8(1), 9€1), 10(1,1-2), 11(2,1-2), 12(2), 13(2), 14(1). Segment V: hair 0(1),

11),20), 3C1), 40545), SCD); 6@,1-2), 7(3,3-6),. 8(1); 9C1), TOC), TiC2, 1-2),

12(1,1-2), 13(2), 14(1). Segment VI: hair O(1), 1(2), 2(1), 3(1), 4(4,3-4), 5(2),

6(1), 7(2), 8(2,2-3), 9(1), 10(€1), 11(2), 12(€1), 13(2), 14€1). Segment VII: hair

OCD); 11), 2G), 3231-3), 41) 502) 606,6-7) 7 A=2), S(6S-6); OC, 1-2), 10th),

11(1), 12(1), 13(2,2-4), 14(1). Segment VIII: Comb scales slender basally, slightly

widened around middle, apex fringed. Hair 0(1), 1(4,3-4), 2(1), 3(5,3-7), 4€1), 5(4,

3-5), 14(1). Siphon: Integument yellowish. Index about 3.6-4.3. Pecten teeth 4

(3-6), usually bifid, sometimes trifid; basal tooth shorter. Hair 1(4,3-7), hair lad

(4,1-5), hair lav(4,3-4); hairs 2,6,7,8 and 9 all single. Anal Segment: Gill about

0.5 of dorsal saddle length. Hair 1-X about the same length as gill. Hair 1(1),

2(5,5-8), 3(1), 4a(10,9-12), 4b(8,8-9), 4c(8,7-9), 4d(9,8-10), 4e(9,8-10), 4f(7,6-9).

SYSTEMATICS. Deinocerites mathesoni is the Atlantic member of the north-

ern complex of the Spanius Group. Its adults are indistinguishable from those of

its geminate Pacific species, belkini, in external features but the 2 species are eas-

ily separated by male genitalic,.pupal and larval characters as noted in the keys

and descriptions. ,

This species is definitely known only from southern Texas but it is very likely

that it occurs along the Atlantic coast of Mexico and that the records of spanius

from the states of Tamaulipas and Veracruz (Vargas, 1956:30) pertain to it.

There is no doubt, now that the Spanius Group has been shown to consist

of 4 distinct species, that mathesoni is indigenous to Texas and that it was not

introduced there by aircraft as suggested by Fisk (1941:548) who first recorded

this species (as spanius) from the United States.

BIONOMICS. The bionomics of mathesoni have been investigated by Fisk (1941,

as spanius) and by Peyton, Reinert and Peterson (1964). Fisk reported the im-

mature stages from holes of the fiddler crab, Uca pugilator, and Peyton et al found

them very commonly in the holes of Uca subcylindrica and less frequently in those

of the Black Land Crab, Gecarcinus lateralis. Deinocerites mathesoni appears to

prefer holes ranging from % to 1% inches but occasionally has been found in holes

of only % inch diameter. Chloride content determinations varied from 8,430 ppm

for a single composite sample by Fisk and from 1,115 to 2,603 ppm for several

individual samples by Peyton et al. The range of pH values, determined colori-

metrically by Fisk and Peyton et al was 7.2-7.6.

Females of mathesoni have not been reported to feed on man and Fisk fed

them on a small turtle. He reports finding both sexes several miles trom any ob-

vious breeding site. Fisk also observed mating of reared individuals in the morning

of the eighth day after the first adults emerged.

Deinocerites pseudes is the only species reported to be associated with mathe-

soni to date (Peyton, Reinert and Peterson (1964)).

DISTRIBUTION (fig. 3). Atlantic coast in Texas from Corpus Christi to Browns-

ville and probably southward into Mexico, at least to Veracruz. Material examined:

78 specimens; 26 males, 28 females, 18 larvae, 6 pupae; 4 individual larval rearings.

TEXAS. Cameron County: Brownsville, F.W. Fisk [UCLA, USNM]; T.N. Burns; E.S. Ross

[USNM]; J.F. Reinert; M.D. Huettel (TEX 13-16,19,20); D. Eyles (KO 32-7) [UCLA]. Har-

34 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

lingen [USNM]. Laguna Vista, S.R. Telford (MT 10) [UCLA]. Locality not specified [USNM].

Nueces County: Corpus Christi, F.R. du Chanois [USNM]. Willacy County: Tenerias triangulation

station, M.D. Huettel (TEX 10,11) [UCLA].

4. Deinocerites belkini Adames, n.sp.

Figs. 3,15-17

TYPES: Holotype male with associated larval and pupal skins and genitalia slide (MT 4-12),

Barra de Navidad, Jalisco, Mexico, 13 July 1963, S.R. Telford, Jr. [USNM]. Allotype female

with associated larval and pupal skins (MT 4-10), same data as holotype [USNM] . Paratypes:

9 IpM (MT 4-13,14,20,25,30,31,33,35,37), 8 IpF (4-11,22-24,26,32,34), 3 pM (4-16,17,27), 2

pF (4-19,28), 1 IP (4-36), 22 L (4), same data as holotype [BM, UCLA, USNM].

FEMALE (fig. 15). Wing 2.57 mm. Proboscis 2.52 mm. Forefemur 1.53 mm.

Abdomen about 2.09 mm. Apparently indistinguishable from mathesoni.

FEMALE GENITALIA (fig. 15). Tergite IX with 1-6 setae on each side. Cercus

and postgenital plate essentially similar to mathesoni.

MALE (fig. 15). Wing 2.41 mm. Proboscis 1.27 mm. Forefemur 1.47 mm. Ab-

domen (not including genitalia) about 2.29 mm. Essentially similar to female.

MALE GENITALIA (fig. 16). Segment IX: Tergite lobe slender, slightly wid-

ened near middle, flattened beyond basal part, constricted subapically and then

expanded; ventral surface with a very strong premedian projection, visible in toto

only in lateral aspect. Sidepiece: Similar to atlanticus. Phallosome: Dorsal para-

mere essentially as in mathesoni except for usually 3 or 4 teeth on sternal sclero-

tization and the width which is about 0.5 of the length of tergal sclerotization.

Aedeagus in general similar to atlanticus.

PUPA (fig. 16). Abdomen 2.57 mm. Trumpet 0.61 mm. Paddle 0.61 mm. Di-

agnostic characters as in the key; general chaetotaxy based on 10 reared speci-

mens. Hair 10-C about 1.3 length of 11-C; hair 1-VII as in mathesoni. Cephalo-

thorax: Integument yellowish. Hair 1(2,2-3), 2(3,2-3), 3(2,2-4), 4(4,3-5), 5(2,2-3),

6(2,1-3), 7(4,3-6), 8(1), 9(3,2-4), 10(1,1-2, forked when double), 11(3,3-4), 12(4,

3-5). Trumpet: Moderately long and slender, almost uniform in width throughout

except for broadened apex; index about 8.7-11.0. Integument yellowish distad,

medium brown on tracheoid; not markedly contrasting with cephalothoracic inte-

gument. Tracheoid about 0.4 of trumpet length; border of pinna without a dis-

tinct emargination. Abdomen: Integument yellowish. Segment I: hair 1(14,12-16

primary branches), ’2(1,1-2), 3(2), 4(3,2-3), 5(5,3-8), 6(1), 7(3,3-5), 9(1). Segment

II: hair 0(1), 1(4-9), 2(2), 3(1,1-2), 4(4,3-6), 5(1), 6(1), 7(2,1-3), 9(1). Segment

III: hair O(1), 1(6,4-7), 2(1), 3(2), 4(2,1-4), 5(€1,1-3), 6(1), 7(2,1-3), 8(7,6-10),

9(1), 10(2,2-3), 11(1), 14(1). Segment IV: hair 0(1), 1(5,4-6), 2(1), 3(4,4-5), 4

(2,1-3), 5(1,1-2), 6(1), 7(3,2-4), 8(3,2-4), 9(1), 10(3,2-4), 11(€1), 14(1). Segment

V: hair O(1), 1(4,3-5), 2(1), 3(3,2-3), 4(5,4-6), 5(1), 6(1,1-3), 7(5,4-5), 8(3,2-3),

9(1), 10(2), 11(1), 14(1). Segment VI: hair 0(1), 1(2,1-3), 2(€1), 3(2,1-2), 465,

3-6), 5(1), 6(1), 7(2,1-3), 8(3,2-5), 9(1), 10(2,1-2), 11(1). Segment VII: hair 0

CT), 1C2), 28), ete, bad) 422-3), OCZ 1-3), OCU), TZ lth, BC St, QF)... 910.128),

10(2,1-2), 11(1,1-2), 14(1). Segment VIII: hair 0(1), 4(2,2-3), 9(1), 14(1). Pad-

dle: Width about 0.75 of length.

FOURTH INSTAR LARVA (fig. 17). Head 0.94 mm. Siphon 1.19 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 7 reared specimens.

Adames: Genus Deinocerites 35

Head: integument yellowish. Mental plate distinctly wider than long, without med-

ian terminal spicule; all spicules with denticulate apex, basal and lateral spicules

longer than terminal. Hair 0(1), 1(1), 2(1), 3(not detectable), 4(7,5-7), 5(5,5-7),

6(3,2-4), 7(10,9-11), 8(3,2-3), 9(4,3-5), 10(2,2-3), 11(5,3-7), 12(4,2-4), 13(5,4-6),

14(1), 15(3,2-3). Antenna: Length about 0.45 of head; shaft with several spicules

on proximal part; hair 1(4,3-5). Thorax: Prothorax: hair 0(6,6-7), 1(1), 2(1), 3

(2,1-2), 4(4,3-5), 5(1), 6(1), 7(4,4-5), 8(3,2-3), 9(1), 10(1), 11(2,1-4), 12(1), 14

(1,1-3). Mesothorax: hair 1(1), 2(1), 3(1), 4(3,3-4), 5(1), 6(1), 7(1), 8(8,6-8), 9

(8,7-9), 10(1), 11(2,1-3), 12(1), 13(€11,10-11), 14(3-5 primary branches). Meta-

thorax: hair 1(1), 2(3,2-4), 3(4,3-6), 4(2,2-3), 5(1), 6(3,2-4), 7(10,8-12), 8(7,5-7),

9(9,6-10), 10(1), 11(2,2-3), 12(1), 13(5,4-5). Abdomen: Segment I: hair 1(1), 2

(1), 3(3,2-4), 4(8,8-10), 5(3,2-4), 6(2), 7(3,2-3), 9(3,2-3), 10(3,2-4), 11(2,2-3),

12(2,1-3), 13(1,1-2). Segment II: hair 0(1), 1(1), 2(€1,1-2), 3(4,3-4), 4(6,5-6), 5

(2,2-3), 6(1), 7(4,3-6), 8(3,3-4), 9(1), 10(3,3-4), 11(2,2-3), 12(2,2-3), 13(12,10-

13). Segment III: hair 0(1), 1(3,2-3), 2(1), 3(3,2-3), 4(2,1-2), 5(1,1-3), 6(1,1-

2), 7(5,4-6), 8(2,1-3), 9(1), 10(2), 11(2), 12(2), 13(1), 14(1). Segment IV: hair

0615, <1 A-2), 20); 3(5;,5-7), 21-2), St 152), 6022-3), G6 466), SUL 2), 9

(1), 10(2,1-2), 11(2,2-3), 12(2,1-2), 13(2), 14(1). Segment V: hair O(1), 1(1),

2(1), 3(1), 4(7,4-7), 5(1,1-2), 6(2,1-2), 7(5,5-7), 8(2,1-2), 9(1), 10(€1,1-2), 11@Q,

2-3), 12(1), 13(2), 14(1). Segment VI: hair 0(1), 1(2,1-3), 2(1), 3(1), 4(4,3-5),

5(2,2-3), 6(1), 7(3,2-3), 8(3,3-5), 9(1), 10(1), 11(2), 12€1), 13(16-22), 14(1). Seg-

ment VII: hair 0(1), 1€1), 2(1), 3(2,2-3), 4(€1), 5(2,1-3), 6(10,10-12), 7(2,1-2), 8(9,6-

11), 9(1,1-2), 10(€1,1-2), 11(€1,1-2), 12(€1), 13(3,2-5). Segment VIII: Comb scales

with basal two-thirds narrow, distal third wide and fringed. Hair 0(1), 1(3,3-4),

2(1), 3(5,5-6), 4(1), 5(4,3-5), 14(1). Siphon: Integument yellowish. Index about

5.2-6.4. Pecten teeth 4(4-5), all teeth about the same size, usually bifid, some-

times trifid. Hair 1(4,3-6), hair lad(2,2-3), hair lav(2,1-3); hairs 2,6,7,8 and 9

all single. Anal Segment: Gills about two-thirds of dorsal saddle length. Hair 1-X

about same length as gill. Hair 1(2,1-3), 2(4,3-5), 3(1), 4a(10,7-12), 4b(8,7-10),

4c(7,6-8), 4d(7,6-10), 4e(8,7-9), 4f(8,5-10).

SYSTEMATICS. Deinocerites belkini is the Pacific member of the northern com-

plex of the Spanius Group. It is easily differentiated from its Atlantic twin, math-

esoni, in the male genitalia, pupae and larvae but cannot be separated from it on

external features of the adults.

This species is known to date only from the 3 localities in the states of Nayarit

and Jalisco, Mexico, but it is very likely that it occurs southward of this area

toward the isthmus of Tehuantepec.

BIONOMICS. Immature stages of belkini have been found only once, in small

crabholes which may have been made by fiddler crabs. Adults have been collected

twice in large crabholes in association with the adults of mcdonaldi.

DISTRIBUTION (fig. 3). Pacific coast of Mexico in the states of Nayarit and

Jalisco. Material examined: 94 specimens; 16 males, 11 females, 42 larvae, 25 pu-

pae; 25 individual rearings (19 larval, 5 pupal, 1 incomplete).

MEXICO. Nayarit: Matanchen, San Blas (MF 6) [UCLA]. Jalisco: Barra de Navidad (MT 4)

[BM, UCLA, USNM]. Puerto Vallarta (MEX 466) [UCLA].

DYARI GROUP

FEMALES. Small species, wing 2.6-3.0 mm. Mesonotum dark, pleuron with

36 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

strongly contrasting whitish portions or almost uniformly dark brown not con-

trasting with mesonotal integument (barretoi). Antenna: Flagellar segment 1 with

scales, about equal to combined length of segments 2, 3 and two-thirds of 4; seg-

ments 2-12 subequal in length but progressively slightly shortened distad; mod-

erately long, exceeding proboscis from base of flagellar segment 8. Thorax: Post-

notum usually with a few setae in lower part. Apn light, slightly darkened anter-

iorly or entirely dark brown; ppn either light or brown, scales all narrow; ppl

light, slightly darkened or brownish; psp, ssp, stp and pra either markedly dark-

ened or concolorous with most of pleuron; paratergite dark; mep light or brown —

and without translucent scales, Imep with 1 very strong bristle; meron light or

brown; metameron and metapleuron either light or pale brownish. Legs: Coxal

integument either light, slightly darkened or brownish. Entire posteroventral sur-

face of all femora whitish; anteroventral margin of forefemur without spiniforms

but with 4-9 bristles, posterodorsal margin with weaker bristles. Abdomen: Ster-

nites III-VII dark scaled, II sometimes pale scaled.

FEMALE GENITALIA. Sternite VIII with sclerotized distal band narrow and

with only a few scales, gradually shortened ventrad; caudal margin not produced

into a submedian ventral angle, this area without strongly differentiated marginal

bristles. Tergite IX with at least 2 small setae on each side. Tergite X poorly dif-

ferentiated dorsally and with a very small indistinct lateral sclerotization. Cercus

with a few moderately strong apical or subapical normal bristles.

MALES. Essentially similar to females. Antenna: Exceeding proboscis from at

least middle of flagellar segment 5; flagellar segments 1, 2 and base of 3 with

scales; segments 1-5 or 1-6 markedly elongated but progressively shorter distad;

segment | always shorter than combined length of segments 2 and 3; segments

7-12 subequal but progressively slightly shortened distad; segment 13 slightly wid-

er than 12. Legs: Claws of foreleg and midleg enlarged; anterior member of a

pair always larger and with a subbasal tooth, posterior simple.

MALE GENITALIA. Segment IX: Tergite lobe cylindrical; angled laterad at base;

distal part varied in shape and length, apex at most reaching base of subapical

lobe. Sidepiece: Without scales. Apicosternal lobe prominent; long, slender and

with a long apical seta. Phallosome: Dorsal parameres widely separated in tergal

aspect, with a slight indication of a very short, narrow, incomplete sclerotized

dorsal bridge; articulated ventrally with a subbasal dorsal sclerotized projection

of aedeagus.

PUPAE. Cephalothorax: Hairs 2,3-C closely approximated, 2-C poorly developed,

shorter than 3-C; hair 5-C single or double, strongly developed, longer than dis-

tance from its alveolus to base of trumpet, at least 1.5 of trumpet length; 7-C

usually double (1-3); hair 8-C single. Metanotum: Hair 10-C poorly developed, al-

ways shorter than 11-C (0.3-0.5), usually single (1 or 2). Abdomen: Hair 1-II us-

ually not reaching apex of tergite III, single or triple, barbed; 1-III-VI branched;

S-II always laterad of 3-II; hair 3-V usually double (1-4); hair 1-VII shorter than

tergite VIII, at most reaching base of 4-VIII. Paddle: Hair 1-P as long as or longer

than paddle.

FOURTH INSTAR LARVAE. Head: Hair 2-C short, inconspicuous, about 0.20-

0.25 of 1-C, either slightly mesad, in line with or laterad of 1-C; hair 5-C usually

with 3 or 4 barbed branches (2-5); hair 6-C single and barbed, about 1.25-1.33

of 5-C. Thorax: Prothoracic hair 9-P single. Abdomen: Hair 6-II double; dorsal

sensillum of segment V laterad of 4-V; hair 6-VI usually double (1 or 2); hair

I-VII short, shorter than 3-VII and not reaching base of segment VIII; hair 1-

Adames: Genus Deinocerites 37

VIII usually with 4-6 branches (4-7). Siphon: Hair 1-S usually double or triple.

Anal Segment: Ventral brush (4-X) with 6 or 7 pairs of hairs. Gill slightly emar-

ginate on apex or towards ventral margin; short, about 0.60-0.70 of dorsal saddle

length. Dorsal saddle narrow; its ventral margin far from hair 1-X.

DISCUSSION. The Dyari Group shows fewer derived features than any other

group except Spanius and in some respects may be regarded as transitional be-

tween the latter and the other groups of the genus. The adults are small species,

only slightly larger than those of the Spanius Group, but they have distinctly long-

er antennae than the latter and show sexual dimorphism in the flagellum as do

all the groups other than Spanius. In general external features, the adults show

more resemblance with the Cancer Group than with the 2 other derived groups

(Epitedeus and Pseudes). They are usually readily distinguished from the Cancer

Group by the presence of postnotal bristles. The male genitalia do not show any

distinctive characters on a group level but sternite VIII of the female is uniquely

developed. The immature stages provide the most reliable diagnostic features for

the group in the reduced length of pupal hair 10-C and larval hair 1-VII.

The Dyari Group as now understood is diagnosed in the adults essentially as

it was by Belkin and Hogue (1959:428) except that a few scales may be present

in the female genitalia and the ninth tergite lobe of the male is not always short

as in dyari. It consists of 3 allopatric species with complementary known distri-

butions extending from Nicaragua and Costa Rica (nicoyae) through Panama and

northern Colombia (dyari) to central Colombia (barretoi). The group as a whole

has the most restricted range in the genus and is confined to the Pacific basin.

On the basis of the development of the dorsal paramere (expanded external

surface) and subapical lobe of the sidepiece (ventromesal spiniforms), nicoyae is

sharply differentiated from the other 2 species and it appears likely that it was

separated early from the stock which gave rise to the dyari-barretoi complex. It

would appear also that the short tergite [IX lobe of dyari is a derived state since

barretoi has this lobe similar to that of nicoyae.

The species of the group appear to be uncommon as they have seldom been

collected even as adults. Because of the elongate antennae they are not as likely

to be confused with unornamented species of the subgenus Melanoconion of Culex

as species. of the Spanius Group. Recent studies have shown that dyari females

have a definite preference for reptilian blood but nothing is known of the blood

feeding habits of the other 2 species. The immature stages of the Dyari Group

are apparently very difficult to locate and only a few collections of nicoyae and

dyari have been made to date. Both of these species were found in the medium-

sized burrows of Ucides occidentalis. The immature stages of barretoi are unknown.

5. Deinocerites dyari Belkin & Hogue

Figs. 4,18-20

1959. Deinocerites dyari Belkin and Hogue, 1959:428-429. TYPE: Holotype male, Corozal,

Panama, Canal Zone, 20 Apr 1919, J. Zetek, 1183 [USNM, 64262].

Deinocerites dyari of Stone, Knight and Starcke (1959:284); Gorgas Memorial Institute (1970:

20); Tempelis and Galindo (1970).

FEMALE (fig. 18). Wing 2.67 mm. Proboscis 1.22 mm. Forefemur 1.53 mm.

38 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Abdomen about 2.55 mm. Mesonotum dark brown, pleuron with strongly con- —

trasting whitish portions. Head: Narrow decumbent scales of vertex creamy; erect

scales brown to very dark brown; lateral patch of broad decumbent scales whitish.

Antenna: Torus occasionally with 1 scale. Thorax: Apn light or slightly darkened

anteriorly; ppn light and often with a few scales in middle; ppl light or slightly

darkened, with several bristles and frequently a few scales; psp, ssp, stp and pra

usually markedly darker than rest of pleuron; mep light, with a few scales in up-

per part, microsetae absent; metameron simple; meron, metameron and metapleu-

ron light. Legs: Coxal integument usually light; anterior surface of forecoxa cov-

ered with bristles and translucent scales, basolateral area with a few bristles; mid-

coxa with scales anterolaterally, with 2 groups of bristles laterally, the anterior

bristles irregular, the posterior in a row and stronger, posterolateral surface sim-

ple; hindcoxa with scales and several bristles laterally, posterior surface with bristles. __

FEMALE GENITALIA (fig. 18). Tergite IX usually with 4 setae on each side

(2-5). Cercus (in lateral aspect) broadened basally, gradually narrowing distally,

sternal margin convex, tergal margin more or less straight throughout except for

a slight subapical concavity; apex slightly upturned, usually with 4-6 moderately

strong apical or subapical bristles. Distal part of postgenital plate (in ventral as-

pect) deeply notched, margins of notch diverging distad; lateral lobe elongate (in

lateral aspect), its apical bristle slightly converging with its mate (in ventral as-

pect), its small dorsal bristle not converging.

MALE (fig. 18). Wing 2.61 mm. Proboscis 1.88 mm. Forefemur 1.78 mm. Ab-

domen (not including genitalia) about 1.88 mm. Flagellar segments 1-6 markedly

elongated. Anterior claw of foreleg with a long, very slender tooth, anterior claw

of midleg with a shorter and heavier tooth.

MALE GENITALIA (fig. 19). Segment IX: Tergite lobe with distal part short,

more or less pointed and conical, not reaching more than halfway to subapical

lobe. Sidepiece: Subapical lobe with a distinct thumb; seta c rather heavy and

spiniform but with an apical attenuation; the 3 setae of ventromesal surface bris-

tlelike and attenuated apically. Phallosome: Dorsal paramere with apical spine long

and strongly curved, ventral teeth large and arising from convex caudolateral bor-

der. Aedeagus (in tergal aspect) almost uniform in width except for a subbasal

expansion; distally the 2 plates come together forming a globular expansion.

PUPA (fig. 19). Abdomen 3.52 mm. Trumpet 0.51 mm. Paddle 0.71 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 8 reared specimens.

Hair 5-IV usually reaching spiracular sensillum of segment V. Cephalothorax: In-

tegument light brown. Hair 1(2), 2(2,2-4), 3(2,1-2), 4(3,1-4), 5(1), 6(1), 7(2,1-2),

8(1), 9(2,1-2), 10(1,1-2), 11(2,1-3), 12(2,1-3). Trumpet: Short, index 3.6-5.0. Inte-

gument brown distad, darker on tracheoid, contrasting with cephalothoracic integu-

ment. Tracheoid about 0.36 of trumpet length. Abdomen: Integument light brown.

Segment I: hair 1(8,4-13 primary branches), 2(1), 3(2,1-2), 4(4,2-4), 5(2,1-3), 6(1),

7(2,2-4), 9(1), 10(€1), 11(1). Segment II: hair 0(1), 1(€1), 2(1,1-2), 3(1), 4(2,1-4),

S(1), 6(1), 7(1,1-2), 9(1). Segment III: hair 0(1), 1(4,2-5), 2(1), 3(2,1-2), 4(1,

1-2), 5(1,1-2), 6(1), 7(€2,1-3), 8(3,2-6), 9(1), 10(2,1-2), 11(1), 14(1). Segment IV:

hair, OC), 412,15), 32C19;; S(3)2-3)5 4), 501), Oh). HE IQ), BE 1-2) 901) 10

(2,1-2), 11(1), 14(1). Segment V: hair 0(1), 1(2,1-3), 2(1), 3(2,1-4), 4(3,2-4), 5

(1), 6(1), 7(2,2-4), 8(2,1-2), 9(1), 10(2,1-2), 11(1), 14(1). Segment VI: hair 0(1),

1(2,2-3), 2(1), 3€1,1-3), 4(3,2-3), 5(1), 6(1), 7(1,1-2), 8(1,1-2), 9(1), 10(2,1-2),

11(1), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(2,1-2), 4(2,1-2), 5(1), 6(1),

7(2), 8(2,2-3), 9(3,3-5), 10(2,1-2), 1101), 14(1). Segment VIII: hair 0(1), 4(2,

Adames: Genus Deinocerites 39

1-2), 9(1), 14(1). Paddle: Width about 0.73 of length; apex usually slightly pro-

duced; hair 1-P longer than paddle.

FOURTH INSTAR LARVA (fig. 20). Head 1.11 mm. Siphon 1.07 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 7 reared specimens.

Head: Integument light brown. Mental plate more or less rectangular, marginal

spicules sharply pointed. Hair 0(1), 2(1), 3(not detectable), 4(3,2-4), 5(3,3-5), 6

(1), 7(9,7-11), 8(3,2-4), 9(5,5-6), 10(2,1-4), 11(3,2-4), 12(2,1-3), 13(4,4-6), 14

(1), 15(3,2-3). Antenna: Length about 0.38 of head; shaft with several spicules

on proximal part. Hair 1(5,4-7). Thorax: Prothorax: hair 0(5,5-7), 1(1), 2(1), 3

(11-2), 4(3,2-4), 5(1), 601), 7G), 82,23), 9G), 10U1), 732-4), 1201), Fac).

Mesothorax: hair 1(1), 2(2), 3(1), 4(3,2-4), 5(1), 6(1), 7(1), 8(6,5-7), 9(5,4-7),

~ 10(1), 11€3,2-3), 12(1), 13(8,6-8), 14(6,4-7). Metathorax: hajr 1(1), 2(3,3-4), 3

(4,3-5), 4(2,2-4), 5(1), 6(2,1-3), 7(9,7-10), 8(7,7-10), 9(8,6-9), 10(1), 11(3,2-3),

12(1), 13(4,4-6). Abdomen: Segment I: hair 1(1), 2(1), 3(3,3-4), 4(6,5-7), 5(3,

2-4), 6(2,2-4), 7(2), 9(3,3-4), 10(4,3-5), 11(3,2-4), 12(3,2-4), 13(1,1-2). Segment

II: hair 0(1), 1€1), 2(1), 3(3,2-3), 4(3,2-4), 5(4,3-5), 6(2), 7(3,2-3), 8(2,1-4), 9(1),

10(1,1-2), 11(2,2-4), 12(3,3-5), 13(7,7-9). Segment III: hair 0(1), 1(2,2-3), 2(1),

3(2,1-3), 4(1,1-2), 5(4,3-6), 6(3,3-4), 7(4,3-5), 8(2,1-3), 9(1), 10(2), 11(2,1-2),

12(2,1-2), 13(1,1-2), 14(1). Segment IV: hair 0(1), 1(2), 2(1), 3(4,2-4), 4(1), 5

(3,2-4), 6(3), 7(5,5-6), 8(1), 9(€1), 10(€2,1-2), 11(2,1-2), 12(2,1-2), 13(2), 14(1).

Segment V: hair 0(1), 1(2,1-2), 2(1), 3(1), 4(4,4-5), 5(2,1-3), 6(2,2-3), 7(5,3-6),

8(1), 9(1), 10(€1), 11(€2,1-2), 12(1), 13(2), 14(1). Segment VI: hair 0(1), 1(3,3-4),

2(1), 3(1), 4(4,3-4), 5(2,2-4), 6(2,1-3), 7(3,2-3), 8(2), 9(1), 10(1,1-2), 11(2), 12

(1), 13(13,9-14), 14(1). Segment VII: hair 0(1), 1(2), 2(1), 3(3,3-4), 4€(1), 5(,

2-3), 6(7,6-8), 7(2,1-2), 8(5,4-7), 9(2,1-2), 10(2,1-2), 11€1,1-2), 12(1), 13(2,2-3),

14(1). Segment VIIT. Comb scales with body very slender, apex slightly wider

and fringed. Hair 0(1), 1(4,4-5), 2(1), 3(4,3-6), 4(1), 5(3,3-4), 14(1). Siphon: In-

tegument medium brown. Index about 4.6-5.8. Pecten teeth 4(3-5), usually bifid,

sometimes trifid, both blades very slender, size variable. Hair 1(2,2-3), hair lad

(2), hair lav(2,1-2); hairs 2,6,7,8,9 all single. Anal Segment: Gill about 0.7 of

dorsal saddle length. Hair 1-X shorter than gill. Hair 1(3), 2(8,6-9), 3(1), 4a(12,

11-14), 4b(10,10-11), 4c(11,9-12), 4d(11,10-14), 4e(13,11-14), 4£(12,11-14).

SYSTEMATICS. This species appears to be the most derived one of the group

judging by the reduction of the IX tergite lobe of the male genitalia. In other

respects the male genitalia are very similar to those of barretoi and I therefore

consider that these 2 species are closely related. As noted above in the descrip-

tion and in the keys, the adults of dyari are readily distinguished from barretoi

by the lighter pleural integument which strongly contrasts with the dark meso-

notum.

BIONOMICS. The immature stages of dyari are known only from Curiche, Co- |

lombia, where they were found in large crabholes in a mangrove area. On 1 oc-

casion adults were found in this locality in a hole in which a Wide Red Land

Crab, Ucides occidentalis, was also collected (COA 26).

Studies on the blood feeding habits of species of Deinocerites in Panama indi-

cate that dyari has a definite preference for reptiles (Gorgas Memorial Institute,

1970:20; Tempelis and Galindo, 1970).

This species has been found associated with pseudes.

DISTRIBUTION (fig. 4). Pacific coast from the Canal Zone, Las Perlas islands

southward to Curiche, Colombia. Material examined: 116 specimens; 35 males,

30 females, 32 larvae, 19 pupae; 19 individual rearings (9 larval, 5 pupal, 5 in-

complete.

40 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

COLOMBIA. El Choco: Curiche (COA 22,24-29) [BM, UCLA, USNM].

PANAMA AND CANAL ZONE. Canal Zone: Corozal, J. Zetek [UCLA, USNM]. La Boca,

A. Busck [USNM]. Darien: Boca Lava, B.F. Eldridge [USNM]. sla San Jose (Archipielago de

las Perlas), J.P.E. Morrison [USNM]. Panama: Punta Paitilla, Panama City, GML [UCLA].

6. Deinocerites barretoi Adames, n.sp.

Figs. 4,21

TYPES: Holotype male with genitalia slide (COL 47f), mouth of Raposo River, Valle, Colom-

bia, 31 Mar 1955, V.H. Lee [USNM]. Allotype female with genitalia slide (COL 47a), same

data as holotype [USNM]. Paratypes: 2 M (COL 47h,g), 3 F (47c-e), same data as holotype

[UCLA, USNM].

Deinocerites dyari of Barreto and Lee (1969:435,436).

FEMALE (fig. 21). Wing 3.06 mm. Proboscis 1.88 mm. Forefemur 1.78 mm.

Abdomen about 2.60 mm. Mesonotum dark brown; pleuron almost uniformly dark

brown, not markedly contrasting with mesonotum. Antenna: Torus without scales.

Thorax: Apn dark brown; ppl dark brown, with a few scales in middle; ppl dark

brown, with several bristles and at least 1 scale; psp, ssp, stp and pra dark brown

and concolorous with most of pleuron; mep dark brown, with a few scales in

upper part; microsetae absent; metameron simple; meron brown; metameron and

metapleuron lighter than adjacent portions of pleuron. Legs: Coxal integument

slightly brownish; scaling and bristles as described for dyari.

FEMALE GENITALIA (fig. 21). Tergite IX with 1-4 setae on each side. Cercus

(in lateral aspect) broadened basally and evenly narrowing distally; sternal and ter-

gal margins convex; apex not upturned, with 4 or 5 moderately strong apical or

subapical bristles which may be twisted apically. Distal part of postgenital plate

(in ventral aspect) deeply notched, margins of notch parallel; lateral lobe elon-

gate (in lateral aspect), its apical and dorsal bristles converging with their mates

(in ventral aspect).

MALE (fig. 21). Wing 2.72 mm. Proboscis 1.78 mm. Forefemur 1.73 mm. Ab-

domen (not including genitalia) about 2.14 mm. Flagellar segments 1-5 markedly

elongated. Anterior claws of foreleg and midleg similar to those of dyari.

MALE GENITALIA (fig. 21). Segment IX: Tergite lobe with distal part long,

cylindrical, slender and reaching base of subapical lobe but not extending beyond.

Sidepiece: Subapical lobe with a distinct thumb; seta c rather heavy, spiniform

and bent apically; the 3 setae of ventromesal surface bristlelike and attenuated

apically. Phallosome: Dorsal paramere only known in tergal aspect, apical spine

long and strongly curved. Aedeagus apparently similar to that of dyari.

PUPA and LARVA. Unknown.

SYSTEMATICS. Deinocerites barretoi appears to be closely related to dyari on

the basis of general similarity in the male genitalia. However, it has an elongate

IX tergite lobe, a primitive character state in my opinion. Therefore I consider

that barretoi is probably the earlier derivative of this phyletic line. The adults of

barretoi, as noted above and in the keys, are readily separated from the other 2

species of the group by the dark pleural integument. The male of barretoi also

differs from the others in having only flagellar segments I -5 elongated instead of

segments 1-6.

Adames: Genus Deinocerites 41

BIONOMICS. Adults of barretoi have been collected from crabholes in a man-

grove area at the mouth of the Raposo River in Colombia. On | occasion a female

was caught in a treehole in the same area.

DISTRIBUTION (fig. 4). Pacific coast of Colombia near Buenaventura. Material

examined: 13 specimens; 5 males, 8 females.

COLOMBIA. Valle: Rio pede (COL 47), type series [BM, UCLA, USNM]; (VL 26,69)

[VALLE].

7. Deinocerites nicoyae Adames & Hogue

Figs. 4,22-24

1970. Deinocerites nicoyae Adames and Hogue, 1970:9. TYPE: Holotype male (CR 254-21)

with associated larval and pupal skins and genitalia slide, Estero El Mero, Boca del

Rio Barranca, Puntarenas Province, Costa Rica, 11 Feb 1969, C.L. Hogue and D.B.

Bright [USNM].

FEMALE (fig. 22). Wing 2.79 mm. Proboscis 1.69 mm. Forefemur 1.61 mm.

Abdomen about 2.83 mm. Apparently indistinguishable from dyari.

FEMALE GENITALIA (fig. 22). Tergite IX usually with 1 seta on each side

(0-3). Cercus (in lateral aspect) broadened basally, gradually narrowing distally,

sternal margin convex, tergal margin more or less concave; apex upturned, usually

with 4-6 moderately strong apical or subapical bristles. Distal part of postgenital

plate (in ventral aspect) deeply notched, margins of notch diverging distad; lateral

lobe elongate (in lateral aspect), its apical bristle converging with that of its mate

(in ventral aspect), its small dorsal bristle not converging.

MALE (fig. 22). Wing 2.91 mm. Proboscis 1.90 mm. Forefemur 1.90 mm. Ab-

domen (not including genitalia) about 1.67 mm. Flagellar segments 1-6 markedly

elongated. Anterior claws of foreleg and midleg both with a heavy tooth.

MALE GENITALIA (fig. 23). Segment IX: Tergite lobe with distal part reach-

ing base of subapical lobe, directed mesad distally by a distinct deep mesal curva-

ture or constriction, expanded distally. Sidepiece: Subapical lobe with distinct

thumb; seta c rather heavy and spiniform, with an apical attenuation; the most

anterior seta of ventromesal surface bristlelike and attenuated apically, the 2 pos-

terior setae spiniform, rather heavy and without apical attenuation. Phallosome:

Dorsal paramere with caudoventral margin expanded laterad as a broad hemispher-

ical ledge; apical spine short, heavy and strongly curved dorsad, visible in toto

only in lateral aspect; ventral teeth large, heavy and progressively shorter toward

apex. Aedeagus (in tergal aspect) broad in middle, then constricted; apex trun-

cate and poorly sclerotized but well defined by a subapical necklike constriction.

PUPA (fig. 23). Abdomen 3.56 mm. Trumpet 0.51 mm. Paddle 0.71 mm. Di-

agnostic characters as in the key; general chaetotaxy based on 6 reared specimens.

Hair 5-IV not reaching spiracular sensillum of segment V. Cephalothorax: Inte-

gument light brown. Hair 1(2), 2(2,1-3), 3(2,1-3), 4(3,2-4), 5(2), 6(1,1-2), 7(2, |

2-3), 8(1), 9(2), 10(1,1-2), 11(2), 12(2,2-3). Trumpet: Short, index about 4.5-

4.6. Integument brown distad, darker on tracheoid; markedly contrasting with cep-

halothoracic integument. Tracheoid about 0.45 of trumpet length. Abdomen: In-

tegument light brown. Segment I: hair 1(13-14,8-15 primary branches), 2(1,1-2),

3(2,1-2), 4(3,2-5), 5(3,2-4), 6(1), 7(2,2-3), 9(1). Segment II: hair 0(1), 1(3), 2

(2,1-2), 3(1), 4(3,2-4), 5(1), 6(1), 7(2,1-2), 9(1). Segment III: hair 0(1), 1(4,3-5),

42 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

2(1), 3(2,1-2), 4(1,1-2), 5(1,1-2), 6(1), 7(2,1-3), 8(4,4-6), 9(1), 10(2), 11(¢1,1-2),

14(1). Segment IV: hair 0(1), 1(2,2-4), 2(1), 3(3,2-3), 4(1,1-2), 5(1), 6(1), 7@,

1-3), 8(2,1-3), 9(1), 10(2), 11(1), 14(1). Segment V: hair 0(1), 1(2,2-3), 2(1),

3(2,1-2), 4(3,2-4), 5(1), 6(1), 7(4,2-4), 8(2,1-3), 9(1), 10(2,1-2), 11(€1), 14(1).

Segment VI: hair 0(1), 1(2,2-4), 2(1), 3(2,2-3), 4(3,2-4), 5(1), 6(1), 7(2,1-2), 8

(2,1-2), 9(1), 10(2,1-2), 11€1), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(2,1-

2), 4(2), 5(1,1-2), 6(1,1-2), 7(1,1-2), 8(2,1-2), 9(3,2-4), 10(2,1-2), 11¢1), 14(1).

Segment VIII: hair 0(1), 4(2), 9(1), 14(1). Paddle: Width about 0.70 of length;

apex more or less rounded; hair 1-P usually as long as paddle.

FOURTH INSTAR LARVA (fig. 24). Head 1.05 mm. Siphon 1.14 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 8 reared specimens.

Head: integument yellowish to very light brown. Mental plate triangular, wider

than long, basolateral spicules sometimes denticulate, all others sharply pointed.

Hair 0(1), 1(1), 2(1), 3(not detectable), 4(3,3-5), 5(4,2-5), 6(1,1-2), 7(9,7-11),

8(3,2-5), 9(3,3-5), 10(2,2-4), 11(4,2-4), 12(2,2-3), 13(4,2-4), 14(1,1-2), 15(2,2-3).

Antenna: Length about 0.33 of head; shaft with a few minute spicules on proxi-

mal part. Hair 1(4,4-7): Thorax: Prothorax: hair 0(5,5-8), 1(1), 2(1), 3(2), 4(3,

2-3), 5(1), 6(1), 7(4,2-4), 8(2), 9(1), 10(€1), 11(4,2-5), 12(1), 14(1). Mesothorax:

11), 202,13), 31), 43,2-4), 5C1), 6(1),. 70D,-807,6-3), 9(6,9-7);: 100K), -TH4.2-

4), 12(1), 13(11,8-11), 14(9,8-10). Metathorax: hair 1(1), 2(2,2-4), 3(5,3-5), 4(4,

2-5), 5(1), 6(2,1-3), 7(7,7-9), 8(7), 9(7,5-8), 10(1), 11(€2,2-3), 12(1,1-2), 13(4,3-

5). Abdomen: Segment I: hair 1(1), 2(1), 3(2,2-3), 4(6,4-7), 5(3,2-4), 6(2,2-3),

7(2,2-3), 9(4,3-4), 10(4,3-4), 11(3,2-4), 12(3,2-3), 13(1). Segment II: hair 0(1),

1(1), 2(1), 3(2,2-3), 4(4,3-4), 5(4,3-5), 6(2), 7(3,2-4), 8(3,2-4), 9(1), 10(1), 11

(2,2-3), 12(3,3-4), 13(6,5-8). Segment III: hair 0(1), 1(3,2-3), 2(1), 3(2,2-3), 4

(2,1-2), 5(3,2-4), 6(2), 7(4,4-5), 8(2), 9(1), 10(2), 11(2), 12(2,2-3), 13(1), 14(1).

Segment IV: hair 0(1), 1(1,1-2), 2(1), 3(4,3-5), 4(1,1-2), 5(2,1-2), 6(2,2-3), 7(5,

4-5), 8(1), 9(1), 10(2), 1102), 12(2,1-2), 13(2), 14(1). Segment V: hair O(1), 1

(121-2), 201), 301), 404.45), 9( 251-2), 6(2); 704545), 81), OC), TOC), Ti Ee

(1), 13(2), 14(1). Segment VI: hair 0(1), 1(3,2-4), 2(1), 3(1), 4(4,3-4), 5(2,1-2),

6(2,1-2), 7(3,2-3), 8(2), 9(1), 10(1), 11(2), 12(1), 13(8,8-9), 14(1). Segment VII:

hair O(1), 1(2,1-2), 2(1), 3(2,2-3), 4(1), 5(2), 6(6,4-6), 7(1), 8(5,3-5), 9(1,1-3),

10(1), 11€1), 12(1), 13(2), 14(1). Segment VIIT: Comb scales similar to those of

dyari. Hair 0(1), 1(6,5-7), 2(1), 3(6,5-7), 4(1), 5(3,3-4), 14(1,1-2). Siphon: Inte-

gument medium brown. Index about 4.7-5.3. Pecten teeth 5(4-6), usually bifid,

sometimes trifid, both blades very slender; distal teeth generally slightly longer.

Hair 1(3,2-3), hair lad(2), hair lav(2,1-3); hairs 2,6,7,8,9 all single. Anal Segment:

Gill about 0.6 of dorsal saddle length. Hair 1-X shorter than gill. Hair 1(3,2-5),

2(7,5-7), 3(1), 4a(12,10-13), 4b(9,8-11), 4c(10,9-11), 4d(11,10-12), 4e(14,10-14),

4f(11,9-11).

SYSTEMATICS. Deinocerites nicoyae clearly belongs to a different phylad than

the other 2 species of the group. This is particularly evident in the male genitalia

(shape of paramere; spiniform setae of ventromesal surface of subapical lobe; [X

tergite lobe) but is also suggested in the larva (reduction of ventral brush and

branching of hairs 6-III,[V). It seems likely therefore that nicoyae represents the

earliest offshoot of the group.

I have tentatively assigned to nicoyae a single female collected in a light trap

at Puerto Somoza, Nicaragua. It is possible that this specimen could represent a

distinct species but this cannot be determined without males and immature stages.

BIONOMICS. Deinocerites nicoyae has been collected in mangrove areas most

Adames: Genus Deinocerites 43

frequently in the burrows of the Wide Red Land Crab, Ucides occidentalis. The

immature stages were found on 1 occasion only, when the water level in these bur-

rows was considerably depressed. Occasionally associated with nicoyae was pseudes,

which more frequently utilizes the burrows of Cardisoma crassum.

DISTRIBUTION (fig. 4). Pacific coast from Puerto Somoza, Nicaragua, to the

mouth of the Gulf of Nicoya in Costa Rica. Material examined: 422 specimens;

151 males, 93 females, 167 larvae, 11 pupae; 11 individual rearings (6 larval, 3

pupal, 2 incomplete). :

COSTA RICA. Puntarenas: Boca del Rio Barranca, Estero El Mero (CR 238,420,254; LCBA-

188) [BM, LACM, UCLA, USNM]. La Angostura (CR 3,27) [UCLA]. .

NICARAGUA: Leon: Puerto Somoza (NI 20) [UCLA].

CANCER GROUP

FEMALES. Medium-sized species, wing 2.78-3.01 mm. Mesonotum dark; pleu-

ral integument whitish to dark, distinctly, slightly or not contrasting with meso-

notal integument. Antenna: Flagellar segment 1 with scales, about equal to com-

bined length of segments 2-4; segments 2-12 subequal in length but progressively —

slightly shortened distad; moderately long, exceeding proboscis from at least basal

half of flagellar segment 9. Thorax: Postnotum without setae. Apn whitish to dark

brown; ppn ‘usually whitish to dark brown, scales all narrow; ppl whitish to dark

brown; psp, ssp, stp and pra slightly (usually) or markedly darker than rest of

pleuron; paratergite either light brown or dark brown; mep whitish to dark brown

and without translucent scales, Imep with 1 very strong bristle; meron, metameron

and metapleuron whitish to light brown. Legs: Coxal integument whitish to brown-

ish. Basal two-thirds of posteroventral surface of all femora pale; anteroventral

margin of forefemur without spiniforms but with 6-8 strong bristles on distal two-

thirds, posterodorsal margin with a rwo of weak bristles. Abdomen: Sternites II-

IV paler than tergites.

FEMALE GENITALIA. Sternite VIII with sclerotized distal band broad and with-

out scales; caudal margin produced into a distinct submedian ventral angle, with

several very strong marginal bristles. Tergite [IX without setae. Tergite X repre-

sented dorsally by a poorly sclerotized cuplike area and laterally by a broad strong

sclerite at base of cercus.Cercus with 2 long apical or subapical spatulate setae.

MALES. Essentially similar to females. Antenna: Exceeding proboscis from at

least base of flagellar segment 5; flagellar segments 1-4 with scales; segments 1-6

markedly elongated but progressively shorter distad; segment 1 always shorter than

combined length of segments 2 and 3; segments 7-12 subequal but progressively

slightly shortened distad; segment 13 either shorter, equal or slightly longer, but

always slightly broader, than 12. Legs: Claws of foreleg and midleg enlarged; an-

terior member of a pair larger and with a variable subbasal to almost basal tooth,

posterior with or without subbasal tooth.

MALE GENITALIA. Segment IX: Tergite lobe not markedly angled laterad at

base; distal part strongly flattened, widened; apex sometimes exceeding the sub-

apical lobe. Sidepiece: Without scales. Apicosternal lobe prominent, with a long

apical seta. Phallosome: Dorsal parameres connected by a variable, but well dif-

ferentiated dorsal sclerotized bridge; articulated ventrally with a dorsal sclerotized

projection from basal third of aedeagus.

PUPAE. Cephalothorax: Hairs 2,3-C closely approximated, 2-C weaker and short-

er than 3-C; hair 5-C double, strongly developed, longer than distance from its

ee. Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

alveolus to base of trumpet, about 1.3-1.5 of trumpet length; 7-C usually double

or triple (2-5); hair 8-C usually with 2-5 branches (1-6). Metanotum: Hair 10-C

moderately to strongly developed, always longer than 11-C, branching varied (1-

7). Abdomen: Hair 1-II not reaching apex of tergite III, multibranched (6-30),

sometimes dendritic and barbed; 1-III-VI branched; 5-II always laterad of 3-II; hair

3-V usually double (1-4); hair 1-VII shorter or sometimes equal to tergite VIII,

never exceeding basal third of 4-VIII. Paddle: Hair 1-P longer than paddle.

FOURTH INSTAR LARVAE. Head: Hair 2-C varied in length from 0.5 to long-

er than 1-C, always markedly mesad of 1-C; hair 5-C usually with 4 barbed branches

(2-5); hair 6-C single and barbed, about 1.33-1.50 of 5-C. Thorax: Prothoracic

hair 9-P single. Abdomen: Hair 6-II double; dorsal sensillum of segment V laterad

of 4-V; hair 6-VI single; 1-VII moderately developed, longer than 3-VII, exceed-

ing base of segment VIII but not reaching base of siphon; 1-VIII usually with

3 or 4 branches (2-5). Siphon: Hair 1-S usually double (1-5). Anal Segment: Ven-

tral brush (4-X) with 6 or 7 pairs of hairs. Gill slightly emarginate on apex; short,

about 0.45-0.60 of dorsal saddle length. Dorsal saddle narrow, its ventral margin

far from hair 1-X.

DISCUSSION. The Cancer Group is well marked by unique features in both

male ([X tergite lobe) and female (cercal setae) genitalia. The immature stages

of this group are very similar to those of the Pseudes Group from which, how-

ever, they can usually be separated by the characters given in the keys.

On the basis of morphological features of the adults (length of antenna) and

larvae (ventral brush), the clearly marked magnus appears to be the representa-

tive of the primitive stock of the group. The other 2 species of the group, cancer

and melanophylum, are obviously closely related and show derived states in these

_ characters.

The group is apparently restricted to the Atlantic basin and its distribution is

centered around the Caribbean Sea with extensions northward into the Bahamas

and Florida and southward to the state of Maranhao in Brazil. The 3 species have

complementary allopatric distributions with a possible gap in eastern Venezuela:

cancer in Central America down to at least Almirante, Panama, in the western

Antilles through Hispaniola, and in the Bahamas and southern Florida; melano-

phylum from central Panama through Colombia to central Venezuela; and magnus

in the eastern Antilles from Mona and Puerto Rico through the Lesser Antilles,

Trinidad and Tobago, the Guianas and in Brazil south of the Amazon. Over much

of the group’s distribution (West Indies, Bahamas, Florida, eastern South America)

either cancer or magnus is the only representative of the genus.

The species of the group are usually common in areas where no other species

of the genus are present. Considerable information has been accumulated on the

bionomics of cancer in recent years but little is known about the other species.

All 3 species may occasionally bite man but in Panama cancer appears to have a

preference for avian blood and melanophylum for reptilian blood. The immature

stages of all 3 species are usually found in large crabholes, probably largely those

made by Cardisoma guanhumi.

8. Deinocerites cancer Theobald

Figs. 1,5,25-28

1901. Deinocerites cancer Theobald, 1901b:215-216,356. TYPE: Lectotype female, Spanish

Adames: Genus Deinocerites 45

Town Road, Kingston, Jamaica, 8 Feb 1900, M. Grabham [BM; designation of Belkin

and Hogue, 1959:432].

1909. Deinocerites tetraspathus Dyar and Knab, 1909:260. TYPE: Lectotype female with gen-

italia slide (472), Bluefields, Nicaragua [USNM, 12109; designation of Stone and Knight,

1957:197]. Synonymy with cancer by Dyar (1928:262).

Deinocerites cancer of Pazos (1904:135); Dyar (1905b:24; 1905c:27; 1905d:53; 1905e:109;

1922:8); Felt (1905:491,492); Coquillett (1906:26; 1910:531); Mitchell (1906:19,20; 1907a:

13; 1907b:264); Knab (1907a:121); Dyar and Knab (1918:183); Johnson (1919:422); Gowdey

(1926:73); Gerry (1932:43,44,71); King, Bradley and McNeel (1944:4,61,68,79,81); Middle-

kauff and Carpenter (1944:89); Roth and Young (1944:84); Pratt, Wirth and Denning (1945:

426); Wirth (1945:200); Carpenter and Chamberlain (1946:88); Carpenter, Middlekauff and

Chamberlain (1946:275-277); Pritchard (1947:13); Thompson (1947:78); Hill and Hill (1945:

2; 1948:51-52); Thurman, Haeger and Mulrennan (1949:171); de la Torre, Alayo and Cal-

deron (1956:80); Pratt (1956:8); Branch, Logan, Beck and Mulrennan (1958:161,162); Belkin

and Hogue (1959:432-434); Branch and Seabrook (1959:216); Evans (1962:255); Dodge (1963:

798,811; 1966:375); Ross (1964:104); Porter (1964:222); Forattini (1965:108); Downes

(1966:1169-1177); Montchadsky and Garcia Avila (1966:47); Provost and Haeger (1967:565-

574); Gentry, Gerberg and Hopkins (1970:68-70); Gorgas Memorial Institute (1970:20); Tem-

pelis and Galindo (1970). |

Deinocerites cancer in part of Theobald (1903:276-281; 1905a:35-36; 1905b:37; 1910:553);

Dyar and Knab (1906:188); Surcouf and Gonzalez-Rincones (1911:91); Howard, Dyar and

Knab (1915:201-205); Bonne and Bonne-Wepster (1925:173-174); Dyar (1928:262-263); Ed-

wards (1932:222); Matheson (1944:250-251); Lane (1953:554-556); van der Kuyp (1953b:

144,146); Carpenter and LaCasse (1955:325-327); Horsfall (1955:599-600); Perez Vigueras

(1956:476-483); Forattini (1958:175); Stone, Knight and Starcke (1959:284).

?2Deinocerites cancer of Martini (1935:57); Vargas (1956:30).

2Deinocerites cancer in part of Knab (1907:95-97).

Dinocerites cancer in part of Blanchard (1905:414).

Deinokerides cancer in part of Giles (1902:472-473).

Culex (Deinocerites) cancer of Dyar (1918:102). 3

Deinocerites tetraspathus of Howard, Dyar and Knab (1915:209-210); Bonne and Bonne-Wepster

(1925:175).

Deinocerites melanophylum in part of Dyar (1925:155).

Brachiomyia magna in part of Giles (1902:474).

FEMALE (figs. 1,25,26). Wing 2.88 mm. Proboscis 2.09 mm. Forefemur 1.73

mm. Abdomen about 3.11 mm. Mesonotum dark brown, pleural integument slight-

ly contrasting with mesonotal integument or concolorous. Head: Narrow decum-

bent scales of vertex creamy to brownish; erect scales brown to dark brown; lat-

eral patch of broad decumbent scales whitish. Antenna: Torus occasionally with

1 scale; variable in length, exceeding proboscis from at least basal half of flagellar

segment 9 to apex of segment 8. Thorax: Apn pale brown to dark brown; ppn

pale brown to dark brown, usually with several scales in middle and lower part;

ppl pale brown to brown, with several bristles and scales; psp, ssp, stp and pra

brown to dark brown; paratergite brown to dark; mep brown to dark brown, up-

per part with several bristles, microsetae absent; metameron simple; meron, meta-

meron and metapleuron very pale brown to medium brown. Legs: Coxal integu-

ment very pale brown to medium brown; anterior surface of forecoxa covered

with bristles and translucent scales, basolateral area with a few bristles; midcoxa

with scales anterolaterally, with 2 rows of bristles laterally, the anterior bristles

more irregular in order, the posterior ones stronger, posterolateral surface simple;

hindcoxa with scales and several bristles laterally, posterior surface with bristles.

Haltere: Stem with dorsoapical part with several bristles.

46 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

FEMALE GENITALIA (fig. 26). Sternite VUIT with differentiated marginal bris-

tles largely restricted to sternal margin, at most continued dorsad on one-third

of caudolateral border. Cercus (in lateral aspect) with sternal margin more or less

convex but slightly sinuous subapically, tergal margin more or less straight but

slightly downturned apically; apex downturned; the 2 specialized setae inserted

side by side on apex and subequal in length; tergal and sternal surfaces each with

a moderately strong normal bristle, the sternal shorter and closer to apex. Distal

part of postgenital plate (in ventral aspect) with width not more than 1.8 of length,

usually with notch as deep as wide; lateral lobe short and more or less rounded

(in lateral aspect), its apical bristle subequal to total length of lobe and displaced

mesad and proximad of apex, convergent with its mate (in ventral aspect), its

small dorsal and subapical bristles appearing almost apical.

MALE (fig. 25,26). Wing 3.34 mm. Proboscis 2.60 mm. Forefemur 2.40 mm.

Abdomen (not including genitalia) about 3.0 mm. Flagellar segment 4 with very

few scales, segment 13 subequal to 12. Proboscis extending to distal third of fla-

gellar segment 4. Anterior claws of foreleg and midleg similar, with a large sub-

basal tooth.

MALE GENITALIA (fig. 27). Segment IX: Tergite lobe extending beyond sub-

apical lobe, with the external and internal margins rather evenly curved, distal

part very broad. Sidepiece: Subapical lobe without a distinct thumb; seta c rather

heavy, spiniform and with apex bent and attenuated. Phallosome: Dorsal para-

meres closely approximated tergally and with a broad dorsal, almost complete

bridge; apical spine slender, moderately curved; ventral teeth slender, long and aris-

ing from a distinct process from the convex caudolateral border, two or more

teeth often with a common base, several additional smaller denticles removed from

border. Aedeagus (in tergal aspect) with outer margins sinuous, expanded before

middle, constricted near middle and expanded subapically, apex more or less round-

ed.

PUPA (fig. 27). Abdomen 3.64 mm. Trumpet 0.53 mm. Paddle 0.76 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 10 reared specimens.

Also differing from melanophylum in hair 10-C usually being double or triple (1-

7) instead of usually with 4-6 branches (2-8). Cephalothorax: Integument yellow-

ish. Hair 1(2,2-3), 2(4,3-5), 3(2), 4(3,2-4), 5(2), 6(1), 7(2), 8(2,2-4), 9(2,1-3), 10

(2), 11(2,1-3), 12(3,2-5). Trumpet: Short, slightly widening apically; index about

4.1-5.1. Integument brown distad, slightly darker on tracheoid; contrasting with

cephalothoracic integument. Tracheoid about 0.36 of trumpet length. Abdomen:

Integument light to medium brown. Segment I: hair 1(16,12-19), 2(1,1-2), 3(2,

1-2), 4(4,3-5), 5(5,3-7), 6(2,1-2), 7(3,2-4), 9(1,1-3). Segment II: hair 0(1), 1(7,

5-8), 2(1,1-2), 3(1), 4(3,3-5), 5(1,1-2), 6(1), 7(2,1-2), 9(1). Segment III: hair 0

(1), 1(4,3-6), 2(1), 3(2,2-3), 4(2,1-2), 5(1,1-2), 6(1), 7(2,2-3), 8(3,2-5), 9(1), 10

(2,1-3), 11(1), 14(1). Segment IV: hair 0(1), 1(3,2-6), 2(1), 3(6,4-7), 4(1,1-2),

S(1), 6(1), 7(2,1-3), 8(2,1-3), 9(1), 10(2,2-3), 11(1), 14(1). Segment V: hair 0

(1), 1(3,1-3), 2(1), 3(2,1-2), 4(3,3-5), 5(1), 6(1), 7(3,1-4), 8(3,2-4), 9(1), 10(1,

1-2), 11(1), 14(1). Segment VI: hair 0(1), 1(3,2-4), 2(1), 3(2,1-3), 4(2,2-3), 5(1),

6(1), 7(1,1-2), 8(2,2-3), 9€1), 10(2,1-3), 11(1), 14(1). Segment VII: hair 0(1), 1

(1), 21), 3(2), 4(2,1-4), 5(2,1-3), 6(1), 7(1), 8(2,2-3), 9(3,2-5), 10(2,1-3), 11(1),

14(1). Segment VIII: hair 0(1), 4(2), 9(1), 14(1). Paddle: Width about 0.71-0.81

of length, apex more or less rounded.

FOURTH INSTAR LARVA (fig. 28). Head 1.16 mm. Siphon 1.25 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 10 reared specimens.

Adames: Genus Deinocerites 47

Head: Integument yellowish to light brown. Mental plate wider than long, triang-

ular, lateral spicules denticulate, terminal ones sharply pointed, median terminal

conspicuously produced. Hair 0(1), 1(1), 2(1), 3(sometimes developed as a minute

spicule), 4(4,2-6), 5(4,3-5), 6(1), 7(8,6-8), 8(3,3-4), 9(5,3-6), 10(2,2-3), 11(5,2-

5), 12(2,2-4), 13(5,3-6), 14(1), 15(2,1-3). Antenna: Length about 0.41 of head;

shaft with a few spicules on proximal part. Hair 1(5,4-6). Thorax: Prothorax: hair

0(8,4-10), 1(1), 2(1), 3(1,1-2), 4(3,2-3), S(1), 6(1), 7(3,2-3), 8(1), 9C1), 10(1),

11(3,2-5), 12(1), 14(1). Mesothorax: hair 1(1), 2(2,2-3), 3(1), 4(3,2-3), 5(1), 6(1),

7(1), 8(6,5-7), 9(7,5-9), 10(1), 11(3,2-4), 12(1), 13(8,8-12), 14(8,6-10). Metatho-

rax: hair 1(1), 2(3,1-4), 3(4,3-6), 4(4,3-6), 5(1), 6(2,1-2), 7(8,6-9), 8(6,6-8), 9(7,

5-9), 10(1), 11(2,2-3), 12(1), 13(4,3-5). Abdomen: Segment I: hair 1(1), 2(1),

3(3,2-3), 4(6,4-8), 5(4,2-4), 6(2), 7(2), 9(2,1-3), 10(2,1-3), 11(2,2-4), 12(2,2-3),

13(2,1-2). Segment II: hair 0(1), 1(1), 2(1), 3(3,2-3), 4(5,4-7), 5(3,2-3), 6(2,2-

3), 7(5,4-6), 8(3,2-4), 9(1,1-2), 10(€1), 11(3,2-3), 12(3,2-3), 13(8,6-11). Segment

III: hair O(1), 1(3,2-4), 2(1), 3(€1,1-2), 4(2,1-4), 5(2,2-3), 6(2), 7(6,4-6), 8(2,1-3),

9(1), 10(1), 11(2,2-3), 12(1), 13(1,1-2), 14(1). Segment IV: hair O(1), 1(€1), 2

(1), 3(3,3-4), 4(1,1-2), 5(2,1-3), 6(2), 7(4,4-7), 8(1), 9(1,1-2), 10(1), 11(2,2-3),

12(2), 13(2), 14(1). Segment V: hair 0(1), 1(1), 2(1), 3(1), 4(5,4-7), 5(2,1-3), 6(2),

7(6,4-6), 8(1), 9(1,1-2), 10(1), 11€1,1-2), 12(2,1-2), 13(2), 14(1). Segment VI: hair

O(1), 1(3,2-3), 21), 3(1), 4€3,24), 5(2,2-4), 6(1), 703,24), 8(2,2-4), 9(1), 10(1),

11(2), 12(1,1-2), 13(12,8-14), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(2,2-4),

4(1), 5(3,2-3), 6(8,6-8), 7(1,1-2), 8(5,4-7), 9(1,1-3), 10(1), 11(1), 12(1), 13(@2,1-

3), 14(1). Segment VIIT: Comb scales slender at base, wider apically, fringe re-

stricted to apex. Hair 0(1), 1(4,3-4), 2(1), 3(4,3-5), 4(1), 5(3,2-3), 14(€1,1-2). Si-

phon: integument light brown. Index about 4.4-5.7. Pecten teeth 5(5-7), usually

bifid, sometimes trifid, longest blade wider; basal tooth shorter, others progres-

sively longer apically. Hair 1(2,2-3), hair lad(2), hair lav(1,1-2), hairs 2,6,7,8 and

9 all single. Anal Segment: Gill about 0.60 of dorsal saddle length. Hair 1-X longer

than gill. Hair 1(1), 2(6,5-7), 3(1), 4a(11,11-13), 4b(10,9-11), 4c(9,8-10), 4d(9,

8-11), 4e(11,10-13), 4f(11,6-13).

SYSTEMATICS. Deinocerites cancer appears to be the dominant and most wide-

spread derivative of the group. It is very similar to melanophylum, particularly

in the female and larva. However it can usually be separated from melanophylum

in all stages by the key characters, except as noted under melanophylum for some

Venezuelan larvae.

I have found no significant differences between the Antillean and Central Amer-

ican populations of cancer and I am convinced that only 1 species is involved.

Such a pattern of distribution is known for a number of other mosquitoes (Belkin,

Heinemann and Page, 1970:9, Middle American element) and suggests that the

present large water gaps between these areas are relatively recent (Belkin and Hogue,

1959:430).

BIONOMICS. Cardisoma guanhumi appears to be the only species of crab def-

initely identified as the maker of the burrows utilized by the immature stages

of cancer (Pratt, Wirth and Denning, 1945:246; Haeger and Phinizee, 1959:34).

In Cuba, the immatures of cancer have been reported from burrows of Cardisoma

and unidentified species of stream inhabiting crabs of the genus Epilobocera (fam-

ily Pseudothelphusidae) by Montchadsky and Garcia (1966:47). Belkin, Heinemann

and Page (1970:49) state that in Jamaica cancer has been collected only in crab-

holes of medium to large size and never in smaller ones like those made by fiddler

crabs. The immature stages of cancer have been reported several times from out-

48 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

side their normal breeding sites: from rockholes by Wirth (1945:200) and by Bel-

kin, Heinemann and Page (1970:49); from a treehole by Porter (1964:222); from

a tin can (T.E. Duffey record in Carpenter, Middlekauff and Chamberlain, 1946:

277); from a wooden bucket, an abandoned septic tank and a tin can (Porter,

1964:222, records of F.H. Stutz and J.H. Heidt); and from flooded areas with

crabholes by several workers. | :

The normal associates of the immature stages of cancer are other normally ob-

ligate breeders in crabholes that occur sympatrically: Culex (C.) janitor and Culex

(Mel.) carcinophilus in the West Indies; D. epitedeus, Culex (Tin.) latisquama and

1 or more species of the Culex (C.) inflictus complex in Central America and

Panama. Other species occasionally found associated with cancer in unusual breed-

ing sites, including shallow or flooded crabholes, are: Aedes (O.) taeniorhynchus,

Anopheles (A.) grabhamii, Anopheles (Nys.) albimanus, Culex (C.) nigripalpus, Cu-

lex (C.) bahamensis, Culex (Mel.) elevator complex, Culex (Mel.) opisthopus and

Psorophora (G.) jamaicensis in various ground pools (Belkin, Heinemann and Page,

1970:49; Pratt, Wirth and Denning, 1945:246; Wirth, 1945:200); Aedes (F.) tri-

seriatus complex and Culex (C.) nigripalpus in treeholes or containers (Carpenter,

Middlekauff and Chamberlain, 1946:277; Porter: 1964:222).

The bionomics of cancer have been studied by Haeger and Phinizee (1959) in

an autogenous colony from the Florida Keys and by Gentry, Gerberg and Hopkins

(1970) in Baltimore in a colony established from material collected on Grand Cay-

man Island. According to the latter authors larval development requires 3 to 4

weeks at 27° C while Haeger and Phinizee (1955:35) state that it took about 2

to 3 weeks.

Howard, Dyar and Knab (1915:205) attribute to M. Grabham the statement

that cancer feeds only at night and is ‘‘a voracious bloodsucker’’. Hill and Hill

(1948:52) dealing with the same topotypic Jamaican population of cancer report

that: ’while they probably feed on crabs, we have caught them full of blood in

animal-baited traps and occasionally they will bite man.” Forattini (1965:108) does

not specify the source of his statement that cancer attacks animals such as horses

and man. Gentry, Gerberg and Hopkins (1970:79) report feeding females on chick-

ens and indicate that although a bared forearm was offered on numerous occa-

sions, bites were experienced only 4 times; no full engorgement was noted either

on chicken or human hosts. Haeger and Phinizee (1959:37) found that in their

colony females would not feed on man or animals until a batch of autogenous

eggs was laid. After that a few would bite man and would also feed to a limited

extent on both land crab and salt water blue crab haemolymph to produce a sec-

ond batch of eggs. Studies on the blood feeding habits of cancer in Almirante,

Panama, have shown that this population has a definite preference for avian blood

(Gorgas Memorial Institute, 1970:20; Tempelis and Galindo, 1970).

The unusual mating behavior of cancer has been studied in detail by Downes

(1966) and by Provost and Haeger (1967). The males exhibit a so-called “pupal

attendance behavior” during which they walk on the surface of the water with

the antennae directed down to the surface. When a female pupa is encountered

its trumpets are touched by the tips of the antennae and it is seized and held

by the claws of the male. Copulation takes place before the female is completely

free of the pupal skin. More than 1 male may attempt to copulate with 1 female

and some females may emerge unattended but are soon contacted by 1 or more

males.

DISTRIBUTION (fig. 5). From Florida and the Bahamas through Cuba and His-

Adames: Genus Deinocerites _ 49

paniola to the Atlantic coast of Central America from the Yucatan Peninsula in

Mexico to the Bocas del Toro Province in Panama. Material examined: 5773 spec-

imens; 668 males, 1294 females, 2868 larvae, uae pupae; 635 individual rearings.

(250 larval, 333 pupal, 52 incomplete).

BAHAMA ISLANDS. Abaco: Marsh Harbor, G.M. Stokes [USNM].

BRITISH HONDURAS. Belize, D.S. Bertram [LOND, UCLA]; [USNM]. Trapp’s Key, W.H.

Sligh [USNM].

CAYMAN ISLANDS. Grand Cayman (CAY 10A,11A,12,13,19,127) [UCLA].

COSTA RICA. Limon: Cahuita (LCBA 233,234,237,238,240-243) [LACM, UCLA].

CUBA. Guantanamo: U.S. Naval base, K.L. Knight (461,462) [UCLA, USNM]. La Habana:

Habana, J.R. Taylor [USNM]. Marianao, H.P. Carr [UCLA, USNM]. Pinar del Rio: Mariel, J.R.

Taylor [USNM].

DOMINICAN REPUBLIC. Azua: Azua, P.A. Ricart [USNM]. Montecristi: Montecristi, P.A.

Ricart [UCLA, USNM]. San Pedro de Macoris: San Pedro de Macoris, P.A. Ricart [USNM].

Santo Domingo: Santo Domingo, A. Busck (1136,580113-11); F.E. Campbell (10.1,10.2,10.4)

[UCLA, USNM].

FLORIDA. Beach County: Boca Raton, Wanamaker [USNM]; [UCLA, USNM]. Broward

County: Ft. Lauderdale, W.W. Wirth and D.G. Denning [UCLA, USNM]. Hollywood [USNM].

County not specified: (1173,1282,1302,1719-8,1946,1990) [USNM] . Dade County: Coral Gables

[USNM]. Miami, G.F. Moznette [UCLA, USNM] ; [USNM]; W.W. Wirth [USNM]. Miami Beach

W.W. Wirth [USNM]. Martin County: Jupiter Island, Wanamaker [USNM]. Monroe County:

Key West Banks [USNM]. St. Lucie County: Ft. Pierce [USNM]; N.G. Piatte [USNM].

GUATEMALA. Izabel: Puerto Matias (GUA 5) [UCLA].

HAITI. Department not specified: W.H.W. Komp, leg (No. T.5) [USNM];S.S. Cook [USNM].

Ouest: Gode (HAT 12) [UCLA]. Gressier (HAT 3,4) [UCLA]. Riviere Froide (HAT 1) [UCLA].

Sud: Les Cayes (HAT 14) [UCLA].

HONDURAS. Puerto Castilla, R.X. Maxwell (100) [UCLA, USNM]; (HON 8,9) [UCLA].

Puerto Cortes (HON 62,66,69,71-75,77,79,81-84,86,87,89,91) [UCLA]. Tela, W.H.W. Komp

[USNM].

JAMAICA. Near the coast all around the island. For specific localities see Belkin, Heinemann

and Page (1970:50).

MEXICO. Cozumel, Isla de: Espiritu Santo Bay [USNM].

NICARAGUA. Zelaya: Bluefields (NI 34,51) [UCLA].

PANAMA. Bocas del Toro: Almirante (PA 1033, GML 03690) [UCLA].

SAN ANDRES, ISLA DE (Colombia). (ISA 209,211,214) [UCLA].

9. Deinocerites melanophylum Dyar & Knab

Figs. 5,29-31

1907. Deinocerites cancer melanophylum Dyar and Knab, 1907b:200. TYPE: Lectotype male,

Colon, Panama, A. Busck, 213 [USNM, 10865; designation of Stone and Knight, 1957:

1977:

1925. Deinocerites monospathus Dyar, 1925:155. TYPE: Holotype female, Fort Sherman,

Panama, Canal Zone, 24 April 1925, D. Baker; unique (mounted on same slide with

normal female of melanophylum) [USNM, 12128]. Synonymy with melanophylum by

Belkin and Hogue (1959:434).

Deinocerites melanophylum of Busck (1908:60-61); Howard, Dyar and Knab (1915:207-209);

Dyar (1923:179); Stone and Knight (1957:197); Belkin and Hogue (1959:434-435); Stone,

Knight and Starcke (1959:285); Downes (1966:1173); Gorgas Memorial Institute (1970:20);

Tempelis and Galindo (1970).

Deinocerites melanophylum in part of Dyar (1925:155).

50 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

2Deinocerites melanophylum of van der Kuyp (1953a:38). :

2Deinocerites melanophylum in part of Bonne and Bonne-Wepster (1925:17 5): van der Kuyp:

(1948b:895-897; 1953b:144,146; 1954:39,46,56,57,64,67,71,72,73,76,82,84 86, 88,90,95,96,

119).

Culex (Deinocerites) melanophylum of Dyar (1918:102).

Deinocerites monospathus of Dyar (1928:263); Horsfall (1955:599); Stone and Knight (1957:

197).

Deinocerites cancer of Anduze (1941:17); Anduze, Pifano and Vogelsang (1947 16); Arnett

(1950:107); Carpenter and Peyton (1952:677,681); Komp (1956:349-351).

Deinocerites cancer in part of Lane (1953:554-556); Carpenter and LaCasse (1955: 327); Horsfall

(1955:599-600); Forattini (1958:175); Stone, Knight and Starcke (1959:284).

2Deinocerites pseudes of Anduze (1941:17); Anduze, Pifano and Vogelsang (1947:16).

FEMALE (fig. 29). Wing 3.01 mm. Proboscis 1.98 mm. Forefemur 1.73 mm.

Abdomen about 2.70 mm. Essentially similar to cancer except for the antenna

exceeding the proboscis from base of antennal flagellar segment 8. —

FEMALE GENITALIA (fig. 29). Essentially as in cancer except for the follow-

ing. Distal part of postgenital plate (in ventral aspect) with width always more

than 2.0 of length, notch often poorly differentiated, anterior border with a dis-

tinct median cephalic sclerotized projection; lateral lobe poorly differentiated, us-

ually appearing only as a small rounded protuberance.

MALE (fig. 29). Wing 3.0 mm. Proboscis 2.39 mm. Forefemur 1. 63 mm. Ab-

domen (not including genitalia) about 2.35 mm. Flagellar segment 4 with several

scales, segment 13 slightly longer than 12. Proboscis extending to distal one-third

of flagellar segment 4. Anterior claw of foreleg with a small subbasal tooth, that

of midleg with a longer and heavier tooth.

MALE GENITALIA (fig. 30). Segment IX: Tergite lobe essentially as in cancer

except not quite as broad apically and with more sinuous margins. Sidepiece: Es-

sentially as in cancer except that seta c usually less attenuated apically. Phallo-

some: Dorsal parameres widely separated in tergal aspect and with a long narrow

incomplete dorsal bridge; apical spine long, slender and strongly curved; ventral

teeth short and heavy to minute and in several rows, directed parallel to convex

margin so that they appear appressed to body of paramere. Aedeagus in tergal

aspect expanded before middle, constricted near middle then expanded subapi-

cally; apex blunt.

PUPA (fig. 30). Abdomen 3.34 mm. Trumpet 0.45 mm. Paddle 0.68 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 10 reared specimens.

Cephalothorax: Integument yellowish to medium brown. Hair 1(2,2-5), 2(4,3-5),

3(2,2-3), 4(4,3-5), 5(2), 6(1), 7(2,1-2), 8(3,1-6), 9(2,1-3), 10(6,4-7), 11(3,2-5), 12

(2,2-4).. Trumpet: Short, slightly widening apically; index about 3.4-4.3. Integu-

ment brown distad, slightly darker on tracheoid; contrasting with cephalothoracic

integument. Tracheoid about 0.38 of trumpet length. Abdomen: Integument light

to medium brown. Segment I: hair 1(16,12-23, primary branches), 2(1), 3(2,1-3),

4(3,3-5), 5(3,3-7), 6(1), 7(3,2-4), 9(1). Segment II: hair 0(1), 1(11-17), 2(2,1-3),

3(1), 4(4,1-6), 5(1,1-4), 6(1), 7(2,1-3), 9(1). Segment III: hair 0(1), 1(6,3-7), 2

(1), 3(4,Je7),. 4( 1162), SC), 601), Fst), O62 bra), 9), FOL 2); TC1), 4

(1). Segment IV: hair 0(1), 1(3,2-5), 2(1), 3(6,3-7), 4(1), 5(1), 6(1), 7(2,1-3), 8

(1,1-2), 9€1), 10(2,1-3), 11€1), 14(1). Segment V: hair 0(1), 1(2,1-4), 2€1), 3(2,

2-3), 4(3,2-6), 5(1), 6(1), 7(4,2-5), 8(2,1-2), 9(1), 10(€1,1-2), 11(€1), 14(1). Seg-

ment VI: hair 0(1), 1(2,1-3), 2(1), 3(2,1-3), 4(4,3-5), 5(1), 6(1), 7(1,1-2), 8(2,

1-3), 9(1), 10(2,1-2), 11(1), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(2,1-2),

Adames: Genus Deinocerites 51

4(2,1-2), 5(1,1-3), 6(1,1-3), 7(1), 8(2,1-3), 9(4,3-6), 10(2,1-2), 11(1), 14(1). Seg-

ment VIII: hair 0(1), 4(2,1-3), 9(1), 14(1). Paddle: Width about 0.70 of length;

apex more or less rounded.

FOURTH INSTAR LARVA (fig. 31). Head 1.16 mm. Siphon 1.31 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 7 reared specimens.

Head: integument yellowish to light brown. Mental plate wider than long; shape

of spicules variable, apex either simple or denticulate. Hair (1) 5 U4), 24}, °3

(sometimes developed as a spicule), 4(7,6-8), 5(5,4-6), 6(1), 7(10,8-12), 8(4,3-4),

9(4,4-5), 10(3,2-5), 11(5,3-6), 12(3,2-3), 13(4,3-6), 14(1), 15(3,2-4). Antenna:

Length about 0.43 of head; shaft with several spicules on proximal part; hair |

(6,4-9). Thorax: Prothorax: hair 0(9,7-12), 11), 2(1), 3(2,2-3), 4(4,3-5), 5(1), 6

(1), 7(4,2-5), 8(1,1-2), 9(1), 10(1), 11(3,3-4, 12(1), 14(1). Mesothorax: hair 1

(1), 2(3,2-4), 3(1), 4(2,2-3), 5(1), 6(1), 7(1), 8(7,4-8), 9(6,5-8), 10(1), 11(3,2-3),

12(1), 13(?), 14(10,8-12). Metathorax: hair 1(1), 2(4,3-6), 3(6,4-7), 4(5,3-5), 5

(1), 6(1), 7(9,8-10), 8(7-10), 9(6-10), 10(1), 11(4), 12(1), 13(6,5-7). Abdomen:

Segment I: hair 1(1), 2(1), 3(3,2-4), 4(9,7-11), 5(3,2-4), 6(2), 7(2,1-3), 9(4,3-4),

10(2,2-3), 11(3,2-6), 12(3,2-4), 13(1). Segment II: hair 0(1), 1€1), 2(1), 3(3,2-4),

4(7,6-9), 5(4,2-5), 6(2,2-3), 7(4,4-7), 8(3,2-4), 9(1), 10(1), 11(2,2-3), 12(3,2-3),

13(7,7-12). Segment III: hair 0(1), 1(3,1-3), 2(1,1-2), 3(1,1-2), 4(2,1-3), 5(3,2-

4), 6(2), 7(6,6-8), 8(2,2-3), 9(1), 10(1,1-2), 11(2,1-3), 12(3,2-3), 13(1), 14(1).

Segment IV: hair 0(1), 1(2,1-2), 2(1), 3(4,3-5), 4(2,1-3), 5(3,2-4), 6(2), 7(6,6-8),

8(1), 9(1), 10(1), 11(2,2-3), 12(2,1-2), 13(2), 14(1). Segment V: hair 0(1), 1(2,

1-2), 201), 3(1)- 403-9), 53,3-5),, 2), 708,79), 30 1-2), 90D), FOC), TIC),

12(1,1-2), 13(2,2-3), 14(1). Segment VI: hair 0(1), 1(4,3-5), 2(1), 3(1), 4(3,2-4),

5(5,4-5), 6(1), 7(4,3-5), 8(3,2-3), 9(1), 10(1,1-2), 11(2,2-3), 12(1), 13(13-17), 14

(1). Segment VII: hair 0(1), 1(1), 2(1), 3(5,4-5), 4(1), 5(2,2-3), 6(10,9-12), 7(1,

1-2), 8(5,4-6), 9(4,2-4), 10(1), 11(€1), 12(1), 13(2,2-4), 14(1). Segment VIIT: Comb

scales slender, slightly wider near middle, fringe conspicuous at apex, less devel-

oped in distal half. Hair 0(1), 1(4,3-5), 2(1), 3(4,4-5), 4€(1), 5(3,3-4), 14(1). Si-

phon: Integument light brown. Index about 5.3-6.5. Pecten teeth 6(5-8), usually

bifid, occasionally trifid, blades slender, size variable throughout. Hair 1(3,2-4),

hair lad(2,2-3), hair lav(2,2-3); hairs 2,6,7,8 and 9 all single. Anal Segment: Gill

about 0:45 of dorsal saddle length. Hair 1-X longer than gill. Hair 1(2,1-2), 2(9,

6-11), 3(1), 4a(11,11-14), 4b(10,10-12), 4c(10,10-12), 4d(12,11-12), 4e(10,11-13),

4f(11,10-11).

SYSTEMATICS. Deinocerites melanophylum is very similar to cancer in all stages

but it can usually be separated from it by the key characters. In some larvae from

Venezuela (VZ 216-218,257) the diagnostic features of melanophylum break down.

The interpretation that melanophylum is distinct from cancer is supported by

the difference in blood hosts found in recent investigations in Panama (see bio-

nomics). The preference for reptilian hosts by melanophylum suggests also that

this species is the earlier derivative of the complex.

It is not known at present if melanophylum comes in contact with cancer in

Panama and with magnus in Venezuela, as no collections of Deinocerites have been

made in these relatively wide gaps (particularly in Venezuela).

BIONOMICS. Komp’s (1957:350) record of the immature stages of melanophy-

lum (as cancer) from the burrows of a large land crab, probably Cardisoma guan-

humi, on the right bank of the lower Chagres River in the Canal Zone is the only

more or less definite association of this mosquito with a specific crab. Elsewhere

melanophylum has usually been found in large crabholes except in Venezuela where

a2 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

it has been collected also in small crabholes.

In Panama, melanophylum has been found associated with panamensis and with

a member of the Culex (C.) inflictus complex; in Colombia, with colombianus;

and in Venezuela, with atlanticus and a member of the Culex (C.) inflictus complex.

Busck (1908:61) reported this species not to bite man in the Canal Zone but

Carpenter and Peyton (1952:677) report occasional specimens (as cancer) taken

in horse-baited traps in this area. Recent studies at the Gorgas Memorial Labora-

tory indicate that melanophylum has a marked preference for reptilian blood in

Panama (Gorgas Memorial Institute, 1970:20; Tempelis and Galindo, 1970).

According to Busck (1908:61), melanophylum (as cancer) adults come out at

dusk “‘and swarm above the hole for copulation”. However, it seems very likely

that this species exhibits the same pupal attendance and mating behavior as its

very close relative, cancer, as suggested by the observations of Komp (1957:350)

who noted several pairs of newly emerged adults copulating in the container in

which he had placed several pupae (and larvae) the night before.

DISTRIBUTION (fig. 5). Atlantic coast from Canal Zone to state of Aragua

in Venezuela. Two records of melanophylum in Belkin and Hogue (1959:434) are

probably erroneous. The specimen from Bocas del Toro is probably from Panama

and not Colombia and in that case should be cancer; however this cannot be de-

termined because of its poor condition. The record from Maracay, which was in-

terpreted as indicating the presence of this species in Lago Valencia, has not been

confirmed in extensive recent search and is probably based on mislabelled material.

Material examined: 1045 specimens; 170 males, 195 females, 522 larvae, 158 pu-

pae; 139 individual rearings (110 larval, 19 pupal, 10 incomplete).

COLOMBIA. Antioquia: Atrato River delta (COA 30) [UCLA] . Turbo (COA 34-41) [UCLA].

PANAMA AND CANAL ZONE. Canal Zone: Cristobal [UCLA]. Ft. Randolph, J. Zetek;

D. Baker [USNM]. Ft. Sherman, various collectors [UCLA, USNM]. France Field, J. Zetek

[USNM]; W.P. Murdoch [UCLA]. Locality not specified, AH. Jennings (248,352) [USNM];

W.H.W. Komp [UCLA]. Margarita, J.B. Shropshire [USNM]. Mt. Hope, S.J. Carpenter [UCLA].

Toro Pt., J.B. Shropshire [USNM]. Colon: Calderas Island, Portobelo Bay, A-H. Jennings [USNM].

Colon, A. Busck [USNM]; (PA 1012,1014-1016,1029,1032) [UCLA]. San Blas: Cuadi River,

B.F. Eldridge [USNM].

VENEZUELA. Aragua: Cuyagua (VZ 414,415) [UCLA]. Maracay (locality questionable), M.

Nunez-Tovar [USNM]. Ocumare de la Costa, M. Nunez-Tovar [USNM]; (VZ 134,170,173,174,

346,351,363,364,386) [UCLA]. Puerto Colombia (VZ 214,216-218) [UCLA]. Turiamo, W.H.W.

Komp (32-19,22); E.B. Winston [UCLA]. Carabobo: Moron (VZ 256,257) [UCLA] . Puerto Ca-

bello (VZ 260) [UCLA].

10. Deinocerites magnus (Theobald)

Figs. 5,32-34

1901. Brachiomyia magna Theobald, 1901:344-345. TYPE: Holotype male (described as fe-

male), St. Lucia, Low, per Daniels [BM; specimen presumably lost, see Belkin, 1968:

22|\..

1909. Deinocerites troglodytus Dyar and Knab, 1909:260. TYPE: Lectotype female, Trini-

dad, W.I., June, A. Busck [USNM, 12128; designation of Stone and Knight, 1957:

197]. Synonymy with magnus by Belkin and Hogue (1959:431).

Deinocerites magnus of Belkin and Hogue (1959:431); Stone, Knight and Starcke (1959:285);

Fauran (1961:44,45); Downes (1966:1173); Fleming and Walsh (1966:425,426); Porter (1967:

Adames: Genus Deinocerites 53

39,40); Aitken, Spence, Jonkers and Downs (1969:210).

Deinocerites magna of Theobald (1905:37).

Brachiomyia magna in part of Giles (1902:474).

Deinocerites cancer of Aiken and Rowland (1906:37); Aiken (1907:76; 1909:25); Root (1922:

405); Senevet (1936: 133,134; 1938: 189); Cerqueira (1938:289-291); Pritchard and Pratt (1944:

233); Weatherbee (1944:645); Floch and Abonnenc (1947:12); van der Kuyp (1948a:748,

749; 1953:144,146); Pratt and Seabrook (1952:27); Fox (1953:179); Fox and Maldonado-

Capriles (1953:165); Maldonado-Capriles, Pippin and Kuns (1958:68).

Deinocerites cancer in part of Theobald (1903:276-281; 1905a:35-36; 1910:553); Surcouf and

Gonzalez-Rincones (1911:91); Howard, Dyar and Knab (1915:201-205); Bonne and Bonne-

Wepster (1925:173-174); Dyar (1928:262-263); Edwards (1932:222); Matheson (1944:250-

251); Lane (1953:554-556); van der Kuyp (1953b:144,146; 1954:39,46,56,57,64,65,67,71,

72,73 ,76,82,84,86,88,90,95,96,112); Carpenter and LaCasse (1955:325-327); Horsfall (1955:

599-600); Perez Vigueras (1956:476); Forattini (1958:175); Stone, Knight and Starcke (1959:

284); Porter (1967:39,40). :

Deinokerides cancer in part of Giles (1902:472-473).

Dinocerites cancer in part of Blanchard (1905:414).

Deinocerites troglodytus of Howard, Dyar and Knab (1915:206,207); Urich (1917:525-530);

Bonne-Wepster and Bonne (1923:124); Bonne and Bonne-Wepster (1925:171-173); Floch and

Abonnenc (1945:38,39,40); Stone and Knight (1957:197).

Culex (Deinocerites) troglodytus of Dyar (1918:101).

FEMALE (fig. 32). Wing 2.78 mm. Proboscis 2.04 mm. Forefemur 1.68 mm.

Abdomen about 2.55 mm. Mesonotum dark; pleural integument usually whitish,

distinctly contrasting with mesonotal integument. Head: Narrow decumbent scales

of vertex creamy; erect scales yellowish to brownish; broad decumbent scales in

lateral patch whitish. Amtenna: Torus occasionally with 1 scale; exceeding pro-

boscis from apex of flagellar segment 8. Thorax: Apn whitish (pale); ppn whitish,

with a few scales in middle; ppl whitish, with bristles and sometimes a few scales;

psp, ssp, Stp and pra usually slightly darker than rest of pleuron; paratergite light

brown; mep whitish, upper part with several bristles, microsetae absent; metameron

simple; meron, metameron and metapleuron whitish. Legs: Coxal integument whit-

ish; scaling and bristles as in cancer.

FEMALE GENITALIA (fig. 32). Sternite VIIT with strong marginal bristles con-

tinued about halfway dorsad on caudolateral border. Cercus with sternal margin

convex, tergal margin more or less concave; apex slightly upturned; the 2 special-

ized cercal bristles not inserted side by side, 1 distinctly apical and shorter; tergal

and sternal surfaces each with a moderately strong normal bristle, tergal more api-

cal. Distal part of postgenital plate (in ventral aspect) deeply notched; lateral lobe

elongate (in lateral aspect), its apical bristle markedly longer than total length of

lobe, parallel not convergent with its mate (in ventral aspect), not markedly dis-

placed from apex of lobe.

MALE (fig. 32). Wing 2.78 mm. Proboscis 2.19 mm. Forefemur 2.18 mm. Ab-

domen (not including genitalia) about 2.60 mm. Flagellar segment 13 slightly short-

er than 12. Proboscis extending to base of flagellar segment 5. Both claws of fore-

leg with a heavy almost basal tooth; anterior claw of midleg with a nearly basal

tooth.

MALE GENITALIA (fig. 33). Segment IX: Tergite lobe barely reaching base

of subapical lobe, very broad and with sinuous external and internal margins. Side-

piece: Subapical lobe with a distinct thumb; seta c spiniform, apex not markedly

attenuated. Phallosome: Dorsal parameres closely approximated in tergal aspect,

with a broad, short dorsal ‘bridge; apical spine heavy, long and only slightly curv-

54 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

ed; ventral teeth few in number, short, heavy, and arising in a single row along

the truncate lateral border. Aedeagus in tergal view slender, expanded in basal

third then narrowing apically; apex rounded.

PUPA (fig. 33). Abdomen 3.46 mm. Trumpet 0.48 mm. Paddle 0.73 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 10 reared specimens.

Cephalothorax: Integument yellowish to medium brown. Hair 1(2,2-3), 2(3,2-4),

3(2,1-3), 4(4,2-6), 5(2,2-4), 6(1), 7(3,2-5); 8(3,2-5), 9(3,2-3), 10(4-5,3-5), 1163,

3-4), 12(3,2-5). Trumpet: Short, slightly widening apically, index about 4.4-5.2.

Integument brown distad, slightly darker qn tracheoid. Tracheoid about 0.33 of

trumpet length. Abdomen: Integument medium brown. Segment I: hair 1(20,11-

30 primary branches), 2(1), 3(2), 4(3,2-4), 5(5,4-6), 6(1), 7(4,2-4), 9(1). Segment

II: hair O(1), 1(10,6-12), 2(2,2-3), 3(1), 4(5,4-6), 5(1,1-2), 6(1), 7(2,2-3), 9(1).

Segment III: hair 0(1), 1(4,2-6), 2(1), 3(3,1-3), 4(2,2-3), 5(2,1-2), 6(1), 7(4,2-5),

8(3,2-6), 9(1), 10(2,2-3), 11(1), 14(1). Segment IV: hair 0(1), 1(2,2-4), 2(1), 3

(4,3-6), 4(1), 5(1), 6(1), 7(2,2-3), 8(2,1-2), 9(1), 10(3,2-4), 11(1), 14(1). Segment

V: hair O(1), 1(2,2-3), 2(1), 3(2,2-4), 4(5,2-5), 5(1), 6(1), 7(4,3-5), 8(2,2-4), 9

(1), 10(2,1-3), 11(1), 14(1). Segment VI: hair 0(1), 1(2,2-3), 2(1), 3(2,2-3), 4(4,

2-5), 5(1), 6(1,1-2), 7(2,1-3), 8(2,1-3), 9(1), 10(2,2-3), 11(1), 14(1). Segment VII:

hair 0(1), 1(€1), 2€1), 3(2,1-2), 4(2), 5(2,1-3), 6(€1,1-2), 7(€1), 8(2,1-2), 9(3,2-4),

10(2,1-2), 11¢€1), 14(1). Segment VIII: hair 0(1), 4(2,1-2), 9(1), 14(1,1-2). Pad-

dle: Width about 0.61 of length; apex slightly produced.

FOURTH INSTAR LARVA (fig. 34). Head 1.14 mm. Siphon 1.11 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 7 reared specimens.

Hairs 2-C often closer to each other on midline than they are to 1-C as opposed

to cancer and melanophylum in which hairs 2-C are closer to 1-C than they are

to each other. Head: Integument yellowish to light brown. Mental plate distinct-

ly wider than long, triangular, at least all distal spicules sharply pointed; median

terminal spicule slightly produced or about level with others. Hair 0(1), 1(1), 2

(1), 3(sometimes developed as a minute spicule), 4(5,3-6), 5(4,3-4), 6(1), 7(8,8-

10), 8(3,3-4), 9(5,4-5), 10(3,2-3), 11(5,3-5), 12(2,2-3), 13(5,5-6), 14(1), 15(3,1-

3). Antenna: Length about 0.41 of head; shaft with several spicules on proximal

part. Hair 1(5,4-6). Thorax: Prothorax: hair 0(7,6-9), 1(1), 2(1), 3(2), 4(5,4-5),

S(1), 6(1), 7(3,2-4), 8(2,1-3), 9(1), 10(€1), 11(4,3-5), 12(1), 14(1). Mesothorax:

hair 1(1), 2(4,2-4), 3(1), 4(3,2-4), 5(1), 6(1), 7(1), 8(6,5-9), 9(7,6-9), 10(1), 11

(2,1-3), 12(1), 13(8,8-12), 14(12,10-13). Metathorax: hair 1(1,1-2), 2(4,2-4), 3(5,

5-6), 4(3,3-5), 5(1), 6(1), 7(8,8-13), 8(7,4-9), 9(8,7-15), 10(€1), 11(3,2-3), 12(2,

1-3), 13(5,4-6). Abdomen: Segment I: hair 1(1), 2(1), 3(€2,2-3), 4(8,7-12), 5(5,

5-7), 6(2,2-4), 7(2), 9(3,2-5), 10(3,2-4), 11(3,2-4), 12(2,2-3), 13(2,1-2). Segment

II: hair 0(1), 1€1), 2(1), 3(3,2-5), 4(5,3-6), 5(3,2-4), 6(2,2-3), 7(3,2-5), 8(2,2-4),

9(1), 10(1,1-2), 11(2,2-3), 12(3,2-4), 13(8,6-10). Segment III: hair 0(1), 1(2,2-3),

2(1), 3(1,1-2), 4(2,2-3), 5(2,2-3), 6(2,2-3), 7(4,3-6), 8(2,2-4), 9(1,1-2), 10(2,2-3),

11(2,2-3), 12(2,2-3), 13(1), 14(1). Segment IV: hair 0(1), 1(2,1-2), 2(1), 3(3,2-

4), 4(1,1-2), 5(3,2-4), 6(2,2-3), 7(6,3-6), 8(1,1-2), 9(1), 10(2,2-3), 11(2,1-2), 12

(2,2-3), 13(2), 14(1). Segment V: hair 0(1), 1(2,1-2), 2(1), 3(6,3-7), 4(1), 5(2,

2-3), 6(2), 7(5,5-7), 8(1,1-2), 9(1), 10(1), 11€1,1-2), 12(2), 13(2,2-3), 14(1). Seg-

ment VI: hair O(1), 1(4,2-5), 2(1), 3(1), 4(3,3-5), 5(3,2-4), 6(1,1-2), 7(3,2-4), 8

(3,2-4), 9(1), 10(€1), 11(2,2-3), 12(1), 13(12,9-13), 14(1). Segment VII: hair 0(1),

1(1), 2(1), 3(3,2-4), 4(1), 5(3,2-4), 6(8,7-11), 7(1,1-2), 8(4,4-8), 9(3,2-3), 10(1),

11(1,1-2), 12¢1), 13(2,1-2), 14(1). Segment VIII: Comb scales slender, wider api-

cally; fringe conspicuous at apex; less developed in distal half. Hair 0(1), 1(4,3-4),

Adames: Genus Deinocerites 55

2(1), 3(4,3-5), 4(1), 5(3,2-3), 14(2,1-2). Siphon: Integument light brown. Index

about 3.8-5.1. Pecten teeth 6(5-7), usually bifid, sometimes trifid, both blades slen-

der, basal shorter, rest variable in size. Hair 1(3), hair lad(2,2-3), hair lav(2,2-3);

hairs 2,6,7,8 and 9 all single. Anal Segment: Gill about 0.5 of dorsal saddle length.

Hair 1-X longer than gill. Hair 1(1,1-2), 2(8,6-9), 3(1), 4a(12,11-13), 4b(11,9-12),

4c(9,8-11), 4d(9,7-10), 4e(9,8-10), 4f(9,7-12), 4g(12,11-12).

SYSTEMATICS. Deinocerites magnus is clearly differentiated from the cancer-

melanophylum complex and is probably an earlier offshoot from the original stock

of the Cancer Group as shown by the absence of the conspicuous derived features

in the adults (increased antennal length) and the larva (reduced ventral brush) found

in cancer and melanophylum.

In spite of the extensive fragmentation of magnus into insular populations over

much of its range no significant differentiation has occurred. All the populations

I have examined conform to the diagnostic features given above and no significant

variation has been noted.

I have not seen the material reported as cancer from the state of Maranhao by

Cerqueira (1938:289-291) but judging by his figures it can be unquestionable re-

ferred to magnus.

BIONOMICS. There is no published record of a specific identification of a crab

with which magnus may be associated but its immature stages have usually been

found in large crabholes and only occasionally in smaller ones. In addition to these

normal breeding sites, magnus has been reported from a beach pool in Sint Maarten

(van der Kuyp, 1948a:748, as cancer) and has been taken recently in ground pools

(flooded crabhole areas) in Trinidad and Grenada.

As in the case of other members of the group, the reports on the blood feed-

ing habits of the females are conflicting. Theobald (1903:279, as cancer) quotes

a report from Dr. Low that the latter could not induce this species to bite on

Barbados and never saw any traces of blood in the stomachs of specimens from

St. Vincent. In Trinidad, Urich (1913:527, as troglodytus) found the species to

be a timid biter in the field and to rarely enter houses to feed. In recent years

females have been caught in Trinidad no. 10 traps baited with mice in Trinidad

(once) and baited with chicks in Antigua, Dominica, St. Lucia and Trinidad. Fe-

males were also collected biting man in the evening in St. Vincent. In Surinam,

Bonne and Bonne-Wepster (1925:175, as troglodytus) captured females indoors

and reported their bite as painful and giving ‘“‘a peculiar sensation different from

one of the ordinary house mosquitoes in Surinam”. Floch and Abonnenc (1947:

11, as cancer) report a female biting man in French Guiana.

Busck (reported in Howard, Dyar and Knab, 1915:207) observed this species

to swarm at dawn and evening and “frequently noted copulation at such times.”

This appears to be the only information available on the mating habits of magnus.

Over much of its range magnus appears to be the only normal obligate crab-

hole breeder, its various known associates being largely encountered in marginal

environments. It has been found with Aedes (O.) taeniorhynchus in Anguilla and

Antigua; with Culex (C.) habilitator in Montserrat and Nevis; with Culex (C.) in-

flictus Theobald, 1901 in Dominica and Trinidad; with species of Culex (C.) and

Culex (Mel.) in Dominica; and with a member of the Aedes (O.) scapularis com-

plex in Trinidad.

DISTRIBUTION (fig. 5). From Mona Island and Puerto Rico eastward and south-

ward throughout the Lesser Antilles, Trinidad and Tobago, the Guianas and the

state of Maranhao in Brazil. Material examined: 2138 specimens; 250 males, 1027

56 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

females, 627 larvae, 234 pupae; 151 individual rearings (97 larval, 33 pupal, 21

incomplete).

BARBADOS. Bridgetown (BAR 3,5,8) [UCLA]. Gracine Hall swamp (BAR 25) [UCLA].

Locality not specified, A. Busck [USNM]..

BRITISH VIRGIN ISLANDS. Tortola: Road Harbor, F.M. Root (LAR 5) [USNM]. Virgin

Gorda: Locality not specified, F.M. Root (LAR 8,8A) [USNM].

DOMINICA. Cabrit swamp, W.W. Wirth [USNM] ; (DOM 76,94,99,100,155,156,165) [UCLA].

Layou, W.W. Wirth; D.F. Bray [USNM]; (DOM 76) [UCLA]. Macoucheri, W.W. Wirth [USNM].

Portsmouth, D.F. Bray [USNM]; F.M. Root (LAR 19a) [USNM}.

FRENCH GUIANA. Stoupan (FG 58B) [UCLA].

GRENADA. Pt. Saline (GR 5,6,8) [UCLA].

GRENADINES. Canouan, F.M. Root (LAR 33e). Carriacou, F.M. Root (LAR 36b-2). Little

Martinique, F.M. Root (LAR 35). Little St. Vincent, F.M. Root (LAR 35b). Mustique, F.M.

Root (LAR 33d). Saline, F.M. Root (LAR 37,37A) [USNM].

GUADELOUPE. Desirade: Baie Mahault (FWI 218). Leproserie (FWI 220) [UCLA]. Island

not specified: A. Busck [USNM]. Grande Terre: Port-Louis (FWI 199,200) [UCLA] . Marie Ga-

lante: Grand Bourg (FWI 232). Grande Anse (FWI 224) [UCLA]. :

GUYANA. New Amsterdam (GUY 18) [UCLA].

LEEWARD ISLANDS. Anguilla: Little Harbour (ANG 12,13) [UCLA]. Antigua: Ballast Bay

(ANT 99,100) [UCLA]. Bethesda (ANT 20,22) [UCLA]. Bolands (ANT 4) [UCLA]. English

Harbour, J.F.G. Clark; R.H. Darsie [USNM]. Fitches Creek (ANT 87,88) [UCLA]. St. Johnston

(ANT 56) [UCLA]. Barbuda: Codrington (BAB 2,4) [UCLA]. Montserrat: Fox’s Bay (MNT

44.45 ,57,83,87,90) [UCLA]. Iles Bay Estate (MNT 16) [UCLA]. Old Road Estate (MNT 53)

[UCLA]. Nevis: Charlestown, F.M. Root (LAR 13,14a) [USNM]; between Ft. Charles and Pin-

ney’s Estate (NEV 3,32,34,49 60) [UCLA]. St. Kitts: Frigate Bay Estate (KIT 36,40,41) [UCLA].

MARTINIQUE. Locality not specified, A. Busck; F.M. Root (LAR 28A) [UCLA, USNM].

NETHERLANDS WEST INDIES. Sint Maarten: Philipsburg, E. van der Kuyp [USNM].

PUERTO RICO. Mona: W.F. Pippin [USNM]. Puerto Rico: Catano, H.D. Pratt [USNM];

T.H.G. Aitken [UCLA]. Central Aguire, G.S. Tulloch [USNM]. Dorado, G.S. Tulloch [USNM].

Ft. Buchanan, H.D. Pratt [USNM]. Isla Verde [UCLA]. Mayaguez, G.S. Tulloch [USNM]. Playa

de Humacao (PR 23-26) [UCLA]. Roosevelt Roads NS, H.C. Hurt [UCLA, USNM]. San Juan,

H.D. Pratt [USNM].

ST. LUCIA. Choc swamp (LU 48,49,51,54,153,155) [UCLA]. Cul de Sac Estate (LU 129)

[UCLA]. Marigot Harbour, R.H. Darsie [USNM]. Port Volet swamp (LU 107,117) [UCLA].

Reduit swamp (LU 76) [UCLA]. Roseau Sugar Estate (LU 24,25,27) [UCLA]. Vieux Fort

(LU 95) [UCLA].

ST. VINCENT. Kingstown, F.M. Root (LAR 29B) [USNM]. Young’s Island (VT 73) [UCLA].

TRINIDAD AND TOBAGO. Tobago: Bon Accord Estate (TOB 12,20) [UCLA]. Crown Pt.

(Brown’s), R.H. Darsie [USNM]. Trinidad: Chaguanas (TR 769) [UCLA]. Chaguaramas (U.S.

Naval Base), J. Leacock; K.L. Knight [USNM]. Guayaguayare (TR 794,800) [UCLA]. Locality

not specified, A. Busck; F. Urich [USNM]. Matelot (TR 185) [UCLA]. Monos Island (TR 592)

[UCLA]. Nariva Swamp, T.H.G. Aitken [UCLA]. Port of Spain, F.M. Root (LAR 54) [USNM].

San Souci (TR 156) [UCLA].

VIRGIN ISLANDS. St. Croix: Locality not specified, M.A. Beatty [UCLA, USNM]. St John:

Caneel Bay, T.H.G. Aitken [UCLA]. St. Thomas: Charlotte Amalie (VI 21-23,25,26,29) [UCLA].

Locality not specified, C. Cregue [USNM].

EPITEDEUS GROUP

FEMALES. Medium-sized species, wing 2.83-3.98 mm. Mesonotum dark brown,

pleuron usually tan and not strongly contrasting with mesonotal integument. An-

tenna: Flagellar segment 1 with numerous scales, about equal to combined length

of segment 2 and about one-half of 3; segment 2 with a few scales; segments

Adames: Genus Deinocerites 57

1-4 greatly elongated; segments 5-12 subequal in length but progressively slightly

shortened distad; extremely long, exceeding proboscis from at least distal third

of flagellar segment 5. Thorax: Postnotum with or without bristles on lower part.

Apn usually tan; ppn usually tan or slightly paler, scales broad on caudoventral

area; ppl tan; psp, ssp, stp and pra tan or brownish; paratergite tan or brownish;

mep usually tan and with a patch of or almost completely covered with trans-

lucent scales, Imep without bristle; meron, metameron and metapleuron usually

dark tan. Legs: Coxal integument usually tan. Forefemur completely dark, basal

two-thirds of posteroventral surface of midfemur often pale, basal half of antero-

ventral and posteroventral surfaces of hindfemur slightly paler; anteroventral mar-

gin of forefemur with a row of spiniforms or bristles, posterodorsal margin with

a row of weak bristles. Abdomen: Sternites II-VII lighter than tergites.

FEMALE GENITALIA. Sternite VIII with sclerotized distal band broad and with

several scales; caudal margin not produced into a submedian ventral angle, this area

without strongly differentiated marginal bristles. Tergite IX with 1 to several setae

on each side. Tergite X poorly differentiated dorsally, laterally with a very small

indistinct sclerotization. Cercus with a few moderately strong apical or subapical

normal bristles or with 5 or 6 apical and subapical specialized setae with twisted

apex.

MALES. Essentially similar to females. Antenna: Exceeding proboscis from api-

cal third of flagellar segment 4; flagellar segments 1-4 with scales; segments 1-6

markedly elongated but progressively shorter distad; segment 1 about equal to com-

bined length of segment 2 and one-third of 3; segments 7-13 subequal but pro-

gressively slightly shortened distad; segment 13 slightly expanded subapically. Legs:

Claws of foreleg markedly enlarged, anterior slightly larger and with a variable

submedian tooth, posterior simple or with a very small denticle; claws of midleg

not markedly enlarged and very similar in shape, anterior slightly larger and either

simple or with a small submedian tooth or denticle, posterior simple.

MALE GENITALIA. Segment IX: Tergite lobe cylindrical; angled at base; distal

part variously shaped; apex sometimes reaching base of subapical lobe. Sidepiece:

Without scales. Apicosternal lobe prominent, with a long apical seta. Phallosome:

Dorsal parameres widely separated but always with an indication of an incom-

plete dorsal bridge; articulated ventrally with a dorsal sclerotized projection from

basal third of aedeagus.

PUPAE. Cephalothorax: Hairs 2,3-C not closely approximated, 2-C as strongly

developed and as long as or longer than 3-C; hair 5-C single, moderately long, at

most equal in length to distance from its alveolus to base of trumpet and to length

of trumpet; 7-C usually single or double (1-4); hair 8-C usually triple (2-5). Meta-

notum: Hair 10-C strongly developed, always longer than 11-C, single or double.

Abdomen: Hair 1-II long, exceeding apex of tergite III, usually double or triple

(1-3), simple; 1-III-VI usually forked (except in colombianus); 5-II always mesad

and cephalad of 3-II; hair 3-V usually single or double (1-3); hair 1-VII shorter

than tergite VIII, at most reaching alveolus of hair 4-VIII. Paddle: Hair 1-P longer

than paddle.

FOURTH INSTAR LARVAE. Head: Hair 2-C short, inconspicuous, about 0.25-

0.33 of 1-C, always mesad of 1-C; hair 5-C usually double, rarely triple (1-3),

with a few very inconspicuous barbs; 6-C single and usually simple, barbs at most

few and inconspicuous, length about 1.5 of 5-C. Thorax: Prothoracic hair 9-P dou-

ble or triple. Abdomen: Hair 6-II double; dorsal sensillum of segment V mesad

of 4-V; hair 6-VI single; 1-VII very long, always longer then 3-VII and exceeding

58 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

base of siphon; 1-VIII usually with 3 or 4 branches (2-4). Siphon: Hair 1-S usu-

ally double (1 or 2). Anal Segment: Ventral brush (4-X) with 6 or 7 pairs of

hairs. Gill slightly emarginate on ventral margin; long, from 0.5-1.0 of dorsal sad-

dle length. Dorsal saddle at least moderately broad, its ventral margin sometimes

reaching alveolus of hair 1-X.

_ DISCUSSION. This group shows more unique derived characters than any other.

The adults share with the Pseudes Group the presence of mesepimeral scales but

are unique in the loss of the lower mesepimeral bristle. In the pupae, the hairs

are more strongly developed than in the other groups and abdominal hair 1-II

is rather long and double or triple instead of being multibranched as in the Span-

ius, Cancer and Pseudes Groups. The larvae are distinct from all the other groups

in a double or triple hair 9-P (always single in the others) and a double hair 5-C

(at least triple in the others).

The Epitedeus Group consists of a pair of clearly marked geminate complexes,

1 restricted to the Atlantic basin and the other to the Pacific basin. The species

within each complex show complementary allopatric distributions and are very

similar to each other but are readily distinguished by male genitalic characters

as well as those in the larvae and pupae. In both complexes the northernmost

Species appears to be the dominant one and also has retained 7 pairs of hairs in the

ventral brush of the larva: epitedeus (Atlantic), costaricensis (Pacific). The degree

of morphological differentiation and the distribution of the species suggest that

the original stock of the group first split into an Atlantic and Pacific component

and that later both of these were subdivided. 3 |

The Atlantic complex of 3 species (epitedeus, panamensis, colombianus) is char-

acterized in the adults by the presence of spiniforms instead of simple bristles

in the anteroventral margin of the forefemur. The IX tergite lobe of the male

genitalia of this complex is varied in shape but not conical and it always reaches

the base of the subapical lobe of the sidepiece. Deinocerites epitedeus occupies

the northern part of the distribution of the complex from the Gulf of Honduras

to at least as far south as Almirante, Bocas del Toro, Panama; panamensis is known

from the central part of Panama in the region of the Canal Zone; and colombi-

anus has been found in the region of San Blas, Panama, and Turbo, Colombia.

The Pacific complex consists of 2 known species only (costaricensis and cu-

riche). It is characterized in the adults by the presence of simple bristles in the

anteroventral row of the forefemur and by a short conical IX tergite lobe not

reaching the subapical lobe of the sidepiece in the male genitalia. Deinocerites

costaricensis is known to date only from Costa Rica but it is possible that it will

be found to the north as well as to the south; curiche is currently known only

from the area of the same name in northern Colombia. There are no records of

the Pacific complex from Panama but it is very likely that it is represented there

by either or both of the species and possibly a third one as well which might

occupy the intermediate area as in the case of the Atlantic complex.

All but 2 species of the complex appear to be rare. Deinocerites epitedeus is

apparently the dominant species of Deinocerites on the Atlantic coast of Central

America from the Gulf of Honduras to the province of Bocas del Toro, Panama;

and panamensis is common on the Atlantic coast of the Canal Zone. The only

reliable information on the blood feeding habits is for epitedeus in Almirante,

Panama, where it has been shown to utilize a wide range of vertebrate hosts from

amphibians to mammals. The immature stages of the group have been collected

only in large crabholes. The species of crab (Cardisoma crassum) has been iden-

Adames: Genus Deinocerites 59

tified only in the case of costaricensis.

11. Deinocerites epitedeus (Knab)

Figs. 6,35-37

1907. Dinomimetes epitedeus Knab, 1907:120-121. TYPE: Lectotype male (344a), with gen-

italia on slide (286), Port Limon, Costa Rica, F. Knab [USNM, 10291; designation of

Stone and Knight, 1957:197].

Deinocerites epitedeus of Bonne and Bonne-Wepster (1925:174); Kumm and Ruiz (1940:392,

402); Stone and Knight (1957:197); Ross (1962:192); Gorgas Memorial Institute (1970:20);

Tempelis and Galindo (1970).

Deinocerites epitedeus in part of Dyar (1925:154,156; 1928:264,537); Horsfall (1955:599); For-

attini (1958:176); Belkin and Hogue (1959:438-441); Stone, Knight and Starcke (1959:284).

Dinomimetes epitedeus of Surcouf and Gonzalez-Rincones (1911:92).

Dinomimetes epitedeus in part of Howard, Dyar and Knab (1915:197-199).

Deinocerites cancer in part of Knab (1906:95-97).

Culex (Deinocerites) epitedeus of Dyar (1918:101).

FEMALE (fig. 35). Wing 3.93 mm. Proboscis 2.75 mm. Forefemur 2.77 mm.

Abdomen about 3.52 mm. Head: Narrow decumbent scales of vertex creamy, erect

scales brownish. Antenna: Torus sometimes with a few scales; exceeding probos-

cis from middle to apex of flagellar segment 5. Thorax: Postnotum usually with

2 or more bristles near lower end; ppn with several bristles on middle and lower

areas; ppl with numerous bristles and sometimes with a few scales; mep with a

patch of translucent scales usually in upper half, umep bristles numerous, body

of sclerite with scattered microsetae; metameron sometimes with a few small bris-

tles. Legs: Anterior surface of forecoxa largely covered by bristles and scales, baso-

lateral area with some bristles; midcoxa with outer surface covered with translu-

cent scales, except for a lower posterior area, and 2 groups of bristles, those in

anterior group more numerous and scattered, those in posterior group in a row

and stronger; hindcoxa with anterolateral surface with scales, lower lateral with

bristles, upper lateroposterior with a few scales, posterior surface with numerous

bristles; forefemur with a row of spiniforms in anteroventral margin.

FEMALE GENITALIA (fig. 35). Sternite VIII sometimes with several scales.

Tergite IX with 3-6 setae on each side. Cercus in lateral aspect with sternal mar-

gin convex; tergal margin slightly sinuous at base and slightly concave distally;

apex slightly upturned, usually with 4 moderately strong apical or subapical nor-

mal bristles. Distal part of postgenital plate (in ventral aspect) with deep notch;

lateral lobe (in lateral aspect) moderately elongate, its apical bristle only slightly

convergent with its mate (in ventral aspect).

MALE (fig. 35). Wing 3.93 mm. Proboscis 3.01 mm. Forefemur 2.65 mm. Ab-

domen (not including genitalia) about 3.21 mm. Anterior claw of foreleg with

a moderately large submedian tooth; that of midleg with a small tooth.

MALE GENITALIA (fig. 36). Segment IX: Tergite lobe with distal part slen-

der, reaching base of subapical lobe but not extending beyond it; directed mesad

by distinct curvatures of inner and outer margins; body of lobe broad basally,

constricted near middle and slender in distal half. Sidepiece: Subapical lobe with

rather small thumb; seta c spiniform and attenuated apically. Phallosome: Dorsal

60 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

parameres with broad but short incomplete bridge; apical spine long and slender;

ventral teeth usually long, frequently several joined at base. Aedeagus in tergal

aspect with external margins sinuous, narrower apically after a constriction distad

of middle, apex rounded.

PUPA (fig. 36). Abdomen 4.36 mm. Trumpet 0.91 mm. Paddie 0.86 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 8 reared specimens.

Hair 1-C forked at about 0.4 from base. Hair 9-C usually forked. Cephalothorax:

Integument yellowish to medium brown. Hair 1(2), 2(2,1-2), 3(2), 4(1), 5(1), 6

(3,2-4), 7(1), 8(3), 9(2-3,2-4), 10(1), 11(2,2-3), 12(3,2-4). Trumpet: Moderately

long, width almost uniform throughout except for slight widening apically; index

about 5.3-7.6. Integument brown distad, darker on tracheoid; markedly contrast-

ing with cephalothoracic integument. Tracheoid about 0.4 of trumpet length. Ab-

domen: Integument yellowish to medium brown. Segment I: hair 1(13,9-19), 2

(1), 3(2,2-3), 4(5,3-6), 5(3,3-5), 6(1), 7(1,1-2), 9(1,1-2). Segment II: hair 0(1),

1(2,2-3), 2(2), 3(1), 4(6,4-6), 5(€1), 6(1), 7(1), 9(1). Segment III: hair 0(1), 105,

2-6), 2(1), 3(2,2-3), 4(2,2-3), 5C1), 6(1), 7(3,2-5), 8(5,3-5), 91), 10(1),. 110),

14(1). Segment IV: hair 0(1), 1(4,3-6), 2(1), 3(5,3-6), 4(2,1-2), 5(1), 6(1), 7(3,

2-4), 8(3,2-4), 9(1), 10(€1), 11(1), 14(1). Segment V: hair 0(1), 1(4,2-5), 2(1),

3(1), 4(5,3-5), 5C1), 6(1), 7(5,2-7), 8(2,1-3), 9(1), 10(1), 11€1), 14(1). Segment

VI: hair 0(1), 1(3,2-6), 2(€1), 3(1,1-2), 4(3,2-3), 5(1), 6(1), 7(1), 8(2), 9(1), 10

(1), 11€1,1-2), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(2), 4(2), 5(3,3-6), 6

(1), 7(1), 8(4,2-4), 9(5,4-6), 10(€1,1-2), 11€1,1-2), 14(1). Segment VIII: hair 0(1),

4(2,1-2), 9(1), 14(1). Paddle: Width about 0.76 of length. |

FOURTH INSTAR LARVA (fig. 37). Head 1.21 mm. Siphon 1.52 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 8 reared specimens.

Head: Integument light brown. Mental plate almost as long as wide, triangular;

marginal spicules usually sharply pointed, sometimes denticulate. Hair 0(1), 1(1),

2(1), 3(not detectable), 4(3,3-5), 5(2,2-3), 6(1), 7(5,4-11), 8(3,1-4), 9(5,4-6), 10

(3,2-3), 11(4,3-4), 12(1,1-2), 13(5,4-5), 14(1), 15(4,3-4). Antenna: Length about

0.63 of head; shaft with numerous spicules on proximal part. Hair 1(3,3-4). Tho-

rax: Prothorax: hair 0(10,8-12), 1(1), 2(1), 3(1), 4(3), 5(1), 6(1), 7(3,2-3), 81,

1-2), 9(3,2-3), 10(1), 11(4,4-10), 12(1), 14(1). Mesothorax: hair 1(1), 2(3,2-4),

3(1), 4€1), 5C1), 6(1), 7(1), 8(6,5-7), 9(7,5-8), 10(1), 11(2,2-4), 12(1), 13(11-16),

14(6,6-10). Metathorax: hair 1(1), 2(4,3-5), 3(6,4-8), 4(4,3-5), 5(1), 6(1), 7(7,6-

9), 8(10-19), 9(7,6-8), 10(1), 11(3,2-3), 12(1), 13(6,5-6). Abdomen: Segment I:

hair 1(1), 2(1), 3(€1,1-2), 4(8,8-10), 5(3,3-5), 6(2,2-3), 7(2), 9(3,2-3), 10(1), 11

(3,2-4), 12(3,2-3), 13(1). Segment II: hair 0(1), 1(1), 2(1), 3(2,1-3), 4(8,7-9), 5

(4,3-5), 6(2), 7(2,1-4), 8(4,3-4), 9(1), 10(1), 11(3,2-3), 12(2,2-5), 13(9,8-11). Seg-

ment III: hair 0(1), 1(1,1-2), 2(1), 3(1), 4(2), 5(3,2-5), 6(2), 7(6,5-9), 8(2,1-3),

9(1), 10€1), 11(2,2-3), 12(2,2-4), 13(1), 14(1). Segment IV: hair 0(1), 1(1), 2(1),

3(3,2-4), 4(2), 5(3,2-4), 6(2), 7(8,5-10), 8(2,1-2), 9(1), 10(1), 11(2), 12(3,2-3),

13(2,1-2), 14(1). Segment V: hair 0(1), 1(1), 2(1), 3(1), 4(7,6-9), 5(4,3-5), 6(2),

7(10,6-10), 8(2), 9(1), 10(1), 11(€1), 12(3,2-4), 13(2), 14(1). Segment VI: hair

O(1), 1(5,2-6), 2(1), 3(1), 4€1,1-2), 5(4,2-7), 6(1), 7(4,3-5), 8(3,2-4), 9(1), 10(1),

11(3,2-4), 12(1), 13(19-28), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(4,4-6),

4(1), 5(4,2-6), 6(12,10-15), 7(1), 8(9,8-12), 9(2,2-4), 10(1), 11(2,2-3), 12(1), 13

(2), 14(1). Segment VIII: Comb scales slender at base, fringe restricted mainly

to the apex. Hair 0(1), 1(3,2-4), 2(1), 3(4,3-5), 4(1), 5(2,2-3), 14(1). Siphon: In-

tegument light brown. Index about 4.1-5.2. Pecten teeth 5(4-6), bifid, both blades

narrow apically, longer blade wider; distal teeth slightly increasing in size. Hair

Adames: Genus Deinocerites 61

1(2,1-2), hair 1ad(4,2-6), hair lav(4,2-6); hairs 2,6,7,9 all single; 8(2,2-4). Anal

Segment: Gill subequal to dorsal saddle length. Ventral margin of dorsal saddle

reaching alveolus of hair 1-X. Hair 1-X shorter than gill. Hair 1(1,1-5), 2(6,6-8),

3(1), 4a(11,10-12), 4b(11,9-11), 4c(10,9-12), 4d(10,9-12), 4e(12,10-13), 4f(13,10-

14), 42(10,7-14).

SYSTEMATICS. This is the dominant species of the Atlantic complex of the

group and occupies the largest part of its known range from the Gulf of Hon-

duras to the province of Bocas del Toro in Panama. It is most readily different-

iated from the other species of the complex by the shape and length of the IX

tergite lobe in the male genitalia and by the presence of 7 pairs of hairs instead

of 6 in the ventral brush of the larva. The female cannot be distinguished from

those of panamensis and colombianus but the pupa is usually readily separated

from all the other species of the group by the key characters.

BIONOMICS. All the records of the immature stages of epitedeus are from large

crabholes but without any indication of the species of crab involved. The origi-

nal collection of epitedeus was made by F. Knab (1906:95, as cancer) in crab-

holes on the side of a hill near Port Limon, Costa Rica, at a considerable eleva-

tion above sea level. These crabholes contained fresh water supplied by a small

brook. Recently, epitedeus has been collected in the same general area near Port

Limon, in a cacao grove about 2 mi from the sea (CR 73), and at another local-

ity about %4-% mi from the sea (CR 76). Although no determination of the chlor-

ide content was made, it is very likely that the water in these crabholes was fresh.

Deinocerites epitedeus has been found associated with the following species:

cancer in Guatemala and Nicaragua; 1 or more species of the Culex (C.) inflictus

complex in Guatemala, Nicaragua and Costa Rica; Culex (Tin.) latisquama in Nic-

aragua; and with a species of Culex (Mel.) in Costa Rica.

Only 2 reports have appeared on the blood feeding habits of epitedeus. Kumm,

Komp and Ruiz (1946:392) caught 11 females on horse bait in Puerto Viejo, Costa

Rica. In Panama, epitedeus has been found to have a very wide range of hosts

ranging from amphibians to mammals (Gorgas Memorial Institute, 1970:20; Tem-

pelis and Galindo, 1970).

DISTRIBUTION (fig. 6). Atlantic coast from the Gulf of Honduras to Bocas

del Toro Province in Panama. Material examined: 984 specimens; 132 males, 119

females, 531 larvae, 202 pupae; 49 individual rearings (30 larval, 15 pupal, 4 in-

complete).

COSTA RICA. Limon: Cahuita (LCBA 232,234,235,238-240,243). Cieneguita, near P. Limon

[USNM]. Puerto Limon, F. Knab, type series; (K 344A) [USNM]. Puerto Viejo (181) [USNM].

Westfalia, near P. Limon (CR 73,76) [UCLA].

GUATEMALA. Jzabel: Puerto Matias (GUA 5,5A,6) [UCLA].

NICARAGUA. Zelaya: Bluefields (NI 34,51,67-70,72,73) [UCLA].

PANAMA. Bocas del Toro: Almirante (PA 271,274,1031) [UCLA].

12. Deinocerites panamensis Adames, n.sp.

Figs. 6,38-40

TYPES: Holotype male with associated larval and pupal skins (PA 598-101), Portobelo, Colon,

Panama, 9 Dec 1963, A. Quinonez [USNM]. Allotype female with associated larval and pupal

skins and genitalia slide (PA 598-104), same data as holotype [USNM]. Paratypes: 4 lpM (PA

598-102,109,110), 2 IpF (598-105,108), 4 pF (598-106,112-114), 1 IP (598-111), 1 1 (598-103),

same data as holotype [BM, UCLA, USNM].

62 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Deinocerites epitedeus of Dyar (1923:180); Arnett (1948:193).

Deinocerites epitedeus in part of Dyar (1925:154,156; 1928:264,537); Lane (1953:556,557,558,

559); Horsfall (1955:599); Forattini (1958:176); Belkin and Hogue (1959:438-441); Stone,

Knight and Starcke (1959:284).

Dinomimetes epitedeus of Busck (1908:75).

Dinomimetes epitedeus in part of Howard, Dyar and Knab (1915:197-199).

FEMALE (fig. 38). Wing 3.41 mm. Proboscis 2.50 mm. Forefemur 1.98 mm.

Abdomen about 2.9 mm. Apparently indistinguishable from epitedeus.

FEMALE GENITALIA (fig. 38). Essentially similar to epitedeus except for the

following. Tergite IX with 2-4 setae on each side. Distal part of postgenital plate

(in ventral aspect) with a deeper notch; lateral lobe basically similar and its apical

bristle parallel with or diverging from its mate.

MALE (fig. 38). Wing 3.21 mm. Proboscis 2.65 mm. Forefemur 2.60 mm. Ab-

domen (not including genitalia) about 2.5 mm. Apparently indistinguishable from

epitedeus.

MALE GENITALIA (fig. 39). Differing from epitedeus apparently only in IX

tergite lobe with distal part very long and very slender, angled laterad at base,

then ‘bent mesad to parallel sidepiece and extending beyond base of subapical lobe.

PUPA (fig. 39). Abdomen 3.58 mm. Trumpet 0.73 mm. Paddle 0.76 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 5 reared specimens.

Hair 1-C branched; 9-C usually forked. Cephalothorax: Integument yellowish. Hair

122-3), 202), 32), 40 )e SUL 1-2), (352-3), 42), 8G), 213,2-3), JOU 2y tt

(2), 12(2). Trumpet: As in epitedeus; index about 6.2-8.1, tracheoid about 0.50

of trumpet length. Abdomen: Integument yellowish. Segment I: hair 1(13,13-18),

2(1,1-2), 3(2,2-3), 4(4,3-5), 5(5,3-5), 6(1), 7(2), 9(1,1-2). Segment II: hair 0(1),

1(3,1-3), 2(2), 3(1), 4(6,3-7), 5(1), 6(1), 701), 9(1). Segment HI: hair 0(1), 165,

3-6), 201), 302), 4021-3), 301), OCL),. 43,24), 866,4-6), 261), 1OC1), LIC), oat

(1). Segment IV: hair 0(1), 1(5,4-5), 2(1), 3(4,3-5), 4(€1,1-2), 5(1), 6(1), 7(2,2-

3), 8(2,2-3), 9(1), 10(1), 11(€1), 14(1). Segment V: hair 0(1), 1(5,4-5), 2(1), 3

(2,1-3),, 405,46), 501), 60), 1(5,4-0), 802,2-3), 9(1), POC); 11(1),.-14C). bee

ment VI: hair 0(1), 1(4,3-4), 2(1), 3(2,1-2), 4(2,2-4), 5(1), 6(1), 7(1), 8(3,2-3),

9(1), 10(1,1-2), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(2,2-3), 4(2,1-2), 5

(2,2-3), 6(1,1-2), 7(1), 8(3,2-4), 9(5,4-6), 10(2,1-2), 11(2,1-3), 14(1). Segment

VIII: hair O(1), 4(2), 9(1), 14(1). Paddle: Width about 0.86 of length.

FOURTH INSTAR LARVA (fig. 40). Head 1.18 mm. Siphon 1.52 mm. All

measurements from skins. Diagnostic characters as in the key; general chaetotaxy

based on 5 reared specimens. Head: Integument yellowish to light brown. Mental

plate similar to that of epitedeus. Hair O(1), 1(1), 2(1), 3(not detectable), 4(4,

3-5), 5(2,1-2), 6(1), 7(5,4-7), 8(4,4-6), 9(6,3-6), 10(3,1-3), 11(4,4-5), 12(2), 13

(6,5-7), 14(1), 15(2,2-3). Antenna: Length about 0.50 of head; shaft with num-

erous spicules on proximal part. Thorax: Prothorax: hair 0(12,10-12), 1(1), 2(1),

3(1), 4(4,4-5), 5(1), 6(1), 7(3,2-4), 8(2), 9(2,1-2), 10(1,1-2), 11(4,3-5), 121), 14

(1). Mesothorax: hair 1(1), 2(3,2-4), 3(1), 4(2,1-2), 5(1), 6(1), 7(1), 8(7,6-8), 9

(9,7-9), 10(1), 11(3,2-3), 12¢€1), 13(2), 14(?). Metathorax: hair 1(1), 2(4), 3(6,

6-8), 4(4,3-5), 5(1), 6(1), 7(8,8-11), 8(?), 9(10,8-10), 10(1), 11(2,2-3), 12(1), 13-—

(6,5-6). Abdomen: Segment I: hair 1(1), 2(1), 3(2,2-3), 4(9,8-11), 5(4,3-6), 6(3,

2-3), 7(2,1-2), 9(3,2-4), 10(2,1-2), 11(2,2-3), 12(4,2-4), 13(1,1-2). Segment II:

hair O(1), 1(€1), 2(1), 3(2,2-3), 4(9,8-10), 5(4,3-5), 6(2,2-3), 7(4,3-4), 8(4,3-4), 9

(1), 10€1), 11(3,2-3), 13(3,2-4), 13(8-11). Segment III: hair 0(1), 1(2,2-3), 2(1),

M1), 42,2-3), 32-3), OL), 18-7), 82), FU4), POC). 1102), F2(3,2-3), 13C1),

Adames: Genus Deinocerites 63

14(1). Segment IV: hair 0(1), 1(€1,1-2), 2(1), 3(3,3-5), 4(2), 5(3,2-4), 6(2), 7(8-

9), 8(2,1-2), 9(1), 10€1), 11€2,2-3), 12(3,2-3), 13(2,1-2), 14(1). Segment V: hair

OG1),. 1C1, 1-2), 204), SOL), Bey. Set 3-4), Ol 2), 1095-11), BZ), OU), TOC),

11(2,2-3), 12(1,1-2), 13(2,1-3), 14(1). Segment VI: hair 0(1), 1(4,4-5), 2(1), 3

CP); 4(3,2-3), 5(4), 6(1), 7(4,3-6), 8(3,3-4), 9(1), 10(1), 11(3,2-3), 12(1), 13(2),

14(1). Segment VII: hair 0(1), 1€1), 2(1), 3(4,4-5), 4(1), 5(3,3-4), 6(12,11-13),

7(1), 8(8,5-9), 9(2,2-3), 10(1), 11(2), 121), 13(2,1-2), 14(1). Segment VIIT: Comb

scales similar to those of epitedeus. Hair 0(1), 1(3,3-4), 2(1), 3(4,4-5), 4(1), 5(4,

2-4), 14(1). Siphon: Integument light brown. Index about 4.8-6.0. Pecten teeth

5(4-6), shape and size similar to those of epitedeus. Hair 1(2), hair lad(4,1-4),

hair lav(4,1-5); hairs 2,6,7,9 all single; 8(2,1-2). Anal Segment: Gill length prob-

ably about the same as in epitedeus. Ventral margin of dorsal saddle reaching al-

veolus of hair 1-X. Hair 1-X shorter than gill. Hair 1(1,1-2), 2(7,6-9), 3(1), 4a

(14,11-15), 4b(12,11-16), 4c(14,10-15), 4d(13,11-16), 4e(14,12-17), 4£(14,10-15).

SYSTEMATICS. Deinocerites panamensis occupies a restricted central part of

the range of the Atlantic complex of the group in Central Panama and Canal Zone

where it seems to be as common as epitedeus is to the north. It is readily differ-

entiated from the other members of the complex by the shape and length of IX

tergite lobe of the male genitalia. Its larva has only 6 pairs of hairs in the ventral

brush as does colombianus as well-as curiche but it can be distinguished from

both by the key and diagnostic characters given above. The female is indisting-

uishable from the other 2 members of the Atlantic complex (epitedeus and co-

lombianus) but the pupa is usually readily separated by the key characters from

all the species of the group.

At the present time, there are wide gaps between the known distributions of

epitedeus, panamensis and colombianus on the inaccessible Atlantic coast of west-

ern and eastern Panama respectively and it is not known if these species actually

come in contact.

BIONOMICS. This species has been collected only in large crabholes. There is

no definite information about the blood feeding habits of panamensis, the only

reference to them being that of Busck (1908:75, as epitedeus from Colon, Pan-

ama) who stated, “Though possessing a well-developed proboscis, this species prob-

ably does not bite man; but this should be definitely ascertained by observations.”

Deinocerites panamensis has been found associated with the following species:

melanophylum, species of the Culex (C.) inflictus complex, Culex (Tin.) latisqua-

ma and Anopheles (A.) eiseni.

_ DISTRIBUTION (fig. 6). Atlantic coast of central Panama and Canal Zone. Ma-

terial examined: 195 specimens; 51 males, 96 females, 23 larvae, 25 pupae; 23

individual rearings (20 larval, 2 pupal, 1 incomplete).

PANAMA AND CANAL ZONE. Canal Zone: Ancon (?), J. Zetek [USNM]. Arenal River,

C.S. Ludlow [USNM]. Caldera, A-H. Jennings (160) [USNM]. Cativa, J.B. Shropshire [UCLA].

Ft. Davis, D. Baker [USNM]. Ft. Gulick, S.J. Carpenter [UCLA]. Ft. Lorenzo, J.B. Shropshire

[USNM]. Ft. Randolph, H.G. Dyar and R.C. Shannon [USNM]. Ft. Sherman, S.J. Carpenter

[UCLA]; W.H.W. Komp (KO 32-22); J. Zetek; L.H. Dunn [USNM]. France Field, J. Zetek

[USNM]. Locality not specified, A-H. Jennings (129,150,158,177,180) [USNM]. Majagual, J.

B. Shropshire [UCLA, USNM]. Colon: Colon, A. Busck (213) [USNM]. El Corredor (PA 1013,

1016) [UCLA]. Lagarto [UCLA, USNM]. Maria Chiquita (PA 1006,1008) [UCLA]. Portobelo

(PA 585,598) [UCLA]. Salud (PA 573) [UCLA].

64 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

13. Deinocerites colombianus Adames, n.sp.

Figs. 6,41-43

TYPES: Holotype male with associated larval and pupal skins and genitalia slide (COA 40-27),

La Punta, Turbo, Antioquia, Colombia, 30 Aug 1967, A.J. Adames and A. Quinonez [USNM].

Allotype female with associated larval and pupal skins. and genitalia slide (COA 40-22), same

data as holotype [USNM]. Paratypes: 2 pM (COA 39-10,13), 1 IpF (39-11), 2 IpF (40-20,28),

1 F (38), 2 L (35), 8 L (39), 1 L (40), same data as holotype [BM, UCLA, USNM].

FEMALE (fig. 41). Wing 3.67 mm. Proboscis 2.09 mm. Forefemur 2.14 mm.

Abdomen about 3.41 mm. Essentially similar to epitedeus from which it is ap-

parently indistinguishable.

FEMALE GENITALIA (fig. 41). Sternite VIII sometimes with a few scales. Ter-

gite IX with 4-6 setae on each side. Cercus essentially similar to epitedeus. Distal

part of postgenital plate basically similar to epitedeus except for slight differences

in contour of lateral lobe (in ventral aspect), apical bristle parallel with its mate.

MALE (fig. 41). Wing 3.13 mm. Proboscis 2.70 mm. Forefemur 2.50 mm. Ab-

domen (not including genitalia) about 2.24 mm. Anterior claw of foreleg as in

epitedeus, that of midleg simple.

MALE GENITALIA (fig. 42). Differing from epitedeus apparently only in IX

tergite lobe with distal part broad and directed laterad; body of lobe broad in

basal two-thirds, slightly attenuated in distal third; reaching base of subapical lobe

but not extending beyond it.

PUPA (fig. 42). Abdomen 4.05 mm. Trumpet 0.71 mm. Paddle 0.76 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 5 reared specimens.

Hairs 1-C and 9-C branched. Cephalothorax: Integument yellowish to medium

brown. Hair 1(2), 2(2), 3(2), 4(1), 5(1,1-2), 6(2,1-3), 7(2,1-4), 8(3), 9(3,2-3), 10

(2,1-2), 11(2,2-3), 12(2). Trumpet: As in epitedeus; index about 5.2-6.1. Trach-

eoid about 0.45 of trumpet length. Abdomen: Integument yellowish to medium

brown. Segment I: hair 1(16,14-18), 2(1,1-2), 3(2), 4(4,3-5), 5(3,3-4), 6(1,1-2),

7(2), 9(1). Segment II: hair 0(1), ‘1(2,2-3), 2(2), 3(1), 4(4,2-4), 5(1), 6(1,1-2),

7(1), 9(1). Segment III: hair 0(1), 1(3,3-4), 2(1), 3(2), 4€1,1-2), 51), 6(1), 7

(3,2-4), 8(4,4-6), 9(1), 10(1), 11¢€1), 14(€1). Segment IV: hair 0(1), 1(4,3-4), 2(1),

3(4,3-6), 4(1), 5(1), 6(1), 7(2,1-2), 8(2,2-3), 9(1), 10(1), 11¢1), 14(1). Segment

V: hair 0(1), 1(3,2-4), 2(1), 3(1,1-2), 4(5,3-6), 5(1), 6(1), 7(4,2-5), 8(2,2-3), 9

(1), 10(€1), 11(1), 14(1). Segment VI: hair 0(1), 1(2,2-4), 2(1), 3(€1,1-2), 4(2,2-

3), 5(1), 6(1), 7(1,1-2), 8(2,1-2), 9(€1), 10(€1,1-2), 11(1), 14(1). Segment VII: hair

OO), 11), 201), 30,1-2),, 4(2, 1-2), 32,23), 6U1), 7CD, 8(2,2-3), 96,5-7), 102,

1-2), 11(€1,1-2), 14(1). Segment VIII: hair 0(1), 4(2), 9(1), 14(1). Paddle: Width

about 0.8 of length.

FOURTH INSTAR LARVA (fig. 43). Head 1.27 mm. Siphon 1.52 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 6 reared specimens.

Head: Integument light brown. Mental plate similar to that of epitedeus except

for the subapical spicules which are usually denticulate. Hair 0(1), 1(1), 2(1), 3

(not detectable), 4(4,3-5), 5(2,2-3), 6(1), 7(7,5-8), 8(4,4-5), 9(5,4-6), 10(3,2-4),

11(5,4-6), 12(2,1-3), 13(7,5-8), 14(1), 15(3,2-4). Antenna: Length about 0.45 of

head; shaft with numerous spicules on proximal part. Hair 1(4,3-6). Thorax: Pro-

thorax: hair 0(12,9-12), 3(1), 201), 3(1,1-2),) 4(4,3-5), 5C11-2),. 61), 7(4,3-4), 3

(2,2-3), 9(2,2-3), 10(1), 11(3,3-4), 12(1), 14(1). Mesothorax: hair 1(1), 2(2,2-3),

3(1), 4(2), 5(1), 6(1), 7(€1), 8(8), 9(7-9), 10(€1), 11(2,2-3), 12(1), 13@2), 14(8-

Adames: Genus Deinocerites 65

12). Metathorax: hair 1(1), 2(2,2-3), 3(5,4-7), 4(4,3-4), 5(1), 6(1), 7(9,8-9), 8(%),

9(9,8-9), 10(1), 11(4), 12(1), 13(5,4-6). Abdomen: Segment I: hair 1(1), 2(1),

3(3,2-3), 4(9,8-9), 5(3,3-4), 6(3,2-3), 7(2), 9(2,1-3), 10(1), 11(2), 12(3,3-4), 13

(1). Segment II: hair 0(1), 1(1), 2(1), 3(2), 4(7,7-12), 5(4,3-5), 6(2,2-3), 7(2,2-

4), 8(4,2-4), 9(1), 10(€1), 11(2,2-3), 12(2), 13(?). Segment III: hair 0(1), 1(1,1-

2); 244), 301), M223) 562:2:3)))-602-3) 5-719 5-9) BU 2) Ob) 10G1 Ind), 41

(2), 12(2), 13(2), 14(1). Segment IV: hair 0(1), 1(2), 2(1), 3(2,2-3), 4(2,2-3),

5(3,2-3), 6(2), 7(8-10), 8(1), 9(1), 10€1), 11(€2,1-2), 12(€1), 13(4,3-5), 14(1). Seg-

ment V: hair O(1), 2(2), 3(1), 4(7,6-9), 5(3,3-4), 6(2), 7(10,7-10), 8(1), 9(1),

10(1), 11(2,1-2), 12(2,2-3), 13(4,4-5), 14(1). Segment VI: hair 0(1), 1(5,3-5), 2

1)s BUA) (3,24); 505,325). GED): TIS 2), 8253), 2C), FOG) (2 Ae

13(?), 14(1). Segment VII: hair 0(1), 1(2), 3(4,3-4), 4€1), 5(3-4), 6(13,12-14),

7(1), 8(7-9), 9(3), 10(1), 11(2,2-3), 12(1), 13(2,2-3), 14(1). Segment VIIT: Comb

scales similar to those of epitedeus. Hair O(1), 1(4,3-4), 2(1), 3(4,4-5), 4(1), 5

(5,5-6), 14(1). Siphon: Integument light brown. Index about 5.1-5.6. Pecten teeth

3(2-5); shape and size as in epitedeus. Hair 1(2), hair lad(4,3-4), hair lav(4,3-4);

hairs 2,6,7,9 all single; 8(2,1-3). Anal Segment: Gill about 0.53 of dorsal saddle

length. Ventral margin of dorsal saddle reaching alveolus of hair 1-X. Hair 1-X

longer than gill. Hair 1(1), 2(8,7-12), 3(1), 4a(13,12-17), 4b(12,11-14), 4c(12,10-

13), 4d(12,10-14), 4e(13,12-14), 4f(9,9-10).

SYSTEMATICS. Deinocerites colombianus is the most clearly marked of the 3

members of the Atlantic complex of the group. Its female however cannot be

differentiated from the other 2 species. In addition to the diagnostic features in

the keys, the larva of colombianus is distinguished from epitedeus by the reduc-

tion of the ventral brush to 6 pairs of hairs (from 7 pairs in the latter). This

species occupies the southern part of the range of the complex from the San Blas

area of Panama into the Turbo area of Colombia.

BIONOMICS. The immature stages of colombianus have been collected in large

crabholes in association with melanophylum and Culex (Tin.) latisquama. Nothing

is known of the blood feeding habits of this species.

DISTRIBUTION (fig. 6). Atlantic coast of eastern Panama and western Colombia.

Material examined: 99 specimens; 47 males, 23 females, 20 larvae, 9 pupae; 9 lar-

val individual rearings.

COLOMBIA. Antioquia: Atrato delta (COA 31) [UCLA]. Turbo (COA 35,39,40), type ser-

ies [BM, UCLA, USNM].

PANAMA. San Blas: Cuadi River, B.F. Eldridge [UCLA, USNM].

14. Deinocerites costaricensis Adames & Hogue

Figs. 6,44-46

1970. Deinocerites costaricensis Adames and Hogue, 1970:12. TYPE: Holotype female (CR

28-213) with associated larval and pupal skins, 1 km north Boca del Rio Barranca,

Hacienda Bonilla, Puntarenas Province, Costa Rica, 17 Nov 1962, C.L. Hogue and W.A.

Powder [USNM].

Deinocerites species A of Belkin and Hogue (1959:438); Ross (1962:192); Hogue and Wirth

(1968:6). |

FEMALE (fig. 44). Wing 3.98 mm. Proboscis 2.37 mm. Forefemur 2.24 mm.

66 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Abdomen about 3.77 mm. Head: Narrow decumbent scales creamy; erect scales

usually yellowish. Antenna: Torus sometimes with a few scales; exceeding pro-

boscis at least from middle of flagellar segment 5. Thorax: Postnotum without

hairs. Middle and lower parts of ppn with numerous bristles; ppl bristles very nu-

merous, scales few; mep with translucent scales usually covering most of the scler-

ite, body of sclerite also with very numerous microsetae, umep bristles numerous;

metameron with several small hairs. Legs: Scaling similar to epitedeus except for

more numerous scales in lateroposterior surface of hindcoxa; forefemur with an-

teroventral margin with a row of bristles.

FEMALE GENITALIA (fig. 44). Sternite VIII usually without scales, at most

1 present. Tergite IX with 1-3 setae on each side. Cercus (in lateral aspect) with

dorsal margin of sclerotized part distinctly longer than ventral, its basal width less

than 0.5 of length of sclerotized ventral margin; ventral margin convex, dorsal

margin more or less straight but slightly directed ventrad distally; apex directed

caudad, with 5 or 6 apical or subapical specialized setae with twisted apex. Dis-

tal part of postgenital plate (in ventral aspect) with deep round notch, apicolateral

margins more or less conical; lateral lobe (in lateral apsect) elongate, its apical

bristle parallel with its mate (in ventral aspect).

MALE (fig. 44). Wing 3.13 mm. Proboscis 2.71 mm. Forefemur 2.37 mm. Ab-

domen (not including genitalia) about 2.79 mm. Anterior claw of foreleg with a

very long slender tooth, posterior claw with a very small denticle; anterior claw

of midleg with a small denticle.

MALE GENITALIA (fig. 45). Segment IX: Tergite lobe broad at base and cone

shaped distally; apical part not strongly attenuated and not reaching base of sub-

apical lobe. Sidepiece: Subapical lobe with rather small thumb; seta c spiniform,

with an apical attenuation. Phallosome: Dorsal parameres widely separated but with

a slight indication of an incomplete dorsal bridge; apical spine long and slender;

ventral teeth about 18, short to moderately long, slender, arising from an evenly

convex ventrolateral border. Aedeagus (in tergal aspect) more or less uniform in

width but with a premedian constriction and an apical expansion.

PUPA (fig. 45). Abdomen 4.07 mm. Trumpet 0.76 mm. Paddle 0.48 mm. Gen-

eral chaetotaxy based on 10 reared specimens. Except for diagnostic characters

in the key apparently indistinguishable from curiche. Hair 1-C forked, main stem

about 0.4 of total length, 9-C usually slightly forked. Cephalothorax: Integument

yellowish. Hair 1(2,1-2), 2(2), 3(2,1-2), 4(1), 5(1,1-2), 6(2,2-5), 7(1), 8(3,3-4),

9(2,1-4), 10(1), 11(€2,1-3), 12(2,1-3). Trumpet: As in epitedeus; index about 5.3-

7.6. Tracheoid about 0.5 of trumpet length. Abdomen: Integument yellowish to

medium brown. Segment I: hair 1(15,7-20), 2(1,1-2), 3(2), 4(3,2-4), 5(5,4-6), 6

(1), 7(2,1-3), 9(€1), 10Coccasionally present, 1), 11(occasionally present, 2,1-3).

Segment II: hair 0(1), 1(2,2-3), 2(2), 3(1), 4(4,4-6), 5(1), 6(1), 7(1), 9(1). Seg-

ment III: hair 0(1), 1(5,4-7), 2(1), 3(2,2-4), 4(2,1-3), 5(1), 6(1), 7(3,2-5), 8(5,

4-6), 9(1), 10(1), 11(€1,1-3), 14(1). Segment IV: hair 0(1), 1(5,3-6), 2(1), 3(3,

2-6), 4(2,1-2), 5(1), 6(1), 7(3,2-4), 8(2,2-3), 9(1), 10(1), 11(1,1-2), 14(1). Seg-

ment V: hair O(1), 1(4,2-5), 2(1), 3(1), 4(4,2-6), 5(1), 6(1), 7(4,2-5), 8(3,1-3),

9(1), 10€1), 11€1,1-2), 14(1). Segment VI: hair 0(1), 1(4,2-5), 2(1), 3(1), 4(2,

1-2), 5(1), 6(1), 7(1), 8(2,1-4), 9(1), 10(1), 11(2,1-2), 14(1). Segment VII: hair

O(1), 11), 2(1), 3(2,1-2), 4(1,1-2), 5(3,1-5), 6(1), 7(1), 8(2,2-3), 9(4,3-6), 10(1),

11(2,1-2), 14(1). Segment VIII: hair 0(1), 4(2,1-2), 9(1), 14(1). Paddle: Width

about 0.8 of length. 3

FOURTH INSTAR LARVA (fig. 46). Head 1.22 mm. Siphon 1.43 mm. Diag-

Adames: Genus Deinocerites | 67

nostic characters as in the key; general chaetotaxy based on 9 reared specimens.

Head: integument yellowish to light brown. Mental plate almost as wide as long,

triangular, with apical and subapical spicules denticulate, lateral ones usually sharp-

ly pointed, sometimes denticulate. Hair 0(1), 1(1), 2(1), 3(not detectable), 4(3,

2-5), 5(2), 6(1), 7(6,4-8), 8(5,3-6), 9(5,3-5), 10(3,2-4), 11(3,2-4), 12(2,1-2), 13

(6,5-8), 14(1), 15(4,3-5). Antenna: Length about 0.33 of head; shaft with num-

erous minute spicules on proximal part. Hair 1(4,3-5). Thorax: Prothorax: hair

0(9,9-13), 1(1), 2(1), 3(1), 4(3,2-4, 5(1), 6(1), 7(3,2-4), 8(1,1-2), 9(3,2-3), 10

(1), 11(4,4-6), 12(1), 14(1). Mesothorax: hair 1(1), 2(3,2-5), 3(1), 4(1,1-2), 5

(1), 6(1), 7(1), 8(8,6-8), 9(8,7-10), 10(1), 11(3,2-4), 12(1), 13(16,12-19), 14(?).

Metathorax: hair 1(1), 2(3,2-4), 3(5,4-7), 4(4,3-6), 5(2,1-2), 6(1), 7(9,7-9), 8(12,

12-24), 9(10,8-10), 10(1), 11(3,3-5), 12(1), 13(6,5-7). Abdomen: Segment I: hair

1(1), 2(€1), 3(2,1-2), 4(9,7-12), 5(5,4-6), 6(2,2-3), 7(2,1-2), 9(2,2-3), 10(2,1-2),

11(2,1-4), 12(2), 13(1). Segment II: hair 0(1), 1(1), 2(1), 3(2), 4(9,5-10), 5(5,

4-5), 6(2,2-3), 7(5,4-6), 8(4,2-5), 9(1), 10(1), 11(3,2-4), 12(2,2-3), 13(14,12-16).

Segment III: hair 0(1), 1(1,1-2), 2(1), 3(€1), 4(2), 5(3,2-4), 6(2), 7(7,5-12), 8(,

(2,1-2), 2(1,1-2), 3(2,1-3), 4(2), 5(4,2-5), 6(2), 7(10,7-10), 8(2,1-2), 9(1), 10(1),

11(2,2-5), 12(2,1-3), 13(1,1-2), 14(1). Segment V: hair 0(1), 1(1), 2¢1), 3(1), 4

(6,5-8), 5(5,2-5), 6(2), 7(6,5-9), 8(2,2-3), 9(1), 10(1), 11(3,2-3), 12(1), 13@,]1-

2), 14(1). Segment VI: hair 0(1), 1(5,3-7), 2(1), 3(1), 4(2,1-2), 5(5,3-5), 6(1),

7(4,2-5), 8(3,2-4), 9(1), 10€1), 11(3,3-4), 12(1), 13(16-25), 14(1). Segment VII:

hair O(1), 1(1), 2(1), 3(5,4-7), 4(€1), 5(2,2-4), 6(11,9-13), 7(1), 8(7,7-11), 9(3,2-

5), 10(1), 11(€2,1-4), 12(1), 13€1,1-2), 14(1). Segment VIIT: Comb scales slender

at base, apex fringed, slightly fringed on lateroapical margin. Hair O0(1), 1(3,3-4),

2(1), 3(4,4-5), 4(1), 5(2), 14(1). Siphon: Integument light brown. Index about

3.8-5.6. Pecten teeth 6(4-6), similar to those of epitedeus. Hair 1(2,1-2), hair lad

(3,2-5), hair lav(3,3-4); hairs 2,6,7 and 9 all single; 8(2,1-2). Anal Segment: Gill

subequal to dorsal saddle length. Ventral margin of dorsal saddle not reaching al-

veolus of hair 1-X. Hair 1-X shorter than gill. Hair 1(2,1-3), 2(9,6-10), 3(1), 4a

(12,12-15), 46b(13,11-14), 4c(12,9-15), 4d(13,12-15), 4e(13,12-15), 4f(14,12-16),

49(12,11-14).

SYSTEMATICS. Deinocerites costaricensis is the better known of the 2 species

comprising the Pacific complex of the group and occupies the northern part of

its range in Costa Rica. It is possible that it occurs also in northern Panama. The

females of the 2 species are usually readily differentiated by the key characters.

Of particular interest are the specialized setae of the cercus which are better de-

veloped in the female of costaricensis. These specialized setae, I believe, have aris-

en independently and are not indicative of any relationship with the Cancer Group

which has a single pair of spatulate cercal setae. Therefore, I reject the suggestion

of Belkin and Hogue (1959:438) that costaricensis (as species A) is a species of

hybrid origin between the Cancer Group and the mcdonaldi stock.

The 2 species of the complex cannot be differentiated on external features of

the adults but are usually readily separated in the other stages and the male gen-

italia by the key characters. The larva of costaricensis retains the primitive con-

dition of 7 pairs of hairs in the ventral brush while that of curiche has the brush

reduced to 6 pairs of hairs.

At present there is a wide gap between the distribution of costaricensis and

that of curiche from the Osa Peninsula in Costa Rica to northern Colombia. It

is probable that 1 or more species of the complex will be found in this area, 1

68 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

or both of the known species and possibly a third one which would occupy the

central area of the distribution of the Pacific complex as in the case of the At-

lantic complex of the group.

BIONOMICS. All records of costaricensis are from the burrows of the Mouth-

less Crab, Cardisoma crassum, where it is usually associated with the more com-

mon pseudes. It has also been found associated with a species of the Culex (C._)

inflictus complex and a species of Culex (Mel.). There is also a record of a single

larva from a mangrove treehole (CR 130) in association with Culex (Anoed.) con-

servator, Haemagogus chalcospilans, Orthopodomyia fascipes and Corethrella (C.)

SD.

DISTRIBUTION (fig. 6). Pacific coast of Costa Rica. Material examined: 689

specimens; 31 males, 41 females, 528 larvae, 89 pupae; 38 individual rearings (23

larval, 5 pupal, 10 incomplete).

COSTA RICA. Puntarenas: Boca del Rio Barranca (LCBA 111,155,156,158,159,174) [UCLA].

Boca del Rio Baru (CR 34,43) [UCLA]. Hacienda Bonilla (CR 28,106,107) [BM, LACM, UCLA, —

USNM]. Rincon, Peninsula de Osa (CR 122-124,130,135,136,142) [UCLA]. Tarcoles (835)

[USNM].

15. Deinocerites curiche Adames, n.sp.

Figs. 6,47-49

TYPES: Holotype male with associated larval and pupal skins (COA 50-14), El Naranjo, Cu-

tiche, El] Choco, Colombia, 31 Aug 1967, A.J. Adames and A. Quinonez [USNM]. Allotype

female with associated larval and pupal skins (COA 50-11), same data as holotype [USNM].

Paratypes: 2 \pM (COA 50-12,19), 1 pF (50-100), 2 IP (50-10,18), 9 L (50), same data as

holotype [BM, UCLA, USNM].

FEMALE (fig. 47). Wing 2.83 mm. Proboscis 2.75 mm. Forefemur 2.39 mm.

Abdomen about 2.5 mm. Apparently indistinguishable from costaricensis.

FEMALE GENITALIA (fig. 47). Sternite VITIT sometimes with several scales.

Tergite [IX with 1-3 setae on each side. Cercus (in lateral aspect) with the dorsal

and ventral margins of sclerotized part subequal; its basal width over 0.6 of length

of sclerotized ventral margin; ventral and dorsal margins convex; apex directed

caudad; 5 or 6 weak to moderately strong apical or subapical bristles, with or

without twisted apex. Distal part of postgenital plate basically as in costaricensis

except for a shallower notch (in ventral aspect).

MALE (fig. 47). Wing 3.16 mm. Proboscis 2.42 mm. Forefemur 2.24 mm. Ab-

domen (not including genitalia) about 2.50 mm. Apparently indistinguishable from

costaricensis except for posterior claw of foreleg which appears to be single.

MALE GENITALIA (fig. 48). Segment IX: Tergite lobe similar to that of co-

staricensis except for a more sinuous inner margin and blunter apex. Sidepiece:

As in costaricensis. Phallosome: Dorsal paramere with about 25 ventral teeth, us-

ually long, moderately heavy and arising from a distinct humplike expansion of

ventrolateral border. Aedeagus similar to that of costaricensis.

PUPA (fig. 48). Abdomen 4.61 mm. Trumpet 0.84 mm. Paddle 0.78 mm. Gen-

eral chaetotaxy based on 5 reared specimens. Except for diagnostic key characters

indistinguishable from costaricensis. Hair 1-C usually forked, main stem about 0.4

of total length, 9-C sometimes slightly forked. Cephalothorax: Integument med-

ium brown. Hair 1(2), 2(2), 3(2), 4(1), 5(1), 6(2,2-4), 7(2,1-2), 8(3), 9(3,1-3),

Adames: Genus Deinocerites 69

10(1), 11(2), 12(1). Trumpet: As in epitedeus; index about 5.4-7.1. Tracheoid

about 0.3-0.4 of trumpet length. Abdomen: Integument yellowish to medium brown.

Segment I: hair 1(13,10-15), 2(1,1-2), 3(2), 4(3,1-4), 5(2,2-3), 6(1), 7(2,1-2), 9

(1), 10(occasionally present, 1), 11(occasionally present, 1). Segment II: hair 0

(1), 1(2,1-2), 2(2,1-2), 3(1), 4(3,1-4), 5(1), 6(1,1-2), 7(1), 9(1). Segment III: hair

O(1), 1(4,4-6), 2(1,1-2), 3(2,1-3), 4(2), 5(1), 6(1), 7(2,2-4), 8(5,4-7), 9(1), 10(1),

11(1), 14(1). Segment IV: hair O(1), 1(5,5-6), 2(1), 3(3,1-4), 4(1), 5(1), 6(1),

7(3,2-3), 8(3,1-3), 9(1), 10(€1), 11€1), 14€1). Segment V: hair 0(1), 1(6,4-8), 2

(2)53( 1,142), 406,2-6); S(T), 601), 733-7), 803,24), 901), 100), TAC 4).

Segment VI: hair 0(1), 1(5,4-7), 2(1), 3(1), 4(2), 5(€1), 6(1), 7(1,1-2), 8(2,2-3),

9(1), 10(1), 11(2), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(1,1-2), 4(2,1-2),

5(3,2-5), 6(1), 7(1), 8(2,1-2), 9(4,4-5), 10(2), 11(2), 14(1). Segment VIII: hair

O(1), 4(2,1-2), 9(1), 14(1). Paddle: Width about 0.8 of length.

FOURTH INSTAR LARVA (fig. 49). Head 1.27 mm. Siphon 1.31 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 5 reared specimens.

Head: integument yellowish to light brown. Mental plate similar to that of co-

staricensis, except for lateral spicules which are usually all denticulate. Hair 0(1),

1(1), 2(1), 3(not detectable), 4(3,2-5), 5(2), 6(1), 7(6,5-7), 8(4,4-6), 9(5,4-7), 10

(3,1-3), 11(3,2-5), 12(1), 13(6,5-7), 14(1), 15(3,3-4). Antenna: Length about 0.44

of head; shaft with numerous spicules on proximal part. Hair 1(4). Thorax: Pro-

thorax: hair 0(8,8-10), 1(1), 2(1), 3€1), 4(3,2-4), 5(1), 6(1), 7(3,2-3), 8(1), 9(2),

10(1), 11(3), 12(1), 14(1,1-2). Mesothorax: hair 1(1), 2(3,2-4), 3(1), 4(1,1-2),

S(1), 6(1,1-2), 7(€1), 8(8-9), 9(6-9), 10(1), 11(3), 12(1), 13(?), 1402). Metatho-

rax: hair 1(1), 2(2,2-3), 3(5,5-7), 4(3,2-4), 5(1,1-2), 6(1), 7(8,8-9), 8(?), 9(7-9),

10(1), 11(2-3), 12(1), 13(7). Abdomen: Segment I: hair 1(1), 2(1), 3(2,1-2),. 4

(7-9), 5(4,3-4), 6(2), 7(2,1-2), 9(2,2-3), 10(1), 11(2,2-3), 12(2,2-3), 13(1). Seg-

ment II: hair O(1), 1(1), 2(1), 3€1,1-2), 4(6,6-7), 5(3,3-5), 6(2), 7(4,1-4), 8(4,

4-5), 9(1); 10(1), 11(2,2-3); 12(€2,2-3), 13(2). Segment: II: ‘hair 0(1), 1(1,1-2),

2{1), 301), 4(2,2+3), 5(3),6(2), 7(5,3-6); 8(2,2-3),’ 9(1); 1061), 1 1022-3), 1262),

13(1), 14(1). Segment IV: hair 0(1), 1(1), 2(1), 3(2), 4(2), 5(3,2-3), 6(2), 7(7,

7-8), 8(2), 9(1), 10(1), 11(2), 12(2), 13(1,1-2), 14(1). Segment V: hair 0(1), 1

(1), 2(1), 3(1), 4(7,6-8), 5(3,3-4), 6(2), 7(6,6-7), 8(2), 9(1), 10(1), 11(3,2-3), 12

(1,1-2), 13(2), 14(1). Segment VI: hair 0(1), 1(5,5-6), 2(1), 3(1), 4(2), 5(3,3-4),

6(1), 7(5,4-5), 8(3,2-4), 9(1), 10(1), 11(3,2-5), 12(1), 13(?), 14(1). Segment VII:

hair O(1), 1(2), 2(1), 3(5,4-6), 4(1), 5(3,3-4), 6(10,9-10), 7(1), 8(6,6-7), 9(2,2-

3), 10(1), 11(3,2-3), 12(1), 13(2), 14(1). Segment VIII: Comb scales slender at

base and fringed only apically. Hair 0(1), 1(3,2-4), 2(1), 3(4,3-4), 4(1), 5(3), 14

(1). Siphon: Integument light brown. Index about 4.4-4.9. Pecten teeth 5(4-7),

similar to those of epitedeus. Hair 1(2), hair lad(3,2-3), hair lav(4,2-4); hairs 2.

6,7,9 all single; 8(2). Siphon: Gill about 0.76 of dorsal saddle length. Ventral mar-

gin of dorsal saddle not reaching alveolus of hair 1-X. Hair 1-X shorter than gill.

Hair 1(1), 2(6,6-8), 3(1), 4a(12,11-14), 4b(11,11-14), 4c(12,11-12), 4d(12,11-14),

4e(13,12-15), 4f(11,10-14).

SYSTEMATICS. This species is the southern member of the Pacific complex

of the group. As indicated under its close relative costaricensis, it can usually be

readily separated from that species in all stages except on external features of

the adults. Its distribution is probably more extensive than reported here.

BIONOMICS. All the collections of curiche are from large crabholes. This spec-

ies has been found associated with pseudes and Culex (C.) inflictus. Nothing is

known of its blood feeding habits.

70 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

DISTRIBUTION (fig. 6). Pacific coast of northern Colombia. Material examined:

71 specimens; 7 males, 9 females, 34 larvae, 21 pupae; 19 individual rearings (9

larval, 5 pupal, 5 incomplete).

COLOMBIA. EI Choco: Curiche (COA 10,15,17,43,48,50) [BM, UCLA, USNM].

PSEUDES GROUP

FEMALES. Medium-sized species, wing 3.0-3.4 mm. Mesonotum dark brown;

pleural integument usually tan or brownish and not strongly contrasting with meso-

notal integument. Antenna: Flagellar segment 1 with scales, length varied, about

equal to combined length of segments 2-4 or 2-5; segments 2-12 subequal in length

but progressively slightly shortened distad; length varied, exceeding proboscis from

base of flagellar segment 9 to distal half of 7. Thorax: Postnotum without bris-

tles or with bristles in middle (usually in mcdonaldi). Apn usually tan to brown-

ish; ppn usually light tan to brownish, scales broad on caudoventral area; ppl us-

ually light tan to brownish; psp, ssp, stp and pra usually brownish; paratergite

usually brownish; mep usually lighter than stp, and with a patch of translucent

scales, Imep with 1 very strong bristle; meron, metameron and metapleuron us-

ually lighter than stp. Legs: Coxal integument usually lighter than stp. Only pos-

teroventral two-thirds of hindfemur pale, other femora dark; anteroventral mar-

gin of forefemur with or without spiniforms; when latter present then a row of

short spiniforms near base of posterodorsal margin also present, when absent then

a row of bristles present on both anteroventral and posterodorsal margins. Ab-

domen: Sternites II,III or (usually) II-VI paler than tergites.

FEMALE GENITALIA. Sternite VIII with sclerotized distal band broad and

without or with only a few scales; caudal margin not produced into a submedian

ventral angle, this area without strongly differentiated marginal bristles. Tergite

IX usually with a few setae on each side. Tergite X poorly differentiated dorsally

and with a very small lateral sclerotization. Cercus with a few moderately strong

apical or subapical normal bristles.

MALES. Essentially similar to females. Antenna: Length varied, exceeding pro-

boscis from base of flagellar segment 5 to middle of segment 3; flagellar segments

1 and 2 to 1-4 with scales; segments 1-7 markedly elongated but progressively

shortened distad; segment 1 subequal to combined length of segments 2 and one-

half of 3; segments 8-13 subequal but progressively slightly shortened distad; seg-

ment 13 sometimes slightly broader than 12. Legs: Claws of foreleg strongly en-

larged, similar in shape, either slender or heavy, usually both with a very minute

submedian projection or a very minute slender subbasal tooth; claws of midleg

slightly enlarged, similar in shape and simple.

MALE GENITALIA. Segment IX: Tergite lobe cylindrical; angled laterad at base;

apex always at least reaching base of subapical lobe. Sidepiece: Without scales.

Apicosternal lobe prominent, with a long apical seta. Phallosome: Dorsal para-

meres with an incomplete dorsal bridge; articulated ventrally with a dorsal scler-

otized projection from basal third of aedeagus.

PUPAE. Cephalothorax: Hairs 2,3-C closely approximated, 2-C weaker and short-

er than 3-C; hair 5-C double, strongly developed, longer than distance from its

alveolus to base of trumpet, about 1.3-1.5 of trumpet length; 7-C usually double ©

or triple (1-3); hair 8-C usually single (1 or 2). Metanotum: Hair 10-C moderately

to strongly developed, equal to or longer than 11-C, variously branched, usually

Adames: Genus Deinocerites 71

double or triple (in howardi and mcdonaldi) to markedly multibranched (pseudes).

Abdomen: Hair 1-II not reaching apex of tergite III, usually multibranched (3-

10 primary branches), if single or double then markedly secondarily dendritic; 1-

III-VI branched; 5-II always laterad of 3-II; hair 3-V often single (1 or 2); hair

1-VII shorter than tergite VIII, at most reaching base of 4-VIII. Paddle: Hair 1-

P subequal to longer than paddle.

FOURTH INSTAR LARVAE. Head: Hair 2-C at least subequal to 0.5 of 1-C,

always markedly mesad of 1-C; hair 5-C usually with 4 barbed branches (3-5);

hair 6-C single to triple, barbed, length varied, about 1.2 to 1.4 of 5-C. Tho-

rax: Prothoracic hair 9-P single. Abdomen: Hair 6-II] double; dorsal sensillum of

segment V laterad of 4-V; hair 6-VI single; 1-VII moderately to strongly devel-

oped, always longer than 3-VII and exceeding base of segment VIII, sometimes

reaching base of siphon; 1-VIII usually with 5 branches (2-7). Siphon: Hair 1-S

usually triple (2-4). Anal Segment: Ventral brush (4-X) with 6 or 7 pairs of hairs.

Gill short to moderately long, either round and slightly emarginate on apex or

more or less cone-shaped and slightly emarginate on ventral margin. Length var-

ied, about 0.45 to 0.9 of dorsal saddle length. Dorsal saddle narrow to moderately

broad, its ventral margin sometimes near base of hair 1-X.

DISCUSSION. The Pseudes Group is proposed here for pseudes, howardi and

mcdonaldi which were included by Belkin and Hogue (1959) in their Epitedeus

Group. The new group contains the most complex and intriguing elements in the

genus. The Pseudes Group shares with the restricted Epitedeus Group the devel-

opment of a patch of mesepimeral scales in the adults but is distinguished from

the latter by the presence of a strongly differentiated lower mep bristle. Its im-

mature stages resemble those of the Cancer Group and their separation from that

group is sometimes rather tenuous. |

The group is known only from the Pacific basin except for the Atlantic intru-

sion of pseudes into the Gulf of Mexico. It is the only group whose species are

not allopatric in distribution. All 3 have been found in 1 locality in the Bahia

de Banderas area near the mouth of the Gulf of California.

Of the 3 included species, mcdonaldi appears to be the most primitive as it

lacks the spiniforms of the forefemur in the adults (present in the other 2 spec-

ies) and has 7 pairs of hairs in the ventral brush of the larva. This species ap-

pears to be restricted to the region of the Gulf of California in Mexico on the

mainland as well as the tip of Baja California. Deinocerites pseudes is the mod-

ern dominant species of the entire genus with a distribution extending from the

Gulf of California and the Gulf of Mexico to the Gulf of Guayaquil. It is thus

the only species in the genus definitely known to be present in both the Atlantic

and Pacific basins and it is the only one to show considerable variation especially

on the periphery of its range. The third species, howardi, shows a remarkable mix-

ture of morphological characters of different stages of mcdonaldi and pseudes and

may have been produced through hybridization or introgression between these spec-

ies aS suggested by Belkin and Hogue (1959:436). The range of howardi is now

known to extend from Bahia de Banderas, Mexico, to Nicaragua.

Two of the species, mcdonaldi and howardi, appear to be rare or uncommon,

but pseudes is the most common species of the genus. Nothing is known of the

bionomics of mcdonaldi or howardi but considerable information has been gath-

ered on pseudes which has been colonized in Panama. The latter species will bite

man and has been shown to have a wide variety of blood hosts including mam-

mals, birds, lizards and amphibians. The immature stages of all 3 species have been

72 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

found primarily in large crabholes (Cardisoma and Gecarcinus); pseudes has also

been recorded from the smaller holes made by Ucides and even Uca.

16. Deinocerites pseudes Dyar & Knab

Figs. 7,50-52

1909. Deinocerites pseudes Dyar and Knab, 1909:260. TYPE: Lectotype female, Ancon, Pan-

ama, Canal Zone, A.H. Jennings, 378 [USNM, 12053; designation of Stone and Knight,

1957:197]. :

Deinocerites pseudes of Howard, Dyar and Knab (1915:210-213); Dunn (1917:167-168); Dyar

(1923a:180; 1925:155; 1928:263-264,537); Bonne and Bonne-Wepster (1925:176); Edwards

(1932:222); Martini (1935:57); Kumm and Zuniga (1942:406); Arnett (1950: 107); Carpenter

and Peyton (1952:677-681); Lane (1953:556,557); Horsfall (1955:599); Vargas (1956:30);

Stone and Knight (1957:197); Forattini (1958:176); Stone, Knight and Starcke (1959:285);

Diaz Najera (1963:190,191); Peyton, Reinert and Peterson (1964:449-458); Galindo (1967:

187-190); Grayson, Shrihongse and Galindo (1967:204); Stone (1967:218); Hogue and Wirth

(1968:6); Gorgas Memorial Institute (1970:20); Tempelis and Galindo (1970); Reinert (1970).

Culex (Deinocerites) pseudes of Dyar (1918:101).

Deinocerites cancer in part of Knab (1906:95-97).

Deinocerites epitedeus of Rueger and Druce (1950:60,62); Breland (1956:95); Evans (1962:255);

Vargas (1956:30; 1960:342).

Deinocerites epitedeus in part of Lane (1953:559, U.S.A. record); Forattini (1958:176).

FEMALE (fig. 50). Wing 2.24-3.4 mm. Proboscis 1.93-2.29 mm. Forefemur 1.53-

1.98 mm. Abdomen about 2.55-3.52 mm. Highly variable in size. Head: Narrow

decumbent scales creamy; erect scales yellowish to brownish; broad decumbent

scales of lateral patch whitish. Antenna: Torus without scales; flagellar segment

1 equal to combined length of segments 2-4 or 2 to basal third of 5; exceeding

proboscis from base of flagellar segment 9 to base of segment 8. Thorax: Post-

notum without bristles; ppn with several bristles in lower part; ppl with numer-

ous bristles and occasionally a few scales; mep with translucent scales usually cov-

ering most of the sclerite, also with numerous microsetae; umep bristles numer-

ous; metameron simple. Legs: Anterior surface of forecoxa largely covered by bris-

tles and scales, basolateral area with several bristles; midcoxa with scales antero-

laterally, laterally with 2 rows of bristles, the anterior row more or less irregular,

posterior bristles stronger, posterolateral surface simple; hindcoxa with scales and

bristles, upper lateroposterior surface with scales, posterior surface with bristles;

forefemur with spiniforms in anteroventral and at base of posterodorsal rows.

FEMALE GENITALIA (fig. 50). Sternite VII usually without scales, rarely with

1. Tergite IX occasionally with 1 seta on each side. Cercus (in lateral aspect) with

ventral margin convex, dorsal margin slightly sinuous in basal part and slightly

concave distally; apex slightly upturned, usually with 3 or 4 moderately strong

apical or subapical bristles. Distal part of postgenital plate (in ventral aspect) with

a very deep notch with parallel margins ending in a more or less round base; lat-

eral lobe (in lateral aspect) elongate, its apical bristle parallel with its mate (in

ventral aspect).

MALE (fig. 50). Wing 2.6-3.9 mm. Proboscis 2.2-3.0 mm. Forefemur 1.7-2.7

mm. Abdomen (not including genitalia) about 2.3-3.7 mm. Highly variable in size.

Flagellar segments 1-3 with scales. Proboscis reaching basal third of flagellar seg-

Adames: Genus Deinocerites 73

ment 4 to base of segment 5. Claws of foreleg heavy, usually with a very minute

submedian projection.

MALE GENITALIA (fig. 51). Segment IX: Tergite lobe extremely variable, us-

ually not extending beyond subapical lobe, with or without a distinct curvature

and expansion. Sidepiece: Subapical lobe with a large thumb; seta c spiniform

and usually rather heavy. Phallosome: Dorsal parameres with a very slight indi-

cation of a narrow incomplete dorsal bridge; apical spine short and broad; ven-

tral teeth short and heavy and on a prominent lobe which in tergal aspect pro-

jects about as far caudad as the apical spine. Aedeagus, as seen in tergal aspect,

with very sinuous outer margins, markedly constricted beyond middle and nar-

rowing apically; apex slightly swollen.

PUPA (fig. 51). Abdomen 4.0 mm. Trumpet 0.45 mm. Paddle 0.76 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 10 reared specimens.

Hair 5-III usually exceeding the spiracular sensillum and sometimes the alveolus

of hair 4 of the second segment following. Hairs 5-IV,V usually reaching and/or

exceeding alveolus of hair 4 of the second segment following. Cephalothorax: In-

tegument yellowish. Hair 1(2), 2(4,2-5), 3(2,1-2), 4(4,4-6), 5(2), 6(1), 7(2), 8(1,

1-2), 9(2,1-2), 10(multibranched), 11(4,2-4), 12(3,3-5). Trumpet: Short, progres-

sively widening apically; index about 2.75-3.28. Integument strongly pigmented,

contrasting with cephalothoracjc integument, slightly darker on tracheoid, brown

distad. Tracheoid about 0.3 and pinna about 0.26 of trumpet length. Abdomen:

Integument yellowish. Segment I: hair 1 (multibranched), 2(1), 3(2,2-4), 4(4,3-

5), 5(6,3-7), 6(1), 7(4,2-6), 9(1), 10(occasionally present, 1), 1 1(occasionally pres-

ent, 2). Segment II: hair O(1), 1(usually multibranched, if single or double then

markedly secondarily dendritic), 2(1,1-2), 3(1), 4(4-5,3-5), 5(1), 6(1), 7(2,1-3),

A(1). Segment III: hair 0(1), 1(5,3-6), 2(1), 3(2,1-4), 4(2,1-3), 5(1), 6(1), 7(2,

2-3), 8(4-5,2-6), 9(1), 10(2,2-3), 11(1), 14(1). Segment IV: hair 0(1), 1(3,3-4),

2(1), 3(4,3-5), 4(1), 5(1), 6(1,1-2), 7(2,1-3), 8(2,1-4), 9(1), 10(2,1-2), 11(1), 14

(1). Segment V: hair 0(1), 1(2,2-3), 2(1), 3(1,1-2), 4(4-5), 5(1), 6(1), 7(4,3-7),

8(2,2-4), 9(1), 10(1), 11(1), 14(1). Segment VI: hair O(1), 1(2,2-4), 2(1), 31,

be) (2. 32-3),.'5(1)), 601) 71), 8EL,1-3), 9G)y -100(151-2); 11(2,1-3), 14(1). Seg-

mont Vil: hair:001)5 101), 201), (2,142); 4p. 5(241-3)y: 6440-2) 97), 8(2,2-4),

9(4,3-5), 10(2,1-2), 11(1,1-2), 14(1). Segment VIII: hair O(1), 4(€1,1-2), 9(1), 14

(1). Paddle: Width about 0.77 of length; hair 1-P longer than paddle.

FOURTH INSTAR LARVA (fig. 52). Head 1.27 mm. Siphon 1.44 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 10 reared Specimens.

Length of hairs often variable throughout. Head: Integument light brown. Mental

plate wider than long, triangular; marginal spicules usually sharply pointed. Hair

O(1), 1(1), 2(1), 3(sometimes developed as a minute spicule), 4(6,5-8), 5(4,3-4),

6(usually single, except in the northernmost populations [Brownsville, Texas] and

| of the southernmost [Playas, Ecuador] in which they are single to triple, see

systematics), 7(10,9-12), 8(4,3-5), 9(5,4-8), 10(2,2-3), 11(4,4-5), 12(2,1-2), 13(6,

5-6), 14(1), 15(2,1-3). Antenna: Length about 0.46 of head; shaft with numerous

spicules on proximal part. Hair 1(6,5-7). Thorax: Prothorax: hair 0( 15,10-16), 1

(1), 2(1), 3(2,2-3), 4(5,4-5), 5(1), 6(1), 7(4,4-5), 8(1,1-2), 9(1), 10(1,1-2), 11(4,

3-5), 12(1), 14(1). Mesothorax: hair 1(1), 2(2,2-3), 3(1), 4(3,2-3), 5(1), 6(1), 7

(1), 8(7,5-7), 9(8,6-8), 10(1), 11(4,3-5), 12(1), 13(7-19), 14(11-14). Metathorax:

hair 1(2,1-2), 2(3,2-4), 3(6,4-7), 4(4,2-6), 5(1), 6(1,1-2), 7(9,8-10), 8(14-20), 9

(8,7-10), 10(1), 11(3,2-4), 12(1), 13(5,4-7). Abdomen: segment I]: hair 1€1), 2

tA), 3(3,2-3),° 4(9,8-13), 5(5,3-6), 6(2,2-3), 7(2,2-3). 9(3,2-3), 10(2,243), 1103,2-

74 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

4), 12(2,2-3), 13(2,1-2). Segment II: hair 0(1), 1(€1), 2(1), 3(2,2-3), 4(7,5-7), 5

(4,4-5), 6(2), 7(4,3-5), 8(4,1-4), 9(1,1-2), 10€1), 11(2,1-3), 12(2,1-3), 13(2). Seg-

ment III: hair O(1), 1(2,2-4), 2(1), 3€1), 4(2,1-3), 5(3,2-4), 6(2,1-2), 7(8,6-10),

8(2,1-3), 9(1), 10(€1,1-2), 11€2,1-2), 12(2,2-3), 13(1), 14(1). Segment IV: hair 0

Cy ty, 2C1) SSO), AU 2) S85), 0022-3) 18,69), 381) OCR), 20)

11(2,1-3), 12(2,2-3), 13(4,2-4), 14(1). Segment V: hair 0(1), 1(2,1-2), 2(1), 3

(1), 4(6,5-8), 5(4,2-5), 6(2), 7(7,7-10), 8(1), 9(€1), 10(€1), 11(2,2-3), 12(€1,1-2), 13

(4,2-4), 14(1). Segment VI: hair 0(1), 1(4,3-7), 2(1), 3(1), 4(4,3-5), 5(4,3-6), 6

(1,1-2), 7(3,2-4), 8(3,2-4), 9(1), 10(1), 11(2,2-4), 12(1), 13(2), 14(1). Segment

VII: hair O(1), 1(1-2), 2(1), 3(4,1-5),-4(1), 5(5,2-5), 6(16,15-20), 7(2,1-2), 8(10,

9-13), 9(3,2-4), 10(€1,1-2), 11(2), 12(1), 13(2,2-3), 14(1). Segment VIII: Comb

scales with fringe conspicuous at apex, less developed in distal half. Hair 0(1),

1(6,5-7), 2(1), 3(5,4-7), 4(1), 5(4,3-5), 14(2,2-3). Siphon: Integument light brown.

Index about 5.2-5.6. Pecten teeth 4(3-6), usually bifid, sometimes trifid, longest

blade markedly wider; size variable, basal tooth usually shortest. Hair 1(4,3-4),

hair lad(3,2-4), hair lav(3,2-4); hairs 2,6,7,8,9 all single. Anal Segment: Gill length

about 0.56-0.9 of dorsal saddle length; gill either round and slightly emarginate

on apex (usually in the northern and southernmost populations) or more or less

conical and slightly emarginate on ventral margin (usually in central populations,

see systematics). Dorsal saddle moderately broad and with its ventral margin near

base of hair 1-X only when gill is conical. Length of hair 1-X variable as com-

pared to gill length. Hair 1(1-2), 2(9,8-12), 3(1), 4a(15,14-17), 4b(12,12-16), 4c

(11,10-14), 4d(12,11-13), 4e(13,13-15), 4f(13,13-16). :

SYSTEMATICS. Deinocerites pseudes is the most widespread and the dominant

species of the genus, with a range extending from the Gulf of Mexico (from south

Texas) to the isthmus of Tehuantepec in the Atlantic basin and from the Gulf

of California to the southern shores of the Gulf of Guayaquil in northern Peru

in the Pacific basin. There are a few specimens apparently collected in the At-

lantic basin of Panama but these may be strays or erroneously labelled as no re-

cent specimens are known from this area.

The most characteristic and constant features of pseudes are the development

of the dorsal paramere and the.claws of the male, and of hair 10-C of the pupa.

In most other characters, including size, pseudes shows a great deal of variation

that is not found to any extent in any other species of the genus. The variability

reaches its peak in the peripheral populations in the north and in the south.

In the adults, there is very striking variation in size, even in the same collec-

tion. I found no correlation between size and any other variation noted in the

adults or the immature stages and consider that it is probably dependent on the

nutrition of the larva. Another striking variation in the adults is in the relative

length of the basal flagellar segments in both sexes. This variation affects greatly

the proportional length of the antenna and proboscis as noted in the description.

The immature stages show a great deal of variability also and tend to be con-

spicuously differentiated at the periphery of the range in the north and in the

south. It appears that in these peripheral populations there is a tendency for the

same phenotypes to predominate possibly because of the marginal or severe en-

vironmental conditions encountered by the species in these areas.

At the northern periphery of the range only the populations in the Atlantic

basin from Tampico northward show striking departures from the central popu-

lations. In the northernmost population in Texas, the larvae have hair 6-C usually

double (1-3) and only slightly longer than 5-C, whereas in the central popula-

Adames: Genus Deinocerites 75

tions it is apparently always single and markedly longer than 5-C. Abdominal hair

1-VII tends to be poorly developed and shorter than in the central populations

in which it is usually extremely long, often reaching the base of the siphon. The

siphon in these northern populations is uniformly short, whereas in central popu-

lations it is variable. The anal gill is more or less round and with a slight apical

emargination in the north while this condition is only rarely found in the central

populations which normally have the gill more or less conical and with a slight

ventral emargination. Specimens from Tampico show gills tending toward the type

in Texas while those from Veracruz are more like the central populations, sug-

gesting a clinal variation in the expression of this character. Both populations have

a moderately long siphon. The length of the pupal hairs tends to be slightly re-

duced in these northern populations.

Of the peripheral southern populations, I have studied 1 each on the north

and the south shores of the Gulf of Guayaquil. In the population from Playas,

Ecuador (north), the larvae are similar to those from Texas with hair 6-C usually

double and 1-VII, gill and siphon similarly developed. In the population from

Tumbes, Peru, the larvae are essentially the same as in Playas except that hair 6-

C is usually single and occasionally double but of the same relative length as in

the Texas and Ecuador populations. The pupae in both populations have the hairs

only slightly reduced in length.

BIONOMICS. Considerable additional information on the bionomics of this dom-

inant and widespread species has been published since the summaries in Howard,

Dyar and Knab (1915:212) and Belkin and Hogue (1959:449).

Knab (1906:95) reared adults of pseudes (as cancer) from immature stages col-

lected in the burrows of Cardisoma crassum at Rio Aranjuez near Puntarenas, Costa

Rica (336,339). Peyton, Reinert and Peterson (1964:451) recorded pseudes im-

matures from the holes of Uca subcylindrica, Cardisoma guanhumi and Gecarcinus

lateralis in several localities in Texas. Other specific records of immature stages

are from the burrows of Cardisoma crassum in Colombia (COA 23) and Costa

Rica (CR 28) and of adults only from burrows of Ucides occidentalis in Colom-

bia (COA 26). Other general records indicate that pseudes usually, but not al-

ways, breeds in large crabholes. Peyton, Reinert and Peterson (1964:453-454) re-

port that pseudes in Texas appears to prefer crabholes of a diameter of 2 inches

or more. These authors also recorded 2 unusual collections of immatures, 1 in a

grassy pool containing a crabhole in Texas, the other from a treehole in Mexico

(collection of R.B. Eads). They found the same range of pH (7.2-7.6) and chlor-

ide content (1,115-2,603 ppm) in the holes containing pseudes as in those uti-

lized by mathesoni.

Through its extensive range, pseudes has been found to breed in association with

a variety of mosquitoes: mathesoni in Texas (Peyton, Reinert and Peterson, 1964);

howardi in Nicaragua (NI 3,5,14,15,26,27); costaricensis in Costa Rica (CR 28,

106,107,118); curiche and dyari in Colombia (COA 10,17,25,27,28,48,50); 1 or

more members of the Culex (C.) inflictus complex in Costa Rica, Panama and

Colombia; Culex (Tin.) latisquama in Panama; Culex (Mel.) sp. in Costa Rica and

Panama; Aedes (O.) taeniorhynchus in Nicaragua and Anopheles (A.) eiseni in Co-

lombia.

Galindo (1967) colonized pseudes in Panama and found that the eggs were laid

singly above the water surface in a hollow cylinder of plaster of Paris simulating

a crabhole. Hatching took place without flooding 48-60 hours after oviposition,

larvae breaking out of the eggshell and either dropping or sliding down the sides

76 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

of the container to the water below. At a constant temperature of 76° F, the

larval cycle from hatching to pupation lasted 3 or 4 weeks and the pupal stage

3 or 4 days. In Texas, Peyton, Reinert and Peterson (1964:454) found that larvae

collected in the field required 2-242 months to complete their development and

the pupae 7 or 8 days. Furthermore, they noted that fourth instar larvae collected

during the latter part of November and in December did not pupate for at least

2 months whereas those collected in March pupate within 30 days or less. No

first instar larvae were found during the winter months.

Considerable information has accumulated regarding the blood feeding habits

of pseudes since Dyar’s (1925:155) statement that “the adults are not known to

bite’. The species has been taken in horse-baited traps in Costa Rica (Kumm,

Komp and Ruiz, 1940:392) and inside houses in El Salvador (Kumm and Zu-

niga, 1942:406). Peyton, Reinert and Peterson (1964:455) reported that in Texas

pseudes will readily bite man from dusk to midnight. They also noted that a num-

ber of specimens reported as Deinocerites sp. from horse-baited traps in Panama

(Blanton, Keenan and Peyton, 1955) were actually pseudes. Galindo (1967) found

Panamanian pseudes to feed readily on man in the field as well as the laboratory

and provided his colony with a source of blood from golden hamsters, guinea

pigs and domestic chicks. More recent investigations on the blood feeding habits

of Deinocerites in Panama (Gorgas Memorial Institute, 1970:20; Tempelis and Ga-

lindo, 1970) have shown that pseudes has a wide range of hosts including mam-

mals, birds, lizards and amphibians.

The mating behavior of pseudes in the laboratory is described in some detail

by Galindo (1967) who did not find either swarming or pupal attendance in this

species.

DISTRIBUTION (fig. 7). Pacific coast from Bahia de Banderas, Jalisco, Mexico,

to Gulf of Guayaquil in northern Peru; also Atlantic coast from Texas to state of

Campeche, Mexico. Material examined: 9285 specimens; 951 males, 1239 females,

6054 larvae, 1041 pupae; 567 individual rearings (375 larval, 141 pupal, 51 in-

complete).

COLOMBIA. El Choco: Curiche (COA 1,3,6-12,21,23,25-27,42-48,50) [UCLA]. Narino: Tu-

maco (PBR 989.65,67,68; 991.41; 994.8,9) [VALLE].

COSTA RICA. Guanacaste: El Coco (CR 201-212) [UCLA]. Samara (CR 195,198) [UCLA].

Puntarenas: Aranjuez, Rio, F. Knab (336) [USNM]. Boca de Barranca (CR 98,99,101-107,118,

233,235,237-244,248,251-254;LCBA2-4,12,104-106,111,112,115-118,122-124,126,135,137-143,

145,146,148 ,150,154,161,167,168,174,188,190,230); F.S. Truxal [LACM, UCLA]. Bonilla, Ha-

cienda, H.W. Kumm (75) [USNM] ; (CR 28,29) [UCLA] . Jicoral, H.W. Kumm et al (982) [USNM].

La Angostura (CR 111,115) [UCLA]. Puntarenas, H.W. Kumm et al (440) [USNM] .Rincon,Osa

Peninsula (CR 142) [UCLA]. San Lucas, Isla, H.W. Kumm et al (1117) [USNM].

lg Guayas: Playas, 7 km E, near Data (ECU 157) [UCLA] ; (LCBA 513,514) [LACM:

UCLA].

EL SALVADOR. Espiritu Santo, H.W. Kumm (570) [USNM]. Estero Ticuiclapa [? Ticui-

state], W.H.W. Komp (791, KO 32-29) [USNM].

GUATEMALA. Escuintla: San Jose [USNM].

HONDURAS. Choluteca: Trujillo, Rio Negro [USNM].

MEXICO. Campeche: Cuidad del Carmen, 36 km W (MF 14,15) [UCLA]. Guerrero: Puerto

Marquez, near Acapulco (MEX 142,144,145) [UCLA]. Zihuatanejo, A. Duges [USNM]. Jalisco:

Barra de Navidad (MT 1,4) [UCLA]. Puerto Vallarta [as Las Penas], A. Duges [UCLA, USNM].

Tabasco: Paraiso, near Puerto Ceiba [ISET]. Tamaulipas: Laguna de Chareil, Tampico (MEX

207A) [UCLA]. La Pesca (TEX 29,30,32) [UCLA]. Tampico (MEX 6-10) [UCLA] ; A. Martinez

[ISET] ; J. Goldberger [USNM]. Veracruz: Boca del Rio, near Veracruz (MEX 80,85-88) [UCLA].

Coatzacoalcos (MEX 113) [UCLA]. Nautla, A. Duges [USNM]. Tuxpan (TEX 22-27) [UCLA].

Veracruz, on steamer [USNM].

Adames: Genus Deinocerites a7

NICARAGUA. Chinandega: Corinto, P.A. Woke (779,793 ,794,806). K.R. Maxwell (14) [USNM].

Punta San Jose [as Monypenny Pt.], Bahia el Rosario, W.H.W. Komp [USNM]. Leon: Puerto

Somoza (NI 1,4,5,14,15,20,26,27) [UCLA]. Rivas: San Juan del Sur, P.A. Woke (KO 32-9)

[USNM].

PANAMA AND CANAL ZONE. Canal Zone: Albrook Field, W.H.W. Komp [UCLA, USNM] ;

(PA 737,743,934) [UCLA]. Ancon, A.H. Jennings (69,352,371,378,411,440,464); L. Roth

[USNM]. Cocoli, W.H.W. Komp [USNM]. Corozal, J. Zetek [UCLA, USNM]; L.H. Dunn;

W.H.W. Komp [USNM]. Ft. Sherman, L.H. Dunn, 1 M [USNM]. Gatunella River, J. Zetek

[USNM]. Howard Air Force Base (PA 468) [UCLA]. Monte Lirio, 2 F [UCLA, USNM]. Rod-

man Naval Station (PA 880,885-892,894,906-910) [UCLA]. Darien: El Real (PA 618,620,621)

[UCLA]. Jaque (PA 608,611,615) [UCLA]. Locality not specified: (55-1, 64-1) [UCLA] . Pan-

ama: El Libano, Corozal (PA 484,565,566) [UCLA]. Juan Diaz (PA 569,1018-1023) [UCLA].

La Chorrera (ASM 11-1). Nueva Gorgona (PA 998,1001) [UCLA]. Panama (49-57-1) [UCLA].

L.H. Dunn [USNM]. Perlas, Archipielago de las: Contadora, Isla, GML (PA 1045- 1047) [UCLA].

San Jose, Isla, J.P.E. Morrison [USNM]. Tobaga, Isla: A. Busck [USNM].

PERU. Tumbes: Tumbes, Playa Hermosa (PER 15) [UCLA]; (LCBA 505-510) [LACM].

TEXAS. Cameron County: Boca Chica Beach, E.L. Peyton [USNM]. Laguna Vista (MT 10)

[UCLA]. Stell-Lind Banco, E.L. Peyton, J.F. Reinert and N.E. Peterson [USNM]; (TEX 1-7,

12) [UCLA].

17. Deinocerites howardi Belkin & Hogue

Figs. 7,53-55

1959. Deinocerites howardi Belkin and Hogue, 1959:441-442. TYPE: Holotype male, Las Penas

(Puerto Vallarta), Jalisco, Mexico, 10 May 1903, A. Duges [USNM, 64264].

Deinocerites howardi of Stone, Knight and Starcke (1959:284); Ross (1962:192).

FEMALE (fig. 53). Wing 3.16 mm. Proboscis 2.16 mm. Forefemur 1.88 mm.

Abdomen about 3.11 mm. Essentially similar to pseudes except for antenna which

exceeds proboscis from at least distal third of segment 7.

FEMALE GENITALIA (fig. 53). Sternite VIIT usually with a few scales. Ter-

gite IX with more than 2 setae on each side (3-7). Cercus (in lateral aspect) bas-

ically similar to that of pseudes. Distal part of postgenital plate basically as in

pseudes except margins of notch converging to a round base; apical bristle of lat-

eral lobe slightly converging basally, then parallel with its mate (in ventral aspect).

MALE (fig. 53). Wing 3.01 mm. Proboscis 2.29 mm. Forefemur 2.29 mm. Ab-

domen (not including genitalia) about 2.75 mm. Flagellar segments 1-3 with scales.

Proboscis at most reaching distal third of segment 3. Claws as in mcdonaldi.

MALE GENITALIA (fig. 54). Apparently indistinguishable from mcdonaldi.

PUPA (fig. 54). Abdomen 3.69 mm. Trumpet 0.56 mm. Paddle 0.66 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 10 reared specimens.

Hairs 5-III-V moderately long, usually not reaching the spiracular sensillum of the

second segment following. Cephalothorax: Integument yellowish. Hair 1(2,2-3), 2

(4,3-6), 3(2,2-3), 4(2,2-3), 5(2,1-2), 6(1), 7(2,2-3), 8(1,1-2), 9(2,1-2), 10(3,2-4),

11(3,2-3), 12(3,2-4). Trumpet: As in mcdonaldi; index about 4.0-6.0. Tracheoid

about 0.3 and pinna 0.12 of trumpet length. Abdomen: Integument yellowish to

medium brown. Segment I: hair 1(18,13-22 primary branches), 2(1,1-3), 3(1-2),

4(2,2-5), 5(5,3-5), 6(1,1-2), 7(4,3-5), 9(1,1-2), 10(occasionally present, 1), 11(oc-

casionally present, 2,1-3). Segment II: hair 0(1), 1(5,3-9 primary branches), 2(1,

78 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

1-2), 3(1), 4(4,2-5), 5(1), 6(1), 7(2,2-3), 9(1). Segment III: hair 0(1), 1(6,3-10),

AC), H2sie3), 421-3), SCL 1-2); 60), 13,1-3),. 8(3,2-5);9C1), T0(2,1-2),: MG),

14(1). Segment IV: hair 0(1), 1(6,4-7), 2(1), 3(4,2-5), 4€1,1-2), 5(1,1-2), 6(1),

7(2,1-3), 8(2,2-3), 9(1), 10(2), 11(€1,1-2), 14(1). Segment V: hair 0(1), 1(5,2-6),

2(1), 3(1,1-2), 4(4,2-5), 5(1), 6(1), 7(3,1-5), 8(2,1-3), 9(1), 10(2,1-2), 11(2,1-3),

14(1). Segment VI: hair 0(1), 1(3,1-4), 2(1), 3(2,1-2), 4(3,2-3), 5(1), 6(1), 7(1),

8(2,1-3), 9(1), 10(€1,1-2), 11€1,1-3), 14(1). Segment VII: hair 0(1), 1(1,1-2), 2

(1), 3(2,1-3), 4(1,1-2), 5(2,2-4), 6(1), 71), 8(2,1-3), 9(4,1-5), 10(€1,1-2), 11(1,1-

2), 14(1). Segment VIII: hair 0(1), 4(2,1-2), 9(1), 14(1). Paddle: Width about

0.82 of length; hair 1-P subequal to paddle.

FOURTH INSTAR LARVA (fig. 55). Head 1.11 mm. Siphon 1.52 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 7 reared specimens.

Head: Yellowish to light brown. Mental plate as broad as long, triangular; termi-

nal spicules long and filamentous, lateral shorter and denticulate. Hair 0(1), 1(1),

2(1), 3(sometimes developed as a minute spicule), 4(6,4-6), 5(4,4-5), 6(2-3), 7

(9,8-11), 8(3,2-5), 9(6,4-7), 10(2,1-2), 11(4,3-7), 12(2,2-3), 13(8,6-8), 14(1), 15

(2,2-3). Antenna: Length about 0.46 of head; shaft with numerous spicules on

proximal part. Hair 1(5,4-6). Thorax: Prothorax: hair 0(8,8-11), 1(1), 2(1), 3(1),

4(4,2-4), 5(1), 6(1), 7(3,2-4), 8(1,1-2), 9(1,1-2), 10(€1), 11(3,3-4), 12(1), 14(1).

Mesothorax: hair 1(1), 2(3,2-3), 3(1), 4(2,1-3), 5(1), 6(1), 7(1), 8(7,6-9), 9(7,6-

9), 1O(1), 11(3,2-4), 12(1), 13(12,9-12), 14(11,5-13). Metathorax: hair 1(1,1-2),

2(3,2-3), 3(5,5-6), 4(4,4-6), 5(1, 2), 6(1), 7(8,7-8), 8(11-14), 9(9,7-9), 10(€1), 11

(3,1-4), 12(1), 13(4,4-5). Abdomen: Segment I: hair 1(1), 2(1), 3(2,2-3), 4(7,7-

10), 5(3,1-5), 6(2,2-3), 7(2,1-2), 9(2,2-3), 10(1,1-2),-11(2,1-3), 12(1,1-2), 13(1,

1-2). Segment II: hair 0(1), 1(1), 2(1), 3(2,2-3), 4(6,4-8), 5(5,3-5), 6(2), 7(3,2-

4), 8(4,3-6), 9(1), 10(1), 11(3,3-4), 12(1), 13(11-13). Segment III: hair 0(1), 1

(2,1-2), 2(1), 3(1), 4(1), 5(3,3-5), 6(2), 7(5,3-6), 8(2,1-2), 9(1), 10(1), 11(2), 12

(2), 13(1), 14(1). Segment IV: hair 0(1), 1(2,1-2), 2(1,1-2), 3(3,1-3), 4(1), 5@3,

2-4), 6(2), 7(6,3-7), 8(1), 9(1), 10€1), 11(2,1-3), 12(1), 13(2), 14(1). Segment

V: hair O(1), 1(2,1-2), 2(1), 3€1), 4(4,4-5), 5(4,3-5), 6(2), 7(6,6-7), 8(1), 9(1),

10(1), 11€1), 12(2,2-4), 13(3,2-4), 14(1). Segment VI: hair 0(1), 1(3,2-3), 2(1),

3(1), 4(3,3-4), 5(4,3-4), 6(1), 7(3,2-3), 8(4,3-6), 9(1), 10(1), 11(2,2-3), 12(1), 13

(15,14-18), 14(1). Segment VII: hair 0(1), 1(€1), 2(1), 3(2,2-3), 4(1), 5(4,4-5), 6

(10,8-12), 7(1), 8(5,4-7), 9(2), 10(1), 11(2), 12(1), 13(1,1-2), 14(1). Segment

VII. Comb scales similar to those of mcdonaldi. Hair 0(1), 1(5,4-7), 2(1), 3(3,

2-5), 4(1), 5(3,3-4), 14(2,1-4). Siphon: Integument light brown. Index about 4.3-

5.6. Pecten teeth 5(2-6), in general similar to those of pseudes. Hair 1(3,3-4), hair

lad(3,2-4), hair lav(3,2-3); hairs 2,6,7,9 all single; 8(2,1-2). Anal Segment: Hair

1-X and gill length probably similar to mcdonaldi. Hair 1(2,1-3), 2(7,6-10), 3(1),

4a(11,8-12), 4b(8,7-11), 4c(9-10,7-12), 4d(8,6-10), 4e(9,7-9), 4£(9,8-11), 42(11,10-

14).

SYSTEMATICS. The nature or origin. of howardi is the most intriguing and

possibly the most significant problem in the genus. Belkin and Hogue (1959:442)

suggested that howardi was formed through hybridization or introgression between

the ancient mcdonaldi stock and the modern dominant pseudes because of the

peculiar combination in howardi adults of general external features (spiniforms

of forefemur and lack of postnotal bristle) indistinguishable from those of pseudes

and of genitalia and claws of the male indistinguishable from those of mcdonaldi

and because of the sympatric occurrence of the 3 species. I have studied the im-

mature stages of howardi (unknown to Belkin and Hogue) and have found them

Adames: Genus Deinocerites 719

to be so similar to mcdonaldi that separation of the 2 is sometimes not feasible

on the basis of the combination of characters in the keys. Belkin and Hogue (loc.

cit.) also noted that the antennae of both sexes were longer than in pseudes or

mcdonaldi, implying this to be a manifestation of hybrid vigor. I find that in

the female of howardi there are usually more numerous setae on IX tergite lobe

of the genitalia than in the other 2 species.

I tend to agree with Belkin and Hogue as to the probable origin of howardi

through hybridization for the reasons given below. First, the presence of some

larval characters of pseudes in some larvae of mcdonaldi (see) suggests that intro-

gression may occur occasionally between these 2 species. Second, such hybridi-

zation could occur in an isolated area marginal to both species (Bahia de Banderas)

where a hybrid might have a selective advantage. Should such a hybrid become

stabilized it would probably be reproductively isolated from both mcdonaldi and

pseudes, thus explaining the unique situation in Deinocerites of the sympatric oc-

currence of 3 members of the same group.

Alternative derivation of howardi either from mcdonaldi or pseudes or inde-

pendently of the others from a stock common to all 3 species would involve the

development of identical combinations of characters in different lines or acqui-

sition and subsequent loss of several characters in 1 line and would appear to be

a more tenuous and less probable explanation.

The problem of the origin of howardi might possibly be resolved in the future

through experimental hybridization and cytogenetic analysis.

BIONOMICS. The immature stages of howardi have been collected in large crab-

holes. In Mexico these were associated with Aedes (O.) taeniorhynchus and in

Nicaragua with taeniorhynchus and pseudes. Nothing is known of the blood feed-

ing habits of howardi.

DISTRIBUTION (fig. 7). Pacific coast from Bahia de Banderas, Mexico, to Nic-

aragua. Material examined: 196 specimens; 50 males, 25 females, 81 larvae, 40

pupae; 39 individual rearings (25 larval, 13 pupal, | incomplete.

EL SALVADOR. Estero Ticuiclapa [? Ticuistate] ,W.H.W. Komp (791, KO 32-29) [USNM].

MEXICO. Colima: Manzanillo, M. Rueda [ISET]. Guerrero: Puerto Marquez, near Acapulco

(MEX 144) [UCLA]. Jalisco: Barra de Navidad (MT 1) [UCLA]. Puerto Vallarta [as Las Penas],

A. Duges [UCLA, USNM].

NICARAGUA. Chinandega: Corinto, P.A. Woke (806) [USNM]. Punta San Jose [as Mony-

penny Pt.|, Bahia el Rosario [USNM]. Leon: Puerto Somoza (NI 3,5,14,15,26,27) [UCLA].

18. Deinocerites mcdonaldi Belkin & Hogue

Figs. 7,56-58

1959. Deinocerites mcdonaldi Belkin and Hogue, 1959:437-438. TYPE: Holotype male (UCLA

199-116) with associated larval and pupal skins, San Blas, Nayarit, Mexico, 26 June

1956, W.A. McDonald [USNM, 64263].

Deinocerites mcdonaldi of Stone, Knight and Starcke (1959:285).

FEMALE (fig. 56). Wing 3.01 mm. Proboscis 2.09 mm. Forefemur 1.73 mm.

Abdomen about 2.5 mm. Head: Narrow decumbent scales of vertex creamy; erect

scales yellowish to pale brown; broad decumbent scales of lateral patch whitish.

Antenna: Torus usually with 1 scale; flagellar segment 1 slightly longer than seg-

80 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

ments 2-4 combined; exceeding proboscis from base of flagellar segment 9. Tho-

rax: Postnotum usually with 1 bristle near middle; ppl with several bristles; mep

with a patch of translucent scales usually in upper two-thirds, body of sclerite

with a few microsetae; only a few umep bristles; metameron simple. Legs: Scaling

similar to pseudes except for upper lateroposterior scales on hindcoxa which are

absent; forefemur with bristles only in anteroventral and posterodorsal rows.

FEMALE GENITALIA (fig. 56). Sternite VIII with a few scales. Tergite IX

with 1-3 setae on each side. Cercus in lateral aspect basically similar to that of

pseudes. Distal part of postgenital plate basically as in pseudes except for shallower

notch and bristles of lobes converging (in ventral aspect). :

MALE (fig. 56). Wing 3.11 mm. Proboscis 2.24 mm. Forefemur 2.24 mm. Ab-

domen (not including genitalia) about 2.75 mm. Flagellar segments 1 and 2 with

scales. Proboscis reaching base of flagellar segment 4. Claws of foreleg slender,

with a very minute slender subbasal tooth.

MALE GENITALIA (fig. 57). Segment IX: Tergite lobe with distal part mod-

erately long, slender, reaching base of subapical lobe but not extending beyond;

slightly hooked apically. Sidepiece: Subapical lobe with a distinct thumb; seta c

spiniform and attenuated apically. Phallosome: Dorsal paramere with short, broad

incomplete dorsal bridge; apical spine moderately long, heavy and markedly curv-

ed; ventral teeth short, arising from the evenly convex ventrolateral border. Ae-

deagus with highly sinuous margins and with a subapical necklike constriction;

apex narrow and slightly emarginate.

PUPA (fig. 57). Abdomen 3.03 mm. Trumpet 0.61. mm. Paddle 0.66 mm. Di-

agnostic characters as in the key; general chaetotaxy based on 8 reared specimens.

Hairs 5-III-V moderately long, usually reaching and sometimes slightly exceeding

sensillum of the second segment following, 5-III usually slightly longer than 5-

IV,V. Cephalothorax: Integument yellowish. Hair 1(2,1-3), 2(3,2-5), 3(2), 4(2,1-

3), 5(2,1-2), 6(1), 7(2,1-2), 8(1), 9(2,1-3), 10(2,2-4), 11(2,1-4), 12(2,2-4). Trum-

pet: Short, slightly widening apically; index about 4.5-5.6. Integument brown dis-

tad, darker on tracheoid; contrasting with cephalothoracic integument. Tracheoid

about 0.33 and pinna about 0.12 of trumpet length. Abdomen: Integument yel-

lowish to medium brown. Segment I: hair 1(18,16-22, primary branches), 2(2,

1-3), 3(2,1-2), 4(5,4-6), 5(6,4-6), 6(1), 7(2,2-4), 9(1), 1O0(occasionally present, 1),

11(occasionally present, 1,1-3). Segment II: hair 0(1), 1(6,3-8), 2(1), 3€1), 4(4,

4-6), 5(1), 6(1), 7(2,1-3), 9(1). Segment III: hair 0(1), 1(4,4-6), 2(1), 3(2,2-5),

4(2), 5(1), 6(1), 7(3,3-4), 8(6,4-6), 9(1), 10(2), 11(€1), 14(1). Segment IV: hair

O(1), 1(4,2-5), 2(1), 3(4,3-5), 4(€1,1-2), 5(1), 6(1), 7(2,2-3), 8(2,2-3), 9(1), 10(2),

11(1), 14(1). Segment V: hair 0(1), 1(2,2-4), 2(1), 3(1,1-2), 4(5,4-6), 5(1), 6(1),

7(4,3-6), 8(2,2-3), 9(1), 10(€1,1-2), 11(€1,1-2), 14(1). Segment VI: hair 0(1), 1(@,

1-3), 2(1), 3(1,1-2), 4(3,2-4), 5(1), 6(€1,1-2), 7(1), 8(2,2-3), 9(1), 10(€1), 11(2,1-

2), 14(1). Segment VII: hair 0(1), 1(€1), 2(€1), 3(2,1-2), 4(1,1-2), 5(2,1-3), 6(1),

7(1,1-2), 8(3,1-4), 9(3,1-4), 10(1,1-2), 11(€1,1-2), 14(€1). Segment VIII: hair 0(1),

4(2,1-2), 9(1), 14(1,1-2). Paddle: Width about 0.82 of length; hair 1-P as long as

or slightly longer than paddle.

FOURTH INSTAR LARVA (fig. 58). Head 1.11 mm. Siphon 1.16 mm. Diag-

nostic characters as in the key; general chaetotaxy based on 7 reared specimens.

Head: Integument yellowish to light brown. Mental plate triangular; length vari-

able, either as broad as or broader than long, spicules long and usually filamen-

tous, sometimes lateral ones denticulate. Hair O(1), 1(1), 2(1), 3(sometimes de-

veloped as a minute spicule), 4(6,6-9), 5(4,4-5), 6(2), 7(10,8-11), 8(2,2-3), 9(5,

Adames: Genus Deinocerites 81

4-6), 10(2,1-2), 11(4,3-6), 12(2), 13(7,6-7), 14(1), 15(3,2-4). Antenna: Length

about 0.46 of head; shaft with a few spicules on proximal part. Hair 1(6,4-8).

Thorax: Prothorax: hair 0(9,8-10), 1(1), 2(1), 3(2,1-2), 4(4,3-5), 5(1), 6(1), 7(4,

2-4), 8(1,1-3), 9(1), 10(1), 11(4,3-5), 12(1), 14(1). Mesothorax: hair 1(1), 2(3,

2-3), 3(1), 4(2,2-3), 5(1), 6(1), 7(1), 8(8,8-9), 9(8,6-12), 10(€1), 11(3,2-4), 121),

13(11,10-13), 14(11,9-12). Metathorax: hair 1(1,1-2), 2(2,2-3), 3(5,4-7), 4(4,3-6),

S(1), 6(1), 7(9,7-10), 8(9-14), 9(9,8-10), 10(1), 11(3,2-4), 12(1), 13(5,5-9). Ab-

domen: Segment I: hair 1(1), 2(1), 3(2,2-3), 4(10,7-10), 5(3,3-4), 6(2,1-3), 7(2),

9(3,2-3), 10(1,1-2), 11(2,2-3), 12(2,1-3), 13(2). Segment II: hair 0(1), 1(1), 2(1),

3(2,1-3), 4(7,6-8), 5(4,4-6), 6(2), 7(4,3-6), 8(4,3-4), 9(1), 10(1), 11(4,2-4), 12

(1), 13€12,12-15). Segment III: hair 0(1), 1(2,1-2), 2(1), 3(1), 4€1), 5(4,3-5), 6

(2), 7(5,5-6), 8(2), 9(1), 10(1), 11(€2), 12(2), 13(1,1-2), 14(1). Segment IV: hair

O(1), 1(2), 2(1), 3(2,1-3), 4€1), 5(4,3-5), 6(2), 7(6,6-8), 8(1), 9(1), 10(1), 1102),

12(2), 13(2,2-3), 14(1). Segment V: hair O(1), 1(1,1-2), 2(1), 3(1), 4(6,4-6), 5

(4,3-6), 6(2), 7(7,6-8), 8(1,1-2), 9(1), 10(1), 11(2,2-3), 12(1), 13(3,2-4), 14(1).

Segment VI: hair 0(1), 1(3,3-4), 2(1), 3(1), 4(3,3-4, 5(5,3-6), 6(1), 7(3,2-3), 8

(3,2-4), 9(1), 10(1), 11(2,2-3), 12(1), 13(18,15-19), 14(1). Segment VII: hair 0

(1), 1€2,1-3), 21), 3(3,2-3), 4(1), 5(4,3-5), 6(12,9-12), 7(1), 8(6,5-8), 9(3,2-5),

10(1), 11(2), 12(1), 13(2,1-3), 14(1). Segment VIIT: Comb scales with slender

body, fringe conspicuous at apex, less developed in distal half. Hair 0(1), 1(5,

2-5), 2(1), 3(5,4-6), 4(1), 5(3,3-4), 14(2,2-3). Siphon: Integument light brown.

Index about 3.2-4.6. Pecten teeth 5(4-6), in general similar to those of pseudes.

Hair 1(3,2-4), hair lad(3,3-4), hair lav(3,2-4); hairs 2,6,7 and 9 all single; 8(2,

1-2). Anal Segment: Gill about 0.45 of dorsal saddle length. Hair 1-X usually sub-

equal to gill length. Hair 1(2,1-2), 2(9,8-10), 3(1), 4a(12,11-14), 4b(11,11-12),

4c(10,10-11), 4d(10,10-11), 4e(12,9-12), 4f(12,11-14), 49(12,8-15).

SYSTEMATICS. Deinocerites mcdonaldi is sharply differentiated from the other

members of the group in the adults by the presence of only simple bristles on

the forefemur instead of spiniforms. The genitalia and claws of its male are in-

distinguishable from howardi and it also shares with the latter species a ventral

brush of 7 pairs of hairs. I interpret mcdonaldi as the earliest derivative of the

Pseudes Group stock.

It has been suggested that howardi (see) arose through hybridization or intro-

gression between mcdonaldi and the modern dominant pseudes. That introgres-

sion between the latter 2 species may occur is also suggested in a sample of lar-

vae of mcdonaldi (MEX 476) which contains some specimens with hair 6-C single

and strongly developed and 1-VII long as in pseudes while all the other charac-

ters are typical of mcdonaldi.

The surmise of Belkin and Hogue (1959:438) that the population from Baja

California is not differentiated from that of the mainland of Mexico is confirmed

here on the basis of larvae, pupae and adults.

BIONOMICS. Immature stages of mcdonaldi have been collected most frequent-

ly in large crabholes. In several collections in San Jose del Cabo, Baja California,

Mexico, the species of crab inhabiting the burrows was identified as Cardisoma

crassum, the pH was found to be 7.6-8.0 and the chloride content ranged from

2,350-8,400 ppm. In Puerto Vallarta, Jalisco, Mexico, mcdonaldi was found as-

sociated with belkini, Aedes (O.) taeniorhynchus and a member of the Culex (C.)

inflictus complex. Nothing is known of the blood feeding habits of mcdonaldi.

DISTRIBUTION (fig. 7). Mouth of the Gulf of California on mainland and lower

part of Baja California. Material examined: 2506 specimens; 287 males, 272 fe-

82 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971 ef

males, 1506 larvae, 441 pupae; 183 individual rearings (76 larval, 62 pupal, 45

incomplete).

MEXICO. Baja California Sur: Isla Espirutu Santo, Bahia San Gabriel [LACM]. San Jose del

Cabo, W.G. Downs [USNM]; (MEX 389-393) [UCLA]. Jalisco: Puerto Vallarta [as Las Penas] ,

A. Duges [UCLA, USNM]; (MEX 460-484) [UCLA, USNM]; (LCBA 532) [LACM]. Nayarit:

San Blas, Matanchen (UCLA 199,204) [UCLA, USNM]; (MF 4,6) [UCLA]. Sinaloa: Mazatlan

(3176,3179,3325) [ISET].

Adames: Genus Deinocerites 83

REFERENCES CITED

Adames, Abdiel J. and C.L. Hogue

1970. Mosquito Studies (Diptera, Cuan XVII. Two new species of Deino-

cerites from Costa Rica. Amer. Entomol. Inst., Contrib. 5(1):9-20. 28 Jan.

Aiken, James

1907. Notes on the mosquitoes of British Guiana. Brit. Guiana Med. Annu. 1906

(1907):59-78.

1909. Notes on the mosquitoes of British ‘inanewielin Brit. Guiana Med. Annu. 1908

(1909): 1-25.

Aiken, James and E.D. Rowland

1906. Preliminary notes on the mosquitoes of British Guiana. Brit. Guiana Med.

Annu. 1905(1906): 13-38.

Anduze, Pablo

1941. Lista provisional de los zancudos hematofagos de Venezuela (Diptera:

Culicidae). Bol. Entomol. Venezolana 1:6-18.

Anduze, Pablo J., F. Pifano and E.G. Vogelsang

1947. Nomina de los artropodos vulnerantes conocidos actualmente en Vene-

zuela. Bol. Entomol. Venezolana 6: 1-16.

Arnett, Ross H.

1948. Notes on the distribution, habits, and habitats of some Panama culicines

(Diptera: Culicidae). N.Y. Entomol. Soc., J. 56:175-193.

1950. Notes on the distribution, habits, and habitats of some Panama culicines

(Diptera: Culicidae). N.Y. Entomol. Soc., J. 58:99-115.

Bachmann, Alex O. and O.H. Casal

1963. Mosquitos argentinos que se crian en aguas salobres y saladas. Soc. Ento.

mol. Argentina, Rev. 25:21-27.

Barreto, Pablo and V.H. Lee

1969. Artropodos hematofagos del Rio Raposo, Valle, Colombia. II. Culicidae.

Caldasia 10:407-440.

Bates, Marston

1949. The natural history of mosquitoes. N.Y., Macmillan. 379 p.

Belkin, John N.

1962. The mosquitoes of the South Pacific (Diptera, Culicidae). Berkeley, Univ.

Calif. Press. 2 vol.

1968. Mosquito Studies (Diptera, Culicidae). IX. The type specimens of New

World mosquitoes in European museums. Amer. Entomol. Inst., Contrib. 3(4).

69 p.

Belkin, John N., S.J. Heinemann and W.A. Page

1970. The Culicidae of Jamaica (Mosquito Studies, XXI). Amer. Ertomol. Inst.,

Contrib. 6(1). 458 p. (Also published as Inst. Jamaica, Bull. Sci. Ser. 20.)

Belkin, John N. and C.L. Hogue

1959. A review of the crabhole mosquitoes of the genus Deinocerites (Diptera:

Culicidae). Calif. Univ. Publications, Entomol. 14:411-458.

Belkin, John N., R.X. Schick, P. Galindo and T.H.G. Aitken

1965. Mosquito Studies (Diptera, Culicidae). I. A project for a systematic study

of the mosquitoes of Middle America. Amer. Entomol. Inst., Contrib. 1(2):

1-17.

84 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Belkin, John N., R.X. Schick and S.J. Heinemann

1965. Mosquito Studies (Diptera, Culicidae). V. Mosquitoes originally described

from Middle America. Amer. Entomol. Inst., Contrib. 1(5). 95 p.

Blanchard, Raphael

1905. Les moustiques. Histoire naturelle et medicale. Paris, de Rudeval. 673 p.

Blanton, Frank S.,C.M. Keenan and E.L. Peyton

1955. Mosquitoes collected in horse-baited traps in Panama during 1951 to 1953

inclusive as an index to malaria control. Mosquito News 15:39-42.

Bonne, Cornelis and J. Bonne-Wepster

1925. Mosquitoes of Surinam. Koninkl. Inst. Trop., Mededeel. 21. 558 p.

Bonne-Wepster, Jean and C. Bonne

1923. A list of mosquitoes from Dutch Guiana. Insecutor Inscitiae Mens. 11:

—123-127.

Branch, Nina, L. Logan, E.C. Beck and J.A. Mulrennan

1958. New distributional records for Florida mosquitoes. Florida Entomol. 41:

155-163.

Branch, Nina and E.L. Seabrook

1959. Culex (Culex) scimitar, a new species of mosquito from the Bahama Islands

(Diptera: Culicidae). Entomol. Soc. Wash., Proc. 65:216-218.

Breland, Osmond P.

1956. Some remarks on Texas mosquitoes. Mosquito News 16:94-97.

Busck, August

1908. Report on a trip for the purpose of studying the mosquito fauna of Panama.

Smithsonian Misc. Collect. 52:49-77.

Carpenter, Stanley J. and R.W. Chamberlain

1946. Mosquito collections at Army installations in the Fourth Service Command,

1943. J. Econ. Entomol. 39:82-88.

Carpenter, Stanley J. and W.J. LaCasse |

1955. Mosquitoes of North America (north of Mexico). Berkeley, Univ. Calif.

Press. 360 p.

Carpenter, Stanley J.,W.W. Middlekauff and R.W. Chamberlain

1946. The mosquitoes of the Southern United States east of Oklahoma and Texas.

Amer. Midland Natur. Monogr. 3. 292 p.

Carpenter, Stanley J. and E.L. Peyton

1952. Mosquito studies in Panama Canal Zone during 1949 and 1950 (Diptera,

Culicidae). Amer. Midland Natur. 48:673-682.

Coquillett, Daniel W.

1906. A classification of the mosquitoes of North and Middle America. U.S. Bur.

Entomol,, Tech. Ser. 11. 31 p.

1910. The type species of the North American genera of Diptera. U.S. Nat. Mus.,

Proc. 37:499-647.

Diaz Najera, Alfonso.

1963. Lista de mosquitos capturados en tres localidades del estado de Veracruz,

Mexico. Inst. Salubr. Enferm. Trop., Rev. 23:187-192.

1966. Mosquitos tropicales de Mexico. Rev. Invest. Salud. Publ. 26:57-64.

Dodge, H. Rodney |

1962. Supergeneric groups of mosquitoes. Mosquito News 22:365-368.

1963. Studies on mosquito larvae. I. Later instars of eastern North American

species. Can. Entomol. 95:796-8 13.

1966. Studies on mosquito larvae. II. The first stage larvae of North American

Culicidae and of world Anophelinae. Can. Entomol. 98:337-393.

Adames: Genus Deinocerites 85

Downes, J.A. |

1966. Observations on the mating behaviour of the crabhole mosquito Deino-

cerites cancer (Diptera, Culicidae). Can. Entomol. 98:1169-1177.

Dunn, Lawrence H.

1917. Report on the mosquitoes caught by hand in the Canal Zone from Febru-

ary 1, 1916 to January 31, 1917. Med. Ass. Isthmian Canal Zone, Proc. 10:

145-169.

Dyar, Harrison G. |

1905a. Remarks on genitalic genera in the Culicidae. Entomol. Soc. Wash., Proc.

7:42-49.

1905b. A synoptic table of North American mosquito larvae. N.Y. Entomol. Soc.,

J; 43:22-26.

1905c. Brief notes on mosquito larvae. N.Y. Entomol. Soc., J. 13:26-29.

1905d. Illustrations of the abdominal appendages of certain mosquitoes. N.Y.

Entomol. Soc., J. 13:53-56.

1905e. Brief notes on mosquitoes. N.Y. Entomol. Soc., J. 13:107-109.

1918. A revision of the American species of Culex on the male genitalia. Insecu-

tor Inscitiae Mens. 6:86-111.

1922. The mosquitoes of the United States. U.S. Nat. Mus., Proc. 62:1-119.

1923. The mosquitoes of Panama (Diptera, Culicidae). Insecutor Inscitiae Mens.

11:167-186.

1925. The mosquitoes of Panama (Diptera, Culicidae). Insecutor Inscitiae Mens.

13: 1015195.

1928. The mosquitoes of the Americas. Wash., Carnegie Inst. Wash. (Publication

387). 616 p. :

Dyar, Harrison G. and F. Knab

1906. The larvae of Culicidae classified as independent organisms. N.Y. Entomol.

Soc., J. 14:169-230.

1907a. On the classification of the mosquitoes. Can. Entomol. 39:47-50.

1907b. Descriptions of new mosquitoes from the Canal Zone. N.Y. Entomol.

p0c6J.157197-242.

1909. Descriptions of some new species and a new genus of Amercain mosqui-

toes. Smithsonian Misc. Collect. 52:253-266.

1918. The genus Culex in the United States (Diptera, Culicidae). Insecutor In-

scitiae Mens. 5:170-183.

Dyar, Harrison G. and R.C. Shannon

1924. The subfamilies, tribes and genera of American Culicidae. Wash. Acad.

Sci., J. 14:472-486.

Eads, R.B., G.C. Menzies and L.J. Ogden

1951. Distribution records of west Texas mosquitoes. Mosquito News 11:41-47.

Edwards, Frederick W.

1932. Diptera. Fam. Culicidae. Genera Insectorum 194. 258 p.

Evans, B.R.

1962. Survey for possible mosquito breeding in crayfish holes in New Orleans,

Louisiana. Mosquito News 22:255-257.

Fauran, Pierre

1961. Catalogue annote des Culicides signales en Guyane Francaise. Inst. Pasteur

Guyane Franc. Inini, Arch. Publication 465. 60 p.

Felt, Ephraim P.

1905. Studies in Culicidae. N.Y. State Mus., Bull. 97:442-497.

86 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Fisk, Frank W.

1941. Deinocerites spanius at Brownsville, Texas, with notes on its biology and a

description of the larva. Entomol. Soc. Amer., Ann. 34:543-550.

Fisk, Frank W. and J.H. LeVan

1941. Mosquito collections at Brownsville, Texas. J. Econ. Entomol. 33:944-945.

Fleming, Milton B. and R.D. Walsh

1966. Mosquitoes of the American Virgin Islands. Mosquito News 26:424-426.

Floch, Herve

1946. Rapport sur le fonctionnement technique de VI.P.G.1. pendant l’annee

1945. Inst. Pasteur Guyane Franc. Ter. Inini, Publication 125. 139 p.

Floch, Herve and E. Abonnenc

1945. Les moustiques de la Guadeloupe (II). Les genres Megarhinus, Aedes, Cu-

lex, Deinocerites, Mansonia et Wyeomyia. Inst. Pasteur Guyane Ter. Inini,

Publication 110. 48 p. |

1947. Distribution des culicines des genres autres que le genre Culex, en Guyane

Francaise. Inst. Pasteur Guyane Ter. Inini, Publication 148. 12 p.

Forattini, Oswaldo P. .

1958. ‘“‘Culicidae” que se crian em buracos de carangueijos (Diptera). Rev. Brazil

Biol. 18:175-179.

1965. Entomologia Medica. Vol. 3. Sao Paulo, Univ. Sao Paulo. 416 p.

Fox, Irving

1953. Notes on Puerto Rican mosquitoes including a new species of Culex. J. Par-

asitol. 39:178-181.

Fox, Irving and J. Maldonado-Capriles

1953. Light trap studies on mosquitoes and Culicoides in western eae Rico.

Mosquito News 13:165-166.

Galindo, Pedro .

1967. Preliminary observations on the colonization and bionomics of the crab-

hole breeding mosquito Deinocerites pseudes Dyar and Knab, 1909. Mosqui-

to News 27:187-190.

Gentry, James W., E.J. Gerberg and T.M. Hopkins

1970. Notes on colonization of Deinocerites cancer Theobald. Mosquito News

30:68-70.

Gerry, Bertram I. |

1932. Morphological studies of the female genitalia of Cuban mosquitoes. Ento-

mol. Soc. Amer., Ann. 25:31-75.

Gibson, William W. and J.S. Carrillo S.

1959. Lista de insectos en la coleccion eee ec de la Oficina de Estudios

Especiales, S.A.G. Mexico, Secretaria de Agricultura y Ganaderia. Folleto

Misc. 9. 254 p.

Giles, George M.

1902. A handbook of the gnats or mosquitoes. Ed. 2. London, Bale, Sahs and

Danielsson. 530 p.

Gorgas Memorial Institute

1970. Annual report of the work and operations of the Gorgas Memorial Labora-

tory. 41(1969). 35 p.

Gowdey, Carlton, C.

1926. Catalogus Insectorum Jamaicensis. Jamaica Dep. Agr., Entomol. Bull. 4.

114 p.

Adames: Genus Deinocerites 87

Grayson, Margaret A., S. Srihongse and P. Galindo

1967. Isolation of St. Louis Encephalitis Virus from Deinocerites pseudes in Pan-

ama. Mosquito News 27:204.

Hill, Rolla B. and C.M. Hill

1945. Catalogus Insectorum Jamaicensis. Supplement. A list of the mosquitoes

found in Jamaica. Jamaica Dep. Agr. 3 p.

1948. The mosquitoes of Jamaica. Inst. Jamaica, Bull. Sci. Ser. 4. 60 p.

Hogue, Charles L. and W.W. Wirth

1968. A new Central American sand fly breeding in crabholes (Diptera, Cerato-

pogonidae). Los Angeles County Mus. Contrib. Sci. 152. 7 p.

Horsfall, William R.

1955. Mosquitoes. Their bionomics and relation to disease. New York, Ronald

Press, 723 p.

Howard, Leland O., H.G. Dyar and F. Knab

1913. The mosquitoes of North and Central America and the West Indies. Vol. 2.

Plates. Wash., Carnegie Inst. Wash. (Publication 159). 150 plates.

1915. The mosquitoes of North and Central America and the West Indies. Vol. 3.

Systematic description (in two parts). Part I. Wash., Carnegie Inst. Wash. (Pub-

lication 159). p. 1-523. 1 Oct.

Johnson, Charles W.

1919. A revised list of the Diptera of Jamaica. Amer. Mus. Natur. Hist., Bull. 41:

421-449.

King, Willard V., G.H. Bradley and T.E. McNeel

1939. The mosquitoes of the southeastern States. U.S. Dep. Agr., Misc. Publica-

tion 336. 91 p. June.

Knab, Frederick

1906. Notes on Deinocerites cancer. Psyche 13:95-97.

1907a. A new genus and species of sabethid mosquito. N.Y. Entomol. Soc., J.

15:120-121.

1907b. Deinocerites again. N.Y. Entomol. Soc., J. 15:121-123.

Knight, Kenneth L. and R.W. Chamberlain

1948. A new nomenclature for the chaetotaxy of the mosquito pupa, based on a

comparative study of the genera (Diptera, Culicidae). Helminthol. Soc. Wash.,

Proc. 15:1-10.

Komp, William H.W.

1957. Copulation in crabhole mosquitoes. Entomol. Soc. Wash., Proc. 58:349-

got, 3) Jan.

Kumm, Henry W., W.H.W. Komp and H. Ruiz

1940. The mosquitoes of Costa Rica. Amer. J. Trop. Med. 20:385-422.

Kumm, Henry W. and H. Zuniga

1942. The mosquitoes of El Salvador. Amer. J. Trop. Med. 22:399-415.

van der Kuyp, Edwin

1948a. Mosquito records of the Netherlands Windward Islands. Amer. J. Trop.

Med. 28:747-749.

1948b. Mosquito records of Aruba and Bonaire. Amer. J. Trop. Med. 28:895-

897.

1953a. Mosquitoes from Curacao. Caribbean Med. J. 15:37-39.

1953b. Culicinae from Netherlands Antilles and some other Caribbean localities.

Stud. Fauna Curacao Caribbean Islands 4:144-148.

1954. Mosquitoes of Netherlands Antilles and their hygienic importance. Stud.

Fauna Curacao Caribbean Islands 5:37-114.

88 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Lane, John

1939. Catalogo dos mosquitos neotropicos. Bol. Biol. Ser. Monogr. 1. 218 p.

1953. Neotropical Culicidae. Vol. 2. Sao Paulo, Univ. Sao Paulo. p. 553-1112.

Lee, Vernon H. and P. Barreto

1969. Artropodos hematofagos del Rio Raposo, Valle, Colombia. II. Aspectos

ecologicos. Caldasia 10:385-405.

Maldonado-Capriles, J., W.F. Pippin and M.L. Kuns

1958. An annotated check list of the mosquitoes of Mona Island, Puerto Rico, and

the larva and male of Aedes obturbator D. & K. (Diptera, Culicidae). Entomol.

Soc. Wash., Proc. 60:65-68. |

Martini, Erich

1935. Los mosquitos de Mexico. Mexico, Dep. Salubr. Publica, Bol. Tec. (A) 1.

65 p.

Matheson, Robert

1944. Handbook of the mosquitoes of North America. Ed. 2. Ithaca, Comstock

Publishing Co. 314 p.

McGregor, Theodore and R.B. Eads

1943. Mosquitoes of Texas. J. Econ. Entomol. 36:938-940.

Middlekauff, Woodrow W. and S.J. Carpenter

1944. New distribution records for the mosquitoes of southeastern United States

in 1943. J. Econ. Entomol. 37:88-92.

Mitchell Evelyn G.

1906. Mouth parts of mosquito larvae as indicative of habits. Psyche 13:11-21.

1907a. Validity of the culicid subfamily Deinoceritinae. Psyche 14:11-13.

1907b. Mosquito life. N.Y., Putnam. 281 p.

Montchadsky, Alexander S. and I. Garcia Avila ;

1966. Las larvas de los mosquitos (Diptera: Culicidae) de Cuba. Su biologia y

determinacion. Poeyana (A) 28. 92 p.

Pazos, Jose H.

1904. List de moustiques de Cuba (Dipt.). Soc. Entomol. France, Bull. 1904:

134-135.

Perez Vigueras, Ildefonso

1905. Los ixodidos y culicidos de Cuba. Su historia natural y medica. La Ha-

bana. 579 p.

Peyton, E.L., P. Galindo and F.S. Blanton

1955. Pictorial keys to the genera of Panama mosquitoes. ree News 15:

95-100.

Peyton, E.L. and S.O. Hill

1957. Keys to the genera of larvae and adult mosquitoes of the United States.

Mosquito News 17:294-296.

Peyton, E.L., J.F. Reinert and N.E. Peterson

1964. The occurrence of Deinocerites pseudes Dyar and Knab in the United

States, with additional notes on the biology of Deinocerites species of Texas.

Mosquito News 24:449-458.

Porter, John E.

1964. Deinocerites cancer Theobald recovered from tree holes at Miami, Florida.

Mosquito News 24:222.

1967. A check list of the mosquitoes of the Greater Antilles and the Bahama and

Virgin Islands. Mosquito News 27:35-41.

Adames: Genus Deinocerites 89

Pratt, Harry D.

1956. A check list of the mosquitoes (Culicinae) of North America (Diptera:

Culicidae). Mosquito News 16:4-10.

Pratt, Harry D. and E.L. Seabrook

1952. The occurrence of Culex iolambdis Dyar in Florida and Puerto Rico, with a

description of the larva (Diptera, Culicidae). Entomol. Soc. Wash., Proc. 54:

(27-32.

Pratt, Harry D., W.W. Wirth and D.G. Denning |

1945. The occurrence of Culex opisthopus Komp in Puerto Rico and Florida

with a description of the larva (Diptera, Culicidae). Entomol. Soc. Wash.,

Proc. 47:245-251.

Pritchard, A. Earl and H.D. Pratt

1944. II. A list of mosquitoes of Puerto Rico. Public Health Rep. 59:232-233.

Pritchard, A. Earl, E.L. Seabrook and J.A. Mulrennan

1947. The mosquitoes of the Florida Keys. Florida Entomol. 30:8-15.

Provost, Maurice W. and J.S. Haeger

1967. Mating and pupal attendance in Deinocerites cancer and comparisons with

Opifex fuscus (Diptera, Culicidae). Entomol. Soc. Amer., Ann. 60:565-574.

Randolph, Neal M. and K. O’Neill

1944. The mosquitoes of Texas. Austin, Texas State Health Dep. 100 p.

Reinert, John F.

1970. Description of the pupa of Deinocerites pseudes (Diptera: Culicidae). Flor-

ida Entomol. 53:17-21.

Root, Francis M.

1922. Notes on mosquitoes and other bloodsucking flies from Porto Rico. Amer.

J. Hyg. 2:394-405.

1927. Note on the mosquito fauna of the Republic of Haiti. Amer. J. Hyg. 7:

463-469.

Ross, Herbert H.

1962. A synthesis of evolutionary theory. Englewood Cliffs, N.J., Prentice-Hall.

387 p.

1964. The colonization of temperate North America by mosquitoes and man.

Mosquito News 24:103-118.

Roth, Louis M. and F.N. Young

1944. Culex (Melanoconion) atratus Theobald 1 in Florida; a new continental North

American record, with notes on the other melanoconions of the southeastern

United States. Entomol. Soc. Amer., Ann. 37:84-88.

Rueguer, Myrtle E. and S. Druce

1950. New mosquito distribution records for Texas. Mosquito News 10:60-63.

Scott, Harold G.

1961. Mosquitoes: key to United States genera based on male genitalia (Diptera,

Culicidae). Entomol. News 72:243-245.

Senevet,. Georges

1936. Les moustiques de la Martinique. Inst. Pasteur Algerie, Arch. 14:123-134-.

1938. Les moustiques de la Guadeloupe (Mission 1936). Inst. Pasteur Algerie,

Arch. 16:176-190.

Shannon, Raymond C.

1931. The environment and behaviour of some Brazilian mosquitoes. Entomol.

Soc. Wash., Proc. 33: 1-27.

90 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

Stone, Alan

1961. A synoptic catalog of the mosquitoes of the world, supplement I. (Diptera:

Culicidae). Entomol. Soc. Wash., Proc. 63:29-52.

1967. A synoptic catalog of the mosquitoes of the world, supplement III. (Dip-

tera: Culicidae). Entomol. Soc. Wash., Proc. 69:197-224.

Stone, Alan and P. Barreto

1969. A new genus and species of mosquito from Colombia, Galindomyia leei

(Diptera, Culicidae, Culicini). J. Med. Entomol. 6:143-146.

Stone, Alan and K.L. Knight

1957. Type specimens of mosquitoes in the United States National Museum: VI,

Miscellaneous genera, addenda, and summary. Wash. Acad. Sci., J. 47:196-202.

Stone, Alan, K.L. Knight and H. Starcke

1959. A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae).

Wash., Entomol. Soc. Amer. (Thomas Say Found., vol. 6). 358 p.

Surcouf, Jacques M.R. and R. Gonzalez-Rincones

1911. Essai sur les dipteres vulnerants du Venezuela. Premiere partie. Dipteres

nematoceres vulnerants. Paris, Maloine. 320 p. |

1912. Dipteres piqueurs et suceurs de sang actuellement connus, de la republique

de Venezuela. Arch. Parasitol. Paris 15:248-314.

Sutil Oramas, Ezequiel and J. Pulido

1969. Hallazgo en Venezuela de Deinocerites spanius (Dyar and Knab) (Diptera,

Culicidae). Venezuela, Div. Malarial. Sanid. Amb., Bol. Inform. IX(3):119-125.

Tempelis, C.H. and P. Galindo |

1970. Feeding habits of five species of Deinocerites mosquitoes collected in Pan-

ama. J. Med. Entomol. 7:175-179.

Theobald, Frederick V.

1901a. The classification of mosquitoes. J. Trop. Med. 4:229-235.

1901b. A monograph of the Culicidae or mosquitoes. Vol. 2. London, Brit. Mus.

(Natur. Hist.). 391 p.

1903. A monograph of the Culicidae or mosquitoes. Vol. 3. London, Brit. Mus.

(Natur. Hist.). 359 p.

1905a. The mosquitoes or Culicidae of Jamaica. Kingston, Inst. Jamaica. 40 p.

1905b. Diptera. Fam. Culicidae. Genera Insectorum 26. 50 p.

1910. A monograph of the Culicidae or mosquitoes. Vol. 5. London, Brit. Mus.

(Natur. Hist.). 646 p.

Thompson, George A.

1947. A list of the mosquitoes of Jamaica, British West Indies. Mosquito News 7:

78-80.

Thurman, Ernestine B., J.S. Haeger and J.A. Mulrennan

1949. The occurrence of Aedes (Ochlerotatus) thelcter Dyar in the Florida Keys.

Mosquito News 9:171-173.

Torre y Callejas, S.L. de la, P. Alayo Delmau and M. Calderon Chapman

1961. Los mosquitos de Cuba. Soc. Cubana Hist. Natur., Mem. 25(2). 95 p.

Urich, F.W.

1913. Mosquitoes of Trinidad. Agr. Soc. Trinidad Tobago, Proc. 13:525-530.

Vargas, Luis :

1956. Especies y distribucion de mosquitos mexicanos no anofelinos (Insecta,

Diptera). Inst. Salub. Enferm. Trop., Rev. 16:19-36.

1960. Los mosquitos de Sonora en relacion con el programa de Encefalitis. Rev.

Mexicana Med. 40:338-345.

Adames: Genus Deinocerites 91

Weatherbee, Albert A.

1944. A note on the mosquito distribution records of Puerto Rico and of the

Virgin Islands. Puerto Rico J. Pub. Health Trop. Med. 19:643-645.

Wirth, Willis W.

1945. The occurrence of Culex (Melanoconion) elevator Dyar and Knab in Flor-

ida, with keys to the melanoconions of the United States (Diptera, Culicidae).

Entomol. Soc. Wash., Proc. 47:199-210.

oe ee

Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

FIGURES

Deinocerites cancer; female

Distribution of the genus Deinocerites

Distribution of the Spanius Group

Distribution of the Dyari Group

Distribution of the Cancer Group

Distribution of the Epitedeus Group

Distribution of the Pseudes Group

Deinocerites spanius; male and female heads and genitalia, female forefemur,

and male foreclaws and midclaws

Deinocerites atlanticus; male and female heads, female forefemur and geni-

talia, and male foreclaws and midclaws

Deinocerites atlanticus; male genitalia and pupa

Deinocerites atlanticus; larva

Deinocerites mathesoni; male and female heads, female forefemur and geni-

talia, and male foreclaws and midclaws

Deinocerites mathesoni; male genitalia and pupa

Deinocerites mathesoni; larva

Deinocerites belkini; male and female heads, female forefemur and genitalia,

and male foreclaws and midclaws

Deinocerites belkini; male genitalia and pupa

Deinocerites belkini; larva

Deinocerites dyari; male and female heads, female forefemur and genitalia,

and male foreclaws and midclaws

Deinocerites dyari; male genitalia and pupa

Deinocerites dyari; larva

Deinocerites barretoi; male and female heads and genitalia, female forefemur,

and male foreclaws and midclaws

Deinocerites nicoyae; male and female heads, female forefemur and genitalia,

and male foreclaws and midclaws

Deinocerites nicoyae; male genitalia and pupa

Deinocerites nicoyae; larva

Deinocerites cancer; male head, female head and thorax and wing

Deinocerites cancer; male claws, female legs and genitalia

Deinocerites cancer; male genitalia and pupa

Deinocerites cancer; larva

Deinocerites melanophylum; male and female heads, female forefemur and

genitalia, and male foreclaws and midclaws

Deinocerites melanophylum; male genitalia and pupa

Deinocerites melanophylum; larva

Deinocerites magnus; male and female heads, female forefemur and genitalia,

and male foreclaws and midclaws

Deinocerites magnus; male genitalia and pupa

Deinocerites magnus; larva

Deinocerites epitedeus; male and female heads, female forefemur and geni-

talia, and male foreclaws and midclaws

Deinocerites epitedeus; male genitalia and pupa

Deinocerites epitedeus; larva

Adames: Genus Deinocerites 93

Deinocerites panamensis; male and female heads, female forefemur and geni-

talia, and male foreclaws and midclaws

Deinocerites panamensis; male genitalia and pupa

Deinocerites panamensis; larva

Deinocerites colombianus; male and female heads, female forefemur and geni-

talia, and male foreclaws and midclaws

Deinocerites colombianus; male genitalia and pupa

Deinocerites colombianus; larva

Deinocerites costaricensis; male and female heads, female forefemur and geni-

talia, and male foreclaws and midclaws

Deinocerites costaricensis; male genitalia and pupa

Deinocerites costaricensis; larva

Deinocerites curiche; male and female heads, female forefemur and genitalia,

and male foreclaws and midclaws

Deinocerites curiche; male genitalia and pupa

Deinocerites curiche; larva

Deinocerites pseudes; male and female heads, female forefemur and genitalia,

and male foreclaws and midclaws

Deinocerites pseudes; male genitalia and pupa

Deinocerites pseudes; larva

Deinocerites howardi; male and female heads, female forefemur and genitalia,

and male foreclaws and midclaws

Deinocerites howardi; male genitalia and pupa

Deinocerites howardi; larva

Deinocerites mcdonaldi; male and female heads, female forefemur and geni-

talia, and male foreclaws and midclaws

Deinocerites mcdonaldi; male genitalia and pupa

Deinocerites mcdonaldi; larva

Ri eae earesenie *]

diapered Asn. Gite fiber ss

ie f viata! Selgathseh LENG. eth oak RS

Me are Rastigiabarabiions cciiiapeabe cate

me ’

. x Rese ev at 21 tee fie Pasha” bate Dit:

Debi peatigries lito otc trues Perec ‘eI

aids Seven a pA IETRY Date

WAR, raahexives arivegenietd +24

: "eirend : stounn Aro os

: ae thie Sdean Seri aien oreo

: ‘norte thik cara sue Tectia it rs ty 5 Nan : “xk Sage

} rust) rat Catburot hhireonterh «ka

dr minsinsimnbiieie vat: aes.

pita).

Ssodteen ee fy Haken ' rit seankery MEM ved a4

roy . hide meats icp eateett Bees:

i ate i apie igen ate Sete (a A. Teer

i\e thd > P *

en Beer

wee. RS tape bl Eten ven Brie!

ee. sees ne ge HE, ide opie Ty Shai Ned pean .

AL ea OPS OD re. ibe eat ay

1D u i

, » a

to [ge Se eae

| ¢

. ua ad

np Ai, oR Tih

Ke et Ray ay

, Fer aa ve!

i ie "ie ad

- ‘ rae ter)

iets ' y x y G at Ae he f et ad :

eater) ans ; | aii

Dabs Paes ; het ru : 7 Ah t ; "Ah, . “iy ¢

¥ thay va aN : fy Le - fety A + +

- t Le, fille Rat.) al ‘ F Fil ay i x ah) ine gar"

Fig. 1

DEINOCERITES

Ef .

“i

j BAL

4“) |

hb |

“fh

Wf |

ZN f

i |

Wy) |

mi |

| ANY,

hy) )

Vid )

rif

bi,

cancer

JA 736

Kingston &

St. Andrew

Jamaica ‘

Fig. 2

_ DEINOCERITES

SATIN ALALVLS

00s

oot

005

SYaLAWOTI

wonHe aad

s \ ? a

dNOUd SNINVdS

, pe ne

x Bes

SATIN ALALVLS

= SUTLIWOTOX i sap

VOIWHNV HICCIN i. ae Bia ees

10}0118q

1B Ap

bre |

Q 2 >. ef te

ae) rn RSS Se

SATIW ALALVLS

000

SYA LAWOTM

V Wi

wnyAqdourjow

SATIW ALMIV.LS

00s

000l

00st

0001 or

SYaLAIWOTI

> SYOIINI

fs 1

L~2= a :

: Le 4 JS RD, Nate

Os Bee

: C0? of |

POG sy

SNUBIquIO]OD

snopojzida 4”

—————|]

—

san0ud SMACaLIda

SATIW ALALVLS

00s

0001 00s

SYA LAWOTIA

VORMANV ATCGIN =~

d sad

DEINOCERITES

PA 1044

Cocle anterior posterior

Panama

FORE

PA 1041

ea Panama

Panama

he PA 1044

Cocle

y Panama

anterior posterior

MID

PA 1039

Panama 0. J -_--__—}

Panama

PA 1036

Panama

PA 1039

Panama

VV AIS

i\d

Nie rir an id

1 Nae Se? fu gy)

PA 1039

Q Panama

Panama

° ° ° °

spanius posterior

-————]—_——————_ i —1.0—————————oI4

SS y

anterior | posterior

FORE

anterior posterior

MID |

-——————0.1 —___——__+

g — anterior

atlanticus

VZ 170 teri

Aragua posterior

Venezuela

-——_—_—__ 1.0 —_________{

Fig.10

—

\\ HF Ch })

i Me Te

AN WY) i YY

HAN \, 4

——

VAS

‘

AU eas

aa VN ESO

Sk NERO

7 SM Ee

”

atlanticus

VZ 170

Aragua

Venezuela

atlanticus

VZ 170 Aor

Aragua

Venezuela

DEINOCERITES Fig.12

—— ee

7 a

—>

———

——S

SSS SSE

—~

—

———

———S——

SSS

iN ,

Wy hiihva\

Hi »

| AN

AN |

iN :

| i anterior posterior Q

Gi HON

i WN .

i ANN

i A

a — HAN FORE fle’ NANG,

: y ‘Ly By!

Re\

anterior | posterior

MID

pon Dl

SS 2 SS Sa SSS

His

—S

ZFS

LD”

ZZzZz

GSA

giao

—

;

mathesoni |

posterior

TEX 19

Texas

HK 1.0 AN ——_

U.S.A.

peinocerites Fig.13

mathesoni

2. 14

DEINOCERITES

tyyy

i Y yf!

Vy)

mathesoni

| AHN

Vj | My a

peINocerites Fig. 15

ees Ss

INN

HIN

HIN anterior posterior

i '

i A

f ak vf

Hi} \ ft N\

Hf \ \ aN AN

Uf AN UN

A Di VAY FORE

1 N HY YM \\

H iN

}

4 } ; anterior posterior

WIN

(ANN

WKN \

W \\

AIM

i )\4

K MID

Sood

I Zz

———

Le ZA

EELLAz

—— a

SOL CH”

==>

—

——

SSS

——_

—S—

Sas

————

Hef;

\ belkini

MT 4

Jalisco

Mexico

posterior

Ferseseereerinmmetiteta at pte AA) ge ee ne

DEINOCERITES Fig. 16

belkini

MT 4

Jalisco

Mexico

bel kini

P 1 10) 11% \g

6 5 | i! i MT 4

Jalisco

Mexico

DEINOCERITES

WPS « 4

= —— PSF

SSNS

—<—.

SSSQ

j PA 1030

iy \ Panama

S\ Panama

‘“

HAN

fq PA 1030

Panama

BY Panama

y anterior posterior :

Hy Q

YAN

R\\\ ' —— FORE

iy,

yy i

Yi\\

A * et

A PA 1030

Panama

! Panama

pee

fj anterior posterior

h z

I é W\\

9 eta iN Canal Zone [fi MID ie ;

) ZN beri Panama Hi bese etc eatin Net as,

iy \

ps — f \ ‘NK r————————_ 0.1 _ -—uuwuWwH 0.1——_______}

h ) i) Ki

Vy pei CM ALIN

I 5 EM

i \

ee Br —

aN. —

DN —

j Canal Zone

fh NO Panama

a ae

posterior

dyari :

1,9 —______,

. DEINOCERITES

\ a

J, SH?

ip-° Z

MA Canal Zone

y) Panama

| keira

Fig. 19

COA 24

El Choco

Colombia

ig. 20

~ 2 ang

S Sune gs

“ Es)

S so:

fo ORS

~~ w

{/ 4

‘

DEINOCERITES

4 \\\

ENN

}

DEINOCERITES Fig. 71

anterior posterior

FORE

\ ‘Nae

\\\ et COL 47g

1 N \"\}) Valle

iN \ Colombia

Hie \ ‘ot

ih \N

eS Z ZG

anterior posterior

==>

i \

) COL 47g Ny

y \ Valle WN MID

i Sh actpiacincineinsnnets i, msicenelabsiimenrsprincel

j 0.5

f ate COL 47a

Valle

Colombia

SSS

COL 47f

Valle

Colombia

ae, —— Sa —

SS—Z Za F

SZ”

Sasa

3 SSS SS

SS) SS SS SS

(27

———

SSS SSS SSS

SSS

COL 47c

Valle Q

| \ Colombia

a ANN

COL 47c a

1.0 \ Valle

\ Colombia

ees,

is)

Valle

te barretoi Salar

posterior

1 1.0 — _—_—__f

anterior posterior

FORE

anterior | posterior

MID

-——————— 0.1———¥1

ee ee

posterior

nicoyae

LCBA 188

Puntarenas

Costa Rica

pe SLY

Fig. 23

CR 254

Puntarenas

Costa Rica

DEINOCERITES

1.0

yy Yt NN o, y a

ES Ay

CR. 3

Puntarenas

Costa Rica

nicoyae

‘sy

°o

QU] | EX <

Boer /

nicoyae

CR 254

Puntarenas

Costa Rica

SSS oS

~ g = Zz LEZ

y SSS

CIP Rex

NNN YUNA

VAT NAAR

NRSTRREARE Ss

i INN \ \\ \\ IR Nine . TT —" SAN iy

WINK Mii ‘ AAW AND

\ AY

meniiearan > romarre eee

—

= —=_

DEINOCERITES

——— <tzzue

SES

(ty vi I

Mi mi \

———

=——

—

SSS

SSSA. 2

\ M

HlivN P

HN uN

f aK \ t}

ji ;

in wn \

f \

A\\

Ya bac

AN aly

1.0 7 \ oan

ZZ ey

2 ye >

SS S

PS gS

cancer

JA 736

Kingston &

St. Andrew

Jamaica

VIII tergite

DEINOCERITES Fig. 26 \

cercus

post genital plate

cowl

VIII sternite

spermatheca

Wh yy HRCA

NTS

ig (ight ait.

. V4

insula

is ae ge ana an

ca n Cc er anterior

| g

JA 736

Kingston &

St. Andrew —

Jamaica

SMS oe ae

1.0 posterior

cancer

JA 705

Kingston &

St. Andrew

Jamaica

DEINOCERITES Fig.28

SASS QO

Sk AY

QQ |e AW e

=~ , - =\

= Be \\ |}

Y::4

=< x 00 4

cancer

JA 705

Kingston &

St. Andrew

Jamaica

DEINOCERITES

Canal Zone

Panama

Yr {

MK \

HN n\

it \

} ANN

A — a

| oN

hh HN

Kin fH

| “iN

NN Q

WIN

f\ i

i \ i \\ anterior posterior

fh i) N\

( WN

: ih |

[\\ ~~ Canal Zone Wi

M7) Wy \

Paxne —- e Na, We

in aN ‘ Udy \ \ PT

i ARM 1 Fe 7 1 gp A\O\\id &

H es i iN FORE Wild | a Mc és 1 \ \ ae \ ]

AN 7 + of Graeme weit y \ YI

iN pay: hee mee LTE ey

k .) cy" vi oh ll rae s ip Pe Pe

a re ae gt OE pe dat

Canal Z aed oles Wr tas i Amtek Ae

f ‘anal Zone i WN \ tah MAGN SBE

Ree Te Panama SAYS) ly fis Ilys fully deste

Pye eh pba

AN aha /

; "eA eran

\ anterior posterior iS

f}|\

UN — PA 1032

y \ Colon

Y\\ Panama

Yi MID

; lanophy!

i fret rial meranopnyium

i ae Y x

Hii ys SEN

\ i

iN —

j PA 1029 anterior

vie a 2 Colon

M Panama

posterior

reer 10 iat

ig. 30

DEINOCERITES

Colon

Panama

Canal Zone

SS Ee eat

PA 1032

Colon

Panama

eal

<>)

<2)

(am

SS) SSeS eS

———

——s ES

—>

St a

—

—— a

——

St. Vincent

Q\y

Cre Ss

1.0

yA SsS

—S SS

SS S SNS

pe SS ee

> Sets es Se

SS SP SSS eat

LLL

SSS

Martinique

St. Lucia

St. Lucia e

anterior posterior

FORE

wi \

i/ /»

Martinique

| { pe \ \ [1

Cie |

anterior posterior tay SEEDED yaaa ee

AN pL GUL ee

iN 1

-————_ 0.1—___

MANN 9

! HAN

AN MID

NIN

WAN prone |

v i

Q St. Lucia

oe er <

posterior

rR 1.0 ——______—__

DEINOCERITES

Pi, Ma

(ip I)

Fig. 33

Nariva Swamp

Trinidad &

Tobago

i=)

Martinique

magnus

—

Nariva Swamp

Trinidad &

Tobago

DEINOCERITES |

iii

DEINOCERITES F ig. 3D

, CR 76

\ Limon

t Costa Rica

posterior

FORE

anterior posterior

| a MID - \

————— 0,141

epitedeus

LCBA 234

Limon

Costa Rica

posterior

1.0

Fig. 36

DEINOCERITES

CR 76

Limon

Costa Rica

\ NAS 3555

Limon

Costa Rica

AN ig

iy ms \ |

epitedeus

DEINOCERITES Fig. 37

epitedeus

CR 76

WE Limon

Costa Rica

pernocerites Fig. 38

——

SSS Sa

SS? SFR SS - 2S? a Ss s = :

anterior posterior

OX)

\\ ——

} FORE

K os

4 anterior posterior

MID

x — Hu ee as ete

ff ee ean ni Me

His BARK Raney was ; eat Clee

| -+————0,1 ——__ ESE MN NONE, Caer

aX - ==

; es

\ H———_—_—_- 0.1 —____

2 Ste? SPD ) SES ES

ETS? SSSR

~panamensis

CAZES,

Canal Zone

Panama

posterior

KH 1.0 —_}

Fig. 39

DEINOCERITES

: PA 1016

\ Colon

aS Panama

1.0

(A)

ye . .

"7 * 0.1 Bate

| os , A ae 2 10 F

ee 144

ig. 40

panamensis

DEINOCERITES

‘ &

= %

Gy,

(i 4)

DEINOCERITES Fig. Al

COA 40

Q Antioquia

Colombia

| COA 40

j Antioquia

Colombia

anterior posterior

ty/ i

wy i

FORE ‘

| : O \ © \

\ \ |

Ss SS

Sen

SSS

——=

—— es

in ‘

iN h

COA 31 x fa

Antioquia “i ik\

Colombia VW)

ian ]

}

COA 31

Antioquia ‘et

: if ARI

| A \ 4 iM \

{ iN

" iN

My) :

Aya 4

) WN

| IN Colombia

4 AN

, COA 40 ff

| Antioquia A

) i AN

gid te ANY anterior posterior

TAN

y i

Q ian

| HNN

hy Uy

AY i a

oe SSSA SS . .

FAL.

—

SSS

Zz —

SSS

== ae

py + 0.1 4

\ 1.0 a

R Y

. ‘peg i]! \

i \

Hi —

] A

Ye — Ips ——

Y \ i

‘J

y aaa Has ieee

, | anterior

COA 40

Q Antioquia

Colombia

=} SS SI) SS

SSS ;

colombianus

posterior

SSS)

tH 1.0 -—-____—__X1

a eS

Fig. 42

) DEINOCERITES

COA 40

Antioquia

Colombia

COA 31

Antioquia 0

Colombia VI

COA 40

Antioquia

Colombia

colombianus

lav

. i

at Wise

COA 39

Antioquia

Colombia

colombianus

DEINOCERITES

SFR 20.

Ml! My

yf LH

—\pEINOcERITES Fig. 44

CR 34

Puntarenas

Costa Rica Ki

CR 142

Puntarenas

Costa Rica

CR 34

Puntarenas e

Costa Rica

anterior posterior

Puntarenas

Costa Rica

eee |

} |

— i FORE

\ CR 34

NA oon CO

At —.

Brin

rae ii f anterior posterior

ae i { \

fi i MID

AAT

PRES 1.0 [0,1 1

[

ay 5 Me (en ae ia.

HR anterior

Q Pinter

ntarenas

Cost Rica

ter

e e

posterior

-——_____—_——- 1.0 ——————____

Puntarenas

Costa Rica

CR 142

Puntarenas

Costa Rica

CR 28

ig. 45

I ' 3 3 |

DEINOCERITES

CR 28

Puntarena

5 or /

eh N Hy)

Vx) ‘ iB es

Zas. +3 ly,

flrs Ae sets X ‘

0 te Chr *

Gs Pts

sites

eps UO yj

a Eee 4

= .

te of) 9

Ss “7

'S oy. |

Seabee tao y |

, 4

Costa Rica

icensis

costar

COA 50

Q El Choco

Colombia

msn P05

j Y

TN | AN

AN A j; . \ \

Mi i} 1 ii

HAWN Hh, \ ae \ ;

M ) TN

i HN hl INN

tN in

OA HN

Hf Ii)

Hin iN

YAN ae

if K | i, i

— \

ft yf il

Aisi ih i anterior posterior

a) \

AN | i

A coazs = fil ¥hi)

An El Choco ii \

W\\ Colombia Hin aN

4 Bi AN

Ht aN HM

dik Wy — HN

fy ) en f i \

1 ) | P i Om

AN\ — iA

f B) COA 48 ef

it MAM El Choco

ny ALIN .

A Hn Colombia

hy — ii

COA 15 AN

ath El Choco BAY)

fk Colombia na

if HAAN

, iM anterior posterior

; Q MM A

Y Ris

y, —_ Mt

}

A - MID

SS ‘

SSSS3

-————0.1_______

> 1

SSS

ZZ,

+S Ss

SP» =

x ee ae

COA 50 anterior

Q El Choco

Colombia

S i = 5

Br.

SSS)

;

SSS

\ et eh ft ht LE

cur i c he posterior

<==

-———________ 1.0 —__———_____j

rane DEINOCERITES

(mh)

Fig. 48

COA 48

El Choco

Colombia

El Choco

Colombia

curiche

ig. 49

DEINOCERITES

Z 7,

SL, /

COA 48

El Choco

Colombia

curiche

COA 50

El Choco

Colombia

SS =

SS —S es =

SSS eae

Hee Or ge

= S S

SPs S

pseudes

EN |

SSS

SS SS

FORE

PA 569

Panama ‘os

Panama

anterior posterior

MID

PA 566

)

posterior

Ene eine nent, | pettec ee ae ae ee See

posterior

PPP PIPPI?

Panama = ee)

pseudes

ig. 52

Q A z

NAN

oNs S\N oN

a Ss

DEINOCERITES

pernocerites Fig. ~¥

BS SS OSS ES

SSS

ae eI fey

——— we

—

MT 1

2 Jalisco

Mexico

)

---———— 0.5

Jalisco

Mexico iy

UMA

‘tess i

iy MT 1

th Jalisco Q

Nk :

yp Mexico

Y MT 1

, Jalisco

. Mexico /

} yy

anterior ! posterior

; :

: : 4

‘ FORE

le \ \

{ Eg (eas a Kah \

! rae

i ar a aN eo Te ee he

p ane’ VyA !

; hin ‘a iss ;

N i \ ust \ > )

AW i - RAL 7 :

\ ar \ \ if |

N WW A Le PY yi?

hy anterior posterior Nae oe

\ h \ Ye ona}

va — a © \ Ae 3

I \ Nt RW RG raat j

\ 1.0 AWN Bee ta MR gh eo Re ee eee Ree he eel” RS ey eae ee 5) © eee os nl ltias ano x

/ r y i! I MID

F Jalisco {iW a4

4 Mexico Yih .

ik i mi #0. 1-1

t - ieee IN

hi Ak Q ANY

i / \ Ay

ys hana Rie

hs Dave i hy

‘

y y p

y iy

ae TS hi

i f anterior

Hes i} —

e Jalisco

i N Mexico

sy \.

fe He i ag EY P- Ves

: I howardi mg st

A owardl Lagan

b———————_—__—————1.0 —______1

Fig. 54

|, DEINOCERITES

11 10

a).

10 11

MT 1 ,

Jalisco

‘ Mexico Yi

, wn ar /:

RNS

BAN

Jalisco

Mexico

a howardi

a DEINOCERITES Pi se

SSS

i ga

= —SSSSS =

| IN anterior posterior

(

hs FORE

: Co

j iy

[i N\

\ Ae \ \__ uN

f i,{ ANN :

\ j HN

WYN Z

ih a \ Wn anterior posterior

‘ 2 iN

i Ws

1) A SIN

\ N MID

-#————_0,1 —_—___}

SSS SSS De ee De

: ee eee

AN a

A — :

iN mcdonaldi aaa,

PS posterior

4 UCLA 199

Nayarit

Mexico -———————1.0-_ —___

DEINOCERITES Fig,57

Wy;

| WD

WW .

\ \ ‘

AW} \

RAN

mcdonaldi

i ‘ | \ UCLA 199

(Na Nayarit

iM TA rc

Wor oll 1

DEINOCERITES Fig.58

LEN NY AS . \

AAS

INS

\ \

SW \

[ * 12

mcdonaldi

UCLA 199 ;

Nayarit

Mexico

Adames: Genus Deinocerites 153

INDEX TO SCIENTIFIC NAMES

albimanus (Anopheles), 9, 48

atlanticus (Deinocerites), 18k, 20k, 23k, 26,

27, 28, 28-31, 31, 32, 34, 52; figs. 3, 9-11

bahamensis (Culex), 9, 48

barretoi (Deinocerites), 18k, 19k, 22k, 23k,

36, 37, 39, 40-41; figs. 4, 21

belkini (Deinocerites), 16, 18k, 20k, 23k, 24k,

26, 27, 32, 33, 34-35, 81; figs. 3, 15-17

Brachiomyia, 11, 12

Brachiosoma, 11

cancer (Deinocerites), 3, 8, 9, 10, 11, 13, 16,

17k, 19k, 20k, 23k, 25k, 44, 44-49, 50,

54.52; 54, 55561: fies t, 9,-25-28

cancer (Deinocerites) of authors, 50, 51, 52,

50,55. 59, Of; Tape

Cancer Group, 4, 5, 6, 8, 17k-20k, 23k, 25k,

37, 43-44, 55, 58, 67, 71; fig. 5

carcinophilus (Culex), 9, 48

Cardisoma, 72

chalcospilans (Haemagogus), 9, 68

colombianus (Deinocerites), 17k, 19k, 21k,

22k, 24k, 52, 57, 58, 61, 63, 64-65; figs.

6, 41-43

conservator (Culex), 9, 68

Corethrella (Corethrella) sp., 9, 68

costaricensis (Deinocerites), 9, 17k, 19k, 21k,

22k, 24k, 58, 59, 65-68, 68, 69, 75; figs.

6, 44-46

crassum (Cardisoma), 8, 43, 58, 68, 75, 81

Culex, 6, 7, 16, 26

Culex (Culex), 3, 7

Culex (Culex) sp., 55

Culex (Melanoconion), 7, 27, 37

Culex (Melanoconion) spp., 9, 55, 61, 68, 75

curiche (Deinocerites), 17k, 19k, 21k, 22k,

24k, 58, 63, 66, 67, 68-70, 75; figs. 6, 47-

Deinocerites, 2, 4, 5, 6, 7, 8, 9, 10, 11-16,

26, 39, 51, 58, 76, 79; fig. 2

Deinocerites sp. of authors, 76

Deinokerides, 11

Dinanamensus, 11

Dinocerites, 11

Dinomimetes, 11

dyari (Deinocerites), 8, 18k, 19k, 21k, 22k,

24k, 37, 37-40, 40, 41, 42, 75; figs. 4, 18-

dyari (Deinocerites) of authors, 40

Dyari Group, 5, 6, 8, 18k, 19k, 21k, 22k,

24k, 35-37; fig. 4

eiseni (Anopheles), 9, 63, 75

elevator complex (Culex), 9, 48

Epibolocera, 8, 47

epitedeus (Deinocerites), 11, 17k, 19k, 22k,

24k, 48, 58, 59-61, 62, 63, 64, 65, 66, 67,

69; figs. 6, 35-37

epitedeus (Deinocerites) of authors, 62, 63, 72

Epitedeus Group, 4, 5, 6, 8, 15, 17k, 19k,

21k, 22k, 24k, 37, 56-59, 71; fig. 6

fascipes (Orthopdomyia), 9, 68

Galindomyia, 3, 6, 7, 26

Gecarcinidae, 8

Gecarcinus, 72

grabhamii (Anopheles), 9, 48

guanhumi (Cardisoma), 8, 44, 47, 51, 75

habilitator (Culex), 9, 55

howardi (Deinocerites), 7, 9, 17k, 19k, 22k,

23k, 25k,-71,.75).77-79, Shi fies. 7, J309

inflictus (Culex), 55, 69

inflictus complex (Culex), 9, 31, 48, 52, 61,

63, 68, 75, 81

jamaicensis (Psorophora), 9, 48

janitor (Culex), 9, 48

latisquama (Culex), 7, 9, 48, 61, 63, 65, 75

leei (Galindomyia), 6 |

magnus (Deinocerites), 11, 17k, 19k, 20k,

23k, 25k, 44, 51, 52-56; figs. 5, 32-34

magnus (Deinocerites) of authors, 45

mathesoni (Deinocerites), 8, 9, 10, 15, 16,

18k, 20k, 23k; 24k, 26, 27, 31-34, 34, 35,

75; figs. 3, 12-14

154 Contrib. Amer. Ent. Inst., vol. 7, no. 2, 1971

mcdonaldi (Deinocerites), 7, 9, 17k, 19k, 22k,

Zk, 25%, 35,07, 70, Th, Fi; 879, Fe

82; figs. 7, 56-58

melanophylum (Deinocerites), 17k, 19k, 20k,

23k, 25k, 31, 44, 46, 47, 49-52, 54, 55,

63, 65; figs. 5, 29-31

melanophylum (Deinocerites) of authors, 45

monospathus (Deinocerites), 49

nicoyae (Deinocerites), 8, 14, 18k, 19k; 21k,

22k, 24k, 37, 41-43; figs. 4, 22-24

nigripalpus (Culex), 9, 48

occidentalis (Ucides), 8, 37, 39, 43, 75

Ocypodidae, 8

opisthopus (Culex), 9, 48

panamensis (Deinocerites), 17k, 19k, 21k, 22k,

24k, 52, 58, 61, 61-63; figs. 6, 38-40

pseudes (Deinocerites), 2, 6, 7, 8, 9, 10, 11,

17k, 19k, 21k, 23k-25k, 33, 39, 43, 68,

69, 71, 72-77, 77, 718, 79, 80, 81; fies... 7,

50-52

pseudes (Deinocerites) of authors, 50

Pseudes Group, 4, 5, 6, 7, 8, 16k, 17k, 19k,

21k, 23k, 24k, 37, 44, 58, 70-72; fig. 7

Pseudothelphusidae, 8, 47

pugilator (Uca), 8, 33

scapularis (Aedes), 9

scapularis complex (Aedes), 55

spanius (Deinocerites), 11, 18k, 20k, 22k, 23k,

26, 27, 27-28, 29, 30; figs. 3, 8

spanius (Deinocerites) of authors, 16, 28, 31,

Spanius Group, 4, 5, 6, 8, 18k, 20k, 23k, 25-

27, 33, 35, 37, 58: fig. 3

sp. A (Deinocerites) of authors, 65, 67

subcylindrica (Uca), 8, 33, 75

taeniorhynchus (Aedes), 9, 48, 55, 75, 79, 81

tetraspathus (Deinocerites), 45

triseriatus (Aedes), 9, 48

troglodytus (Deinocerites), 52, 53, 55

Uca, 6:27, 72

Ucides, 72

A apes Reus nya: tan

Feu,

is bal

ay: Ys

=i}, aA SHIM

ait "bade 5 i

: mt

a) “th ”

Bi de ey!

Cae Nt

WON

aye

cali ge

pe Rea

No.

(Continued from inside front cover)

Parts of Volume 4, with prices

Mosquito studies (Diptera, Culicidae), 1968.

X. Peters, T. Michael. Dixinae originally described from North America. 7 pages.

XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes |

originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay.

29 pages. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XII. Berlin, O. G. W. A revision of the Neotropical subgenus Howardina of Aedes.

Carlson, Robert W. and Fred B. Knight, 1969.

Biology, taxonomy, and evolution of four sympatric Agrilus beetles (Coleoptera:

Buprestidae). 104 pages, 47 figures. Price: $3.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XIII. Barr, Ralph A. and Sylvia Barr. Pupae of the genus Psorophora in America

north of Mexico. 19 pages, 11 figures.

XIV. Berlin, O. G. W. Redescription of Toxorhynchites (T.) gigantulus from the

Philippines. 7 pages, 2 figures.

XV. Zavortink, Thomas J. A new species of treehole breeding Anopheles from

the southwestern United States. 12 pages, 4 figures.

Price: $1.25, postpaid.

Matthews, Robert W., 1970. A revision of the genus Spathius in America north of Mexico

(Hymenoptera, Braconidae). 86 pages, 31 figures. Price: $3.50, postpaid.

postpaid.

Parts of Volume 5, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XVI. Zavortink, Thomas J. A new species of treehole breeding Aedes (Ochlerotatus)

from southern California. 6 pages, 2 figures.

XVII. Adames, Abdiel J. and Charles L. Hogue. Two new species of Deinocerites from

Costa Rica. 6 pages, 6 figures.

XVI. Berlin, O. G. W. The subgenus Micraedes of Culex. 30 pages, 12 figures, 1 map.

Price: $2.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XIX. Zavortink, Thomas J. The treehole Anopheles of the New World. 35 pages, 12

figures. Price: $1.25, postpaid. .

Mosquito studies (Diptera, Culicidae), 1970.

XX. Schick, Robert X. The Terrens Group of Aedes (Finlaya). 158 pages, 60 figures.

Price: $5.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

V. Reinert, John F. Genus Aedes, subgenus Diceromyia Theobald in southeast Asia.

43 pages, 15 figures. Price: $1.50, postpaid.

Howden, Anne T. The Tanymecini of the West Indies (Coleoptera: Curculionidae). 74 pages,

76 figures. Price: $2.75, postpaid.

Kethley, John B., 1970. A revision of the family Syringophilidae (Prostigmata: Acarina).

76 pages, 38 figures. Price: $2.75, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

VI. Mattingly, P. F. The genus Heinzmannia Ludlow in southeast Asia. 104 pages,

5i figures. Price: $4.00, postpaid.

Parts of Volume 6, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXI. Belkin, John N., Sandra J. Heinemann, and William A. Page. The Culicidae of

Jamaica. 458 pages, 126 figures. Price: $15.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

Vil. Tyson, W. H. Genus Aedeomyia Theobald in southeast Asia. 27 pages, § figures.

vil. Tyson, W. H. Genus Aedes, subgenus Mucidus Theobald in southeast Asia. 54 pages

21 figures. Price: $3.50, postpaid.

Parts of Volume 7, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXII. Zavortink, Thomas J. A new subgenus and species of Aedes from Arizona.

11 pages, 2 figures.

XXII. Schick, Robert X. Additions and corrections to the revision of the Aedes

terrens group. 27 pages, 8 figures. Price: $1.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1971.

XXIV. Adames, Abdiel J. A revision of the crabhole mosquitoes of the genus

Deinocerites. 154 pages, 58 figures. Price: $6.00, postpaid.

Contributions

of the

American Entomological Institute

Volume 7, Number 3, 1971

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA

IX. The genus Orthopodomyia Theobald in Southeast Asia.

By Thomas J. Zavortink

X. The genus Culisefa Felt in Southeast Asia.

By N. V. Dobrotworsky

XI. A proposed reclassification of Neoculex Dyar based

principally on the male terminalia.

By Sunthorn Sirivanakarn

AN THSON

a a) ie

CONTRIBUTIONS

of the

AMERICAN ENTOMOLOGICAL INSTITUTE

The Contributions are for larger papers on insects. Each paper is a separate number, with separate

pagination and index. Separate numbers aggregating about 500 pages constitute a volume. Issues appear

irregularly, as suitable manuscripts are available.

Copies are sold separately or in subscriptions to complete volumes. Complete volumes are $12.00.

The price of separate numbers varies. Subscribers are billed for each volume with its beginning number,

and receive the parts as issued. Orders for separate numbers that total less than $8.00 must be accom-

panied by payment. Address orders or correspondence to the American Entomological Institute, 5950

Warren Road, Ann Arbor, Michigan 48105, U.S.A.

No.

Parts of Volume 1, with prices

Dasch, Clement E., 1964. The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae).

75 pages, 69 figures. Price: $2.25, postpaid. .

Mosquito studies (Diptera, Culicidae), 1965.

I. Belkin, John N., etal. A project for a systematic study of the mosquitoes of Middle Americ:

II. Belkin, John N., et al. Methods for the collection, rearing and preservation of mosquitoes.

78 pages, 4 figures. Price: $2.25, postpaid.

Same as no. 2, but in Spanish. Price: $2.25, postpaid.

Matthews, Robert W., 1965. The biology of Heriades carinata Cresson. (Hymenoptera,

Megachilidae). 33 pages, 23 figures. Price: $1.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1965.

Ill. Ramalingam, Shivaji, and John N. Belkin. Two new Aedes from Tonga and Samoa.

IV. Belkin, John N. The mosquitoes of the Robinson-Peabody Museum of Salem expedition to

the southwest Pacific, 1956.

34 pages, 3 figures. Price: $1.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1965.

V. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from Middle America. 95 pages. Price: $2.75 postpaid.

Mosquito studies (Diptera, Culicidae), 1966.

VI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from North America. 39 pages. Price: $1.00, postpaid.

DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae (Homopotera,

Cicadellidae). A synopsis of the genus Ponana.

86 pages, 257 figures. Price: $2.50, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1967.

I. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald in 1 Thailand.

55 pages, 20 figures. Price: $1.75, postpaid.

Parts of Volume 2, with prices

Contributions to the mosquito fauna of southeast Asia, 1967.

Il. Bram, Ralph A. The genus Culex in Thailand (Diptera: Culicidae).

296 pages, 104 figures. Price: $9.00, postpaid.

Schuh, Toby, 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region.

35 pages, 30 figures. Price: $1.25, postpaid.

Snider, Richard J., 1967. The chaetotaxy of North American Lepidocyrtus s. str. (Collembola:

Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1968.

Ill. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald of southeast

Asia. 76 pages, 26 figures. Price: $2.25, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1968.

IV. Knight, Kenneth L. Species of the subgroup Chrysolineatus of group D, genus Aedes,

subgenus Finlaya Theobald. 45 pages, 12 figures. Price: $1.50, postpaid.

Parts of Volume 3, with prices

Mosquito studies (Diptera, Culicidae), 1968.

VII. Belkin, John N. The Culicidae of New Zealand.

182 pages, 30 figures. Price: $5.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1968.

VIN. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia.

221 pages, 41 figures. Price: $6.75, postpaid.

Fleming, Richard C., 1968. Head musculature of sphinx moths (Lepidoptera: Sphingidae).

32 pages, 37 figures. Price: $1.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1968.

IX. Belkin, John N. The type specimens of New World mosquitoes in European museums.

69 pages. Price: $2.00, postpaid.

(Continued on back cover)

Contributions

of the

American Entomological Institute

Volume 7, Number 3, 1971

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA

IX. The genus Orthopodomyia Theobald in Southeast Asia.

By Thomas J. Zavortink

X. The genus Culiseta Felt in Southeast Asia.

By N. V. Dobrotworsky

XI. A proposed reclassification of Neoculex Dyar based

principally on the male terminalia.

By Sunthorn Sirivanakarn

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. IX.

CONTENTS

DOC ON ire a we ik ea Se ee Se eS ew ae

Genus Orthopodomyia Theobald ...........6080e+c48-8

KEYS TO THE SOUTHEAST ASIAN SPECIES

POUL TS cai aes s ee ae ee uta gyre gang a ae

NEARS PRIN LAs te oS A ee eA ee

Oe cae ie ee ee ee ee ee Ee eee

a POUR TH OI AGE DGARVAG I. 6 os ee ik Nee Se

DESCRIPTIONS OF THE SUBGROUPS AND SPECIES

PLA VIC OSTASUDGROUP 66 ok Se re ee ee

ORV CHECS; AISA SS ee es Hk ee ee ES

PEN VHCHE ES DION ae aha ype eg wee. bob ese a

SIO ESTs DAVOLEIN a a Shae A ae ee

ES ie ae iy ake ee la be eo a eee

COLE SOME ACO OAs Bs eC ae oe Se tek WE LE os ROW, We eS

ST ee Oe oe ae de ke Gk ew te ee) Wace wets

GIVES MiCICOGIEL oS Suan oA Ra Se SE eee

ANOPHE LOIUES SUBGROUP oo nos i we bo ee Bees

CUA HOHEISTS ‘DATT AUG 205045 oe es See ee eS

Coppin (Glee oa age a be ae ee a

HYBRIDS

CID IPE SS CUERPO OIUES 48a Ais eee ie

NOMEN DUBIUM | :

mcegregovi (Banks)...... SSC e ee oe eae ea

PC CN ey a a ae eS ee ow We eck ote

ay Ee ea ee oo Re ae eS Oe

co CO) Od Of Ol > Go |

THE GENUS ORTHOPODOMYIA THEOBALD IN SOUTHEAST asta?

Thomas J. Zavortink 2

INTRODUCTION

This paper treats the 7 species of Orthopodomyia which occur in the

Southeast Asian area. Although it has been largely extracted from a more

comprehensive study of the entire genus (Zavortink 1968), several significant

changes and additions have been made. These include the following: keys to

species have been rewritten and made more artificial; descriptions of the spe-

cies have been shortened and restricted to characters which are diagnostic or

at least characteristic of one or more species; discussions and group descrip-

tions have been modified to pertain only to species found within the Southeast

Asian region; descriptions have been corrected, where necessary, to accomo-

date variations observed during examination of additional specimens; sections

on distribution and biology now include more extensive data for the Thailand

collections; and a recently described species has been included.

The terminology of the present paper generally conforms to that of

Belkin es and the format to that of Bram (1967), Delfinado (1967, 1968), and

Knight (1968). An asterisk following the abbreviations used (“ = male, ? = fe-

male, P = pupa, and L = larva) indicates that all or some portion of that sex or

stage is illustrated. Abbreviations used for the references to the literature

conform to the World List of Scientific Periodicals, 3rd ed., Academic Press,

1952. Types of the included species which are in the British Museum (Natural

History) and the United States National Museum have been studied by me.

In addition to material accumulated by the Southeast Asia Mosquito

Project, specimens from the following institutions were examined: Bernice P.

Bishop Museum; British Museum (Natural History); Liverpool School of Trop-

ical Medicine; Taiwan Provincial Malaria Research Institute; United States

National Museum; University of California at Davis; and, University of Cali-

fornia at Los Angeles. 3

Orthopodomyia is a small genus of container breeding mosquitoes with

a nearly world wide distribution. Eight species groups have been recognized

(Zavortink 1968), but only one of these, the albipes group, occurs in the

Oriental region. The albipes group is, however, the largest and most diverse

in the genus. It is the only group in which sympatry of species is common and

it is the group in which the greatest discordance between classifications based

on adults and immature stages is found.

Knight and Mattingly (1950) were the first to study this group of mos-

quitoes. Their anopheloides subgroup is nearly coextensive with that portion

of the albipes group considered in the present paper.

Contribution from the Southeast Asia Mosquito Project supported by U.S.

Army Medical Research and Development Command Research Contract

DA- 49-193-MD- 2672.

‘i ho acai of Zoology, University of California, Los Angeles, California

90024. :

2 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

GENUS ORTHOPODOMYIA THEOBALD

Orthopodomyia Theobald 1904, Entomologist 37: 236-237. Type species: O.

albipes Leicester 1904.

FEMALE. Head. , Frons narrow, with light decumbent scales; vertex

and occiput with numerous narrow decumbent and broad erect scales; decum-

bent scales light, erect scales dark posteriorly, light or light-tipped anteri-

orly; dorsolateral, lateral and ventral surfaces with broad flat scales; clypeus

small, normally bare; labium slightly swollen apically, dark scaled with light

scales usually in a narrow to broad band near or distad of middle, in a dorsal

preapical spot, and streaked on ventral surface from or basad of band to or

near apex; palpus long, 0.4 to 0.6 length of proboscis, 4 or 5 segmented, dark

scaled with 2 or 3 light patches beyond the base; antenna longer than probos-

cis, pedicel and flagellomere 1 with light scales. Thorax. Paratergite moder-

ately broad, bare; mesonotal bristles unusually numerous, strongly developed,

and long; scutum largely covered with light and dark scales of various sizes

in a complicated and variable pattern; scutellar lobes with long narrow light

scales; apn, ppn, ppl, lower sip, and upper mep bristles present, pra bris-

tles present or absent, sp and psp bristles absent; apn, ppn, pst, ppl, ssp,

pra, stp, and upper and lower mep scales present, psp and pcx scales absent.

Legs. Coxae with light and sometimes dark scales; femora, tibiae, and, toa

lesser extent, tarsomere 1 with light and dark scales in a complicated pattern

of streaks, spots, bands and speckles; fore and mid legs with tarsomere 1

longer than the remaining 4 tarsomeres combined and tarsomere 4 shorter

than 5; hind tarsomere 5 all white; claws simple. Wing. Dark scaled with

conspicuous light patches on costa and other veins, the number and size of

light patches, especially at base of costa, exceedingly variable. Halter.

Scales light or dark. Abdomen. Terga dark with light scales in distinct or

connected basolateral and apicolateral patches which become smaller on dis-

tal segments and in small to moderately large median subdorsal patches on

all or more distal segments. Tervminalia. Segment VIII not retractile; cerci

large, compressed.

MALE. Similar to female. Head. Proboscis dark with light scales

in complete or incomplete band basad of middle; palpus straight, 0.7 to 0.9

length of proboscis, 5 segmented, dark scaled with 4 light patches; antenna

subequal to proboscis, strongly plumose, flagellomeres 1 to 4 or 5 with tufts

of light scales. Legs. Anterior fore and mid claws large and unidentate,

posterior fore and mid claws and hind claws simple. Abdomen. Usually more

extensively light scaled than in female. Terminalia. Tergum VII with dis-

tinct lobe on posterior margin; tergum IX not strongly sclerotized, without

lobes, usually with fine setae; basimere well developed, long conical, without

apical or basal lobes; basal mesal lobe with 6-8 terete apical bristles, the

more ventral usually finer and more strongly curved; distimere simple, with

numerous setae and usually 1 simple terminal appendage; phallosome simple;

aedeagus large, varied in shape, sclerotization, and dentition; paraproct with

2-5 apical teeth; cercal setae fine, 1-6.

PUPA. Cephalothorax. Hairs 6, 7-C close together; 8,9-C relatively

close together, far caudad of trumpet, and in line perpendicular to middorsal

ridge. Trumpet. Widely spaced; moderate in length; tracheoid portion ab-

sent; meatus without slit. Abdomen. Hair 1-I strongly developed and strongly

dendritic; hair 1-II generally well developed and with many branches; hair 9-

II- VIII cephalad of caudolateral angle of segment, on II-VI small and single,

on VII moderately to strongly developed and 7-13b, on VIII strongly developed

and usually 11-16b (9-19). Terminal Segments. Hairs 1-IX, X absent.

Paddle. Wonger than broad; without spicules; hair 1-P present, small, usually

branched; 2-P absent.

Zavortink: Orthopodomyia in Southeast Asia 3

LARVA. Head. Slightly to distinctly broader than long; mental plate

well developed, usually with 8-13 teeth on each side; integument without orna-

mentation. Antenna. Short to moderately long, smooth; hair 1-A in basal

0.3-0.5, moderately developed, and usually 4-9b. Thovax. Deep red or pur-

ple epidermal pigment usually present; 1 pair of conspicuous tracheal dilations

present. Abdomen. Mature fourth instar larva with large sclerotized plates

on segments VII and VII; hair 6-I, II strongly developed and multiple, 6-III,V,

VI very long and single, 6-IV shorter and usually double; comb scales in 2

rows, those of posterior row larger; siphon moderately long to quite long,

pecten absent; only hair 1-S well developed, placed in basal 0.3-0.5; anal

saddle complete, margin without spicules; 1-X small to moderately developed,

2-X very strongly developed and multiple, 3-X long and single; ventral brush

well developed, usually with 7 pairs of hairs. |

DISTRIBUTION. The Oriental albipes group of the genus Orthopodo-

myia is found from India, southern China, and southern Japan, south to Ceylon,

the Malay Archipelago, and northeastern Australia.

TAXONOMIC DISCUSSION. The albipes group of Orthopodomyia con-

tains 10 species. Seven of these, albipes Leicester, andamanensis Barraud,

anopheloides (Giles), lanyuensis Lien, madrensis Baisas, siamensis Zavortink

and wilsoni Macdonald, are found within the Southeast Asian area. The 3 exo-

tic species are flavicosta Barraud and flavithorax Barraud from coastal south-

western India and Ceylon and papuensis Zavortink from Ceram and New Guinea.

The 10 species of the albipes group fall into 4 subgroups. All 4 of

these, the albipes, anopheloides, flavicosia, and wilsoni subgroups, occur in

the Southeast Asian region and the monotypic albipes and wilsoni subgroups

are restricted to it.

Adults of all species treated in this paper are very similar and those

of each species are quite variable, particularly in ornamentation of the pro-

boscis, wing, abdomen, and hind leg. Consequently, specific identification is

frequently difficult or impossible. The male terminalia of most species are

distinct, but the differences between them are not of sufficient magnitude to

make identification easy. The immature stages show the greatest divergence

and the species and subgroups are defined largely on the basis of differences

in these stages. Unfortunately, the variation in the chaetotaxy of both the

pupa and larva for several of the species is so great that identification, even

to subgroup, is often not simple. Despite the difficulties encountered in iden-

tifying any single stage, specific determinations are quite reliable when made

from a combination of larval, pupal, and adult characters.

BIOLOGY. The immature stages of species in the albipes group occur

in water which accumulates in bamboo stumps, bamboo internodes, rot cavi-

ties in trees, and artificial containers. Adults are rarely encountered in the

field. Females of 2 species, albipes and andamanensis, are known to bite

man. |

KEYS TO THE SOUTHEAST ASIAN SPECIES

ADULTS

1. Pra usually with 1 bristle; ssp scale patch forked

or-ancled (Figs fa. 6 er Oe ee ae ee ee 2

Pra without bristles; ssp scale patch simple

CPV OB Ati, aia al olor w elit ae oat se xe i eee 3

2(1). Joint between hind tarsomeres 1 and 2 without

Bi CaCl WATE Te NY, a ee madrensis (p. 6)

Joint between hind tarsomeres 1 and 2 witha

PEMAG WHATS) PIN Big ee Sih aig unk, a aE Bons lanyuensis (p. 8)

3(1).

4(3).

5(4).

2(1).

3(2).

4(3).

5(1).

Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

Hind tarsomere 2 with a conspicuous subequal

white ring at each end (Fig. 1g); tergum II or

terga II and III of female usually with a large

middorsal basal light patch (Fig. lc). ....

Hind tarsomere 2 usually entirely or largely

dark at either base or apex (Figs. 1d,e, f);

terga II and III of female without a large mid-

dorsal basal light patch (Fig. 2e).......

Hind tarsomere 2 usually with more extensive

white scaling at base than apex (Fig. 1d);

palpus of female with 3 light patches beyond

the base (Fig. a). oc. sea bn ale ek ew

Hind tarsomere 2 with more extensive white

scaling at apex than base (Figs. le, f); palpus

of female with 2 light patches beyond the base

Hind tarsomere 4 with a dark spot or ring distad

GE middle (HIG. BO) ce cee nce a cues: anopheloides (in part)(p.

Hind tarsomere 4 entirely white scaled (Fig. 1f)

. . stlamensis

andamanensis a

anopheloides (in part)

MALE TERMINALIA

Aedeagus with a pair of sclerotized teeth on

ventral surface and without a conspicuous

projection between or below ventral parameres ........

Aedeagus without sclerotized teeth on ventral

surface and with a conspicuous projection

between or below ventral parameres .....

Basal mesal lobe with 7 to 9 more or less

equally developed stout apical bristles

CPs SENS Ate restk Bid ce ay uerdeine duck Edt ays

Basal mesal lobe with 4 or 5 stout and 1 or 2

finer apical bristles (as in Fig. 5a). .....

Aedeagus not pyriform, broadest in apical

Bea C2 1c i ie aces acels eabidcas sauroecebenehisay ids Geecd

Aedeagus more or less pyriform, broadest

Mi bases Wal (as.in Fic. OD). 6c ek Sew ee:

Mesal surface of basimere distad of basal

mesal lobe with strong, elongate, special-

izec bristles (Fig. 4a). 20 fees ee

These bristles weaker and shorter (Fig. 6a) . .

Basal projection of aedeagus usually extending

conspicuously between ventral parameres

more ventrally and not extending conspicu-

ously between ventral parameres (Fig. 8a).

. madrensis tg

lanyuensis

O65 > \OF- ati Oe ee OY FOr OS Of

s+. 2. @loipes

. « « wilsoni

anopheloides (p.

. andamanensis (p.

tt)

4(3).

5 (4).

2(1).

3(1).

Zavortink: Orthopodomyia in Southeast Asia 5

PUPAE

Trumpet nearly cylindrical no eS HE

Trumpet not. cylindrical; 460%..care So eee PP

Hair 5-C strongly developed, usually 5, 6b(3- 8);

hair 2-II-IV thickened (Fig. 9)

Hair 5-C moderately developed, usually single

or double (1-3b); hair 2-II-IV fine (Fig. 9) .

Trumpet broadening to nearly maximum width

in basal portion, remainder almost parallel

sided (Fig. 10)

Trumpet broadening gradually from base to apex .

Hair 5-VI usually single or double and long,

extending nearly to or beyond alveolus of

hein 40eU ie (is. ay ae ss PAR hos

Hair 5-VI usually 3-7b(2-8) and short, extending

nearly to or slightly beyond alveolus of hair

A~VIl, longer in hairy forms::.... 0.9.) ¢ea04%

Hair 1-III usually 12-14b(10-17); hair 9-C usu-

gily 2:SH(A-a) (Pleo Oh iw wales ecg ivet rae,

Hair 1-III usually 5-7b(3-11); hair 9-C usually

single (single, double) (Fig. 11)

FOURTH STAGE LARVAE

Branches of hair 6-I, II usually at least as stout

as 6-III; hair 1-M, T single and short, sub-

equal in length to 3-M or 2-T, and posterior

comb scales expanded at apex and with 5-7

elongate spines .

Branches of hair 6-I,II finer than 6-III; hair

1-M, T branched and/or long, much exceeding

3-M or 2-T, and/or posterior comb scales

narrow at apex and with a single elongate spine .

Branches of 6-I, II tapering to a fine apex and

barbed from near base to near apex; segment

VI of mature larva with dorsal sclerotized

DAAO i a eee a ee ee ee

Branches of 6-I,II coarse to near apex and

barbed only in basal portion (Fig. 3); seg-

ment VI of mature larva without a sclero-

CLEGG TES Sig. a at Ge, he By tat eee tg bos

Posterior comb scales narrow at apex and

with only 1 spine of fringe elongated (Figs.

4, 6); sclerotized body parts exclusive of

siphon largely straw-colored to tan

Posterior comb scales expanded at apex and

with fan-like fringe of several elongate

spines (Figs. 7, 8); sclerotized body parts

exclusive of siphon largely brown to dark brown

madrensis

Co DO

siamensis (p. 9)

wilsoni (p. 11)

. albipes (p. 12)

car 4

madrensis (p. .

lanyuensis (p. 8

andamanensis (p. 14)

anopheloides (p. 16)

siamensis (p. 9)

lanyuensis

6 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

4(3). Elongate spine of posterior comb scales

usually very long; siphon long and thin, |

index usually 7.0-12:0 (Pig. 6)... 0 Ses es ee es Wilsoni (p. 11)

Elongate spine of posterior comb scales

shorter; siphon broader, index usually

406.0 (3.6- 729) (ies 4) A bake cise albipes (p. 12)

5(3). Hair 6-I usually 9, 10b(7-10); hair 6-II |

iegually 9- LibCi- 82) 6S ers SU Sea Gs andamanensis (p. 14)

Hair 6-I usually 5-8b; hair 6-II usually :

aOR (P18. 1) Ses ee Fe ee anopheloides (p. 16)

DESCRIPTIONS OF THE SUBGROUPS AND SPECIES

FLA VICOSTA SUBGROUP

PUPA. Trumpet. Varied. Abdomen. Hair 1-II strongly developed,

long and usually with 9-20 mostly simple branches, Some much finer than

others, arising from the base; 5-V, VI usually long, extending to or beyond

ste ae of hair 4 of second following segment; 5-VI usually single (single,

ouble).

LARVA. Thorax. Hair 1-M, T single, short, usually subequal in

length to 3-M and 2-T. Abdomen. Hair 6-I,II with branches as stout as or

stouter than 6-III, form varied; posterior comb scales expanded at apex and

usually with 5-7 large spines.

DISTRIBUTION. Species in this subgroup are found in southwestern

India, Ceylon, peninsular Thailand, Luzon, and Lanyu.

TAXONOMIC DISCUSSION. As currently interpreted, the flavicosta

subgroup contains the 5 species flavicosta, flavithorax, lanyuensis, madrensis,

and siamensis. Only the last 3 of these occur in the Southeast Asian region.

O. lanyuensis and madrensis are very similar to each other in all

stages. O. siamensis is quite distinct from them, but is largely indistinguish-

able in the adult stage from andamanensis and anopheloides, 2 Southeast Asian

species in the anopheloides subgroup.

The flavicosta subgroup is probably the oldest in the albipes group.

The species, except for madrensis and lanyuensis, are relatively well marked

and all have very limited, relictual distributions in areas characterized by ex-

tremely heavy precipitation. Additional species undoubtedly exist on biologi-

cal islands in and around the periphery of the Southeast Asian area.

BIOLOGY. Larvae and pupae are usually found in tree-holes. Habits

of the adults are unknown.

ORTHOPODOMYIA MADRENSIS BAISAS

(Figures la, 3, 11)

Orthopodomyia madrensis Baisas 1946, Mon. Bull. Bur. Hlth. Philipp.,

Manila 22:41 (c*, 9, P*, L*); Knight & Mattingly 1950, Proc. ent. Soc.

Wash. 52:17 (o, 2, L); Delfinado 1966, Mem. Amer. ent. Inst. 7:68

re L); Zavortink 1968, Contr. Amer. ent. Inst. 3(2):122 (o*, 9,

p*, L*).

FEMALE. (Figure la) Head. Dorsolateral scales yellow; labial light

scales white to yellow, band near middle, preapical spot sometimes com-

pleted ventrally, and sometimes with additional light scales in dorsal streak

basad of middle band; palpus 0.42 length of proboscis, light scales white to

Zavortink: Orthopodomyia in Southeast Asia 7

yellow, in 2 patches beyond the base. Thorax. Pra usually with 1 bristle;

pleural scales largely pure white; ssp scale patch well developed, a moder-

ately long forked or angled row (upper extension erroneously called hypostigial

scales by Zavortink (1968)). Legs. Fore and mid tarsomeres 2 and 3 usually

light scaled at both ends; hind tarsomere 1 dark scaled at apex; hind tarsomere

2 with at most a few light scales at base, broadly white ringed at apex; hind

tarsomere 3 entirely white or with dark patch or ring; hind tarsomere 4 all

white. Wing. Vein Sc without presectoral light patch; 1A dark at base. Abdo-

men. Light scales white or rarely yellowish; terga sometimes with additional

light scales in weak narrow basal band or middorsal basal patch; sternum II

or sterna II and III light scaled, II-V or IV, V light scaled with median lateral

dark patch, VI-VIII dark scaled with light basolateral patch and sometimes

light median streak.

MALE. Head. Proboscis with additional light scales in broad preapi-

cal band, usually in ventral streak joining 2 bands and sometimes scattered

dorsally; palpus 0.77 length of proboscis. Abdomen. Generally as in female.

Terminalia. (Fig. 3) Tergum VIII lobe with apex emarginate and serrate;

basimere with strong sinuous bristles distad of basal mesal lobe; basal mesal

lobe with 7 or 8 more or less equally developed stout bristles; aedeagus weak-

ly sclerotized, more or less irregularly oval in dorsal aspect, with a pair of

nearly contiguous to widely separated ventral teeth about 0.55-0.67 distance

from base and without a conspicuous projection between or below ventral para-

meres.

PUPA. (Fig. 11) Cephalothorax. Hair 1-C moderately developed,

single or double; 2-C moderately developed, usually 2, 3b(2-4); 3-C strongly

to very strongly developed, usually 4-6b(3-8); 4,5-C very strongly developed,

usually 3, 4b(2-4); 7-C very strongly developed, usually double (1-3b); 8-C

moderately to strongly developed, single; 9-C moderately to strongly devel-

oped, usually double or single (1-3b). Trumpet. Largely dark brown, gradu-

ally widening from base. Abdomen. Hair 2-II slightly thickened, mesad of

O-II; 2-III, IV fine; 1-III usually 4-6b; 5-IV usually 3b(2, 3); 5-V usually double

(single, double); 5-VI usually single.

LARVA. (Fig. 3) Head. Integument largely tan to brown; hair 13-C

moderately developed, usually 3-5b. Antenna. Short. Abdomen. Hair 6-1, Il

4,5b(3-5), with branches coarse to near apex and barbed only in basal portion;

1-III moderately long, single; 1-V long, single; 1-VI shorter than 4-VI; seg-

ment VI without sclerotized plate; siphon index usually 3. 8-6. 6; hair 1-S lo-

cated 0: 44-0.53 from base of siphon and usually 8-11b(6-13).

TYPE DATA. Orthopodomyia madrensis Baisas, holotype male with

associated larval and pupal skins (TH1B-4) lost; type locality: Llavac, Laguna,

Luzon (Sunico), PHILIPPINES.

DISTRIBUTION. Specimens examined: PHILIPPINES, Luzon, 22

males, 15 females, 25 pupae, 43 larvae, 19 rearings. Records from litera-

ture: PHILIPPINES, Luzon (Baisas 1946, Delfinado 1966, Zavortink 1968).

TAXONOMIC DISCUSSION. O. madrensis and lanyuensis, a species

only recently described by Lien, are presently considered to be very closely

related. The two are distinguished from each other only in the adult stage and

are easily separated from all other species by having the subspiracular scale

patch forked or angled and a single bristle on the prealar knob. O. madrensis

is distinguished from lanyuensis by lacking a large white ring over the joint

between hind tarsomeres 1 and 2. The male terminalia of both species are un-

usual in having all the apical setae of the basal mesal lobe more or less equally

developed and strong. Pupae of madrensis and lanyuensis have no unique char-

acters; they are separated from the other species by the combination of a long,

Single or double hair 5-VI and the trumpet broadening gradually from the base.

The larvae of both species have abdominal hair 6-I, Il peculiarly developed;

each branch is stout nearly to its apex and barbed only near its base. This

character alone sets them apart from all other Orthopodomyia larvae.

Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

O. madrensis is not distinct from the other members of the albipes

group, including lanyuensis, in ornamentation of the proboscis, base of the

costa, or abdominal sterna, as thought by Baisas (1946: 41), Knight &

Mattingly (1950: 6, 19), and Lien (1968: 6). The light scaling of the proboscis

is frequently white rather than yellow and is subject to the same variation in

extent as in the other species. The basal portion of the costa varies from

completely dark scaled through distinct light basal, prehumeral and humeral

patches to completely light scaled. Only abdominal sterna II and III are light

scaled. As mentioned above, the number of white hind tarsomeres is vari-

able in this species.

O. madrensis is apparently endemic to the island of Luzon in the

Philippines.

BIOLOGY. This species breeds in rot holes in trees and treeferns; it

is frequently associated with anopheloides in those habitats. Nothing is known

of the habits of the adults.

ORTHOPODOMYIA LANYUENSIS LIEN

Orthopodomyia lanyuensis Lien 1968, Trop. Med. 10: 4 (o*, 9, P*, L*).

FEMALE. Head. Dorsolateral scales yellow; labial light scales

cream-colored, band near middle, preapical spot weakly complete ventrally;

palpus 0. 44 length of proboscis, light scales white to yellow, in 2 patches be-

yond the base. Thorax. Pra with 1 bristle; pleural scales largely pure white;

ssp scale patch well developed, a moderately long forked row. Legs. Fore

and mid tarsomeres 2 and 3 light at both ends; hind tarsomere 1 with broad

apical white ring, hind tarsomeres 2 and 3 conspicuously white ringed at both

ends, hind tarsomere 4 all white. Wing. Vein Sc without presectoral light

patch; 1A dark at base. Abdomen. Light scales white to yellowish; distal ter-

ga with additional light scales in basal bands; sternum II largely light scaled,

IlI- VU dark scaled with light basal band and basolateral patch which becomes

smaller on distal segments.

MALE. Head. Proboscis with additional light scales in large preapi-

cal patch; palpus 0.85 length of proboscis. Abdomen. Terga with conspicuous

basal bands. Terminalia. (Probably indistinguishable from madrensis. )

Tergum VIII lobe with apex rounded and serrate; basimere with strong sinuous

bristles distad of basal mesal lobe; basal mesal lobe with 8 or 9 more or less

equally developed stout bristles; aedeagus weakly sclerotized, more or less

irregularly oval in dorsal aspect, with a pair of slightly separated ventral

teeth about 0.58-0.60 distance from base, and without a conspicuous projec-

tion between or below ventral parameres. -

PUPA and LARVA. Apparently indistinguishable from madrensis.

TYPE DATA. Orthopodomyia lanyuensis Lien, holotype male with as-

sociated larval and pupal skins (78400. 6) in Taiwan Provincial Malaria Re-

search Institute; type locality: south of Hongtou, Lanyu (Chen), TAIWAN.

DISTRIBUTION. Specimens examined: TAIWAN, Lanyu, 1 male, 1

female, 2 pupae, 2 larvae, 2rearings. Records from the literature: TAIWAN,

Lanyu (Lien 1968). |

TAXONOMIC DISCUSSION. O. lanyuensis, only recently described by

Lien, is very similar to, and perhaps conspecific with, madrensis. The 2

are distinguishable only as adults, and then only by characters which are

known to be extremely variable in Orthopodomyia species of the albipes group.

The ways by which adults, male terminalia, pupae, and larvae of lanyuensis

and madrensis are separated from the other species in Southeast Asia are

found in the taxonomic discussion of madrensis. O. lanyuensis differs from

madrensis by having a large white ring over the joint between hind tarsomeres

1 and 2. Since lanyuensis differs from madrensis by having a different type of

Zavortink: Orthopodomyia in Southeast Asia 9

leg banding, rather than _ in the number of white hind tarsomeres, I am

considering it to be a distinct species.

The pupae of lanyuensis which I have seen have a slightly greater num-

ber of branches in several hairs, including hair 5-VI which is double, % than

any specimens of madrensis. I believe these differences, rather than being

specific, are the result of hairiness in these individual pupae. Hairy larval

forms are unknown.

O. lanyuensis is presently known only from Lanyu, a small island east

of the southern end of Taiwan. O. madrensis occurs about 250 miles further

south on Luzon, |

BIOLOGY. The immature stages are found in tree-holes. Habits of

the adults are unknown.

ORTHOPODOMYIA SIA MENSIS ZAVORTINK

(Figures 5, 9)

Orthopodomyia siamensis Zavortink 1968, Contr. Amer. ent. Inst. 3(2): 126

(i %, fy PAL):

FEMALE. Head. Dorsolateral scales brown anteriorly, white or yel-

low posteriorly; labial light scales cream-colored, band distad of middle; pal-

pus 0.42 length of proboscis, light scales white, in 2 patches beyond the base.

Thorax. Pra bristles absent; pleural scales largely white to cream-colored

or yellow-tinged; ssp scale patch well developed, a simple short to moderately

long row. Legs. Fore and mid tarsomeres 2 and 3 usually light scaled at

both ends; hind tarsomere 1 white scaled at apex, hind tarsomere 2 with at

most a very narrow white ring at base, broadly white ringed at apex, hind

tarsomere 3 entirely white or less commonly with dark patch or ring distad

of middle, hind tarsomere 4 entirely white. Wing. Vein Sc without presector-

al light patch; 1A usually dark at base. Abdomen. Light scales white or

white and yellow; additional light scales sometimes in basal bands on distal

terga; sterna II- VII dark scaled with basolateral light patch which becomes

smaller on distal segments.

MALE. Head. Proboscis with additional light scales in dorsal and

ventral preapical patches or a broad preapical band which extends further ba-

sally on ventral surface; palpus 0. 82 length of proboscis. Abdomen. As in

female.. Terminalia. (Fig. 5) Tergum VII lobe with apex rounded or emar-

ginate and with or without teeth; basimere without strong specialized bristles

distad of basal mesal lobe; basal mesal lobe with 4 or 5 stout bristles and 1 or

2 finer ones; aedeagus strongly sclerotized, shape in dorsal aspect variable,

, but never pyriform and always broadest in apical half, with a pair of nearly

contiguous ventral teeth about 0.50-0.55 distance from base, and without a

conspicuous projection between or below ventral parameres.

PUPA. (Fig. 9) Cephalothorax. Hairs 1, 2-C strongly developed, usu-

ally 6, 7b(5-8); 3-C strongly developed, usually 7-9b(4-10); 4,5-C strongly

developed, usually 5, Lh By 7-C strongly developed, double; 8-C strongly

developed, usually 6, 7b(3-8); 9-C strongly developed, 3-5b. Trumpet. Large-

ly light brown, nearly cylindrical. Abdomen. Hair 2-II-IV thickened, 2-II

laterad of 5-II; 1-III usually 5, 6b(5-7); 5-IV double; 5-V, VI single.

LARVA. (Fig. 5) Head. Integument largely tan to light brown; hair

13-C moderately strong, usually 8, 9b(8-11). Antenna. Moderately long.

3 In one paratype in SEAMP collection hair 5-VI is single on one side, double

on the other (B.de M.).

10 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

Abdomen. Hair 6-I,II 6, 7b with branches tapering to a fine apex and barbed

from near base to near apex; 1-III long, single; 1-V very long, double; 1-VI

shorter than 4-VI; segment VI with dorsal sclerotized plate; siphon index usu-

ally 6.4-7.8 (6.4-10.1); hair 1-S located 0. 41-0.49 from base of siphon and

usually 6-8b(5-8).

TYPE DATA. Oritkopodomyia siamensis Zavortink, holotype male with

associated larval and pupal skins (TG 102-30) in U.S. National Museum; type

locality: Muang, Trang. THAILAND.

DISTRIBUTION. Specimens examined: THAILAND, Nakhon Si :

Thammarat, Narathiwat, Phangnga,Ranong, Trang, 14 males, 8 females, 20

pupae, 58 larvae, 21 rearings. Records from the literature: THAILAND,

Narathiwat, Trang (Zavortink 1968).

TAXONOMIC DISCUSSION. The adult of this species differs from

madrensis and lanyuensis, the other Southeast Asian members of the flavi-

costa subgroup, by not having a forked or angled subspiracular scale patch or

a bristle on the prealar knob. It is, however, apparently indistinguishable

from andamanensis and many specimens of anopheloides, Southeast Asian spe-

cies of the anopheloides subgroup. The male terminalia are differentiated

from those of all other species by the shape of the aedeagus. The pupa has

hair 2-II-IV thickened and thorn-like. In the larva the branches of hair 6-I, II

are stout, yet taper to a fine apex and are barbed throughout. Abdominal seg-

ment VI of the mature larva bears a dorsal sclerotized plate; while this plate

is commonly found in Orthopodomyia larvae of other groups, siamensis is the

only species in the albipes group to have it developed. ’

Hind tarsomere 3 of the adult is entirely white scaled or has a dark

patch or ring beyond the middle. Ornamentation of the base of the costal vein

is almost as variable in this species as in madrensis. Hairy forms of the lar-

va and pupa are not yet known.

O. siamensis is a relictual species restricted, as far as is known, to

peninsular Thailand. Despite the extreme similarity with members of the

anopheloides subgroup in the adult stage, the species seems to be quite re-

moved from all others. |

BIOLOGY. The immature stages have been collected in rot holes in

trees, stumps, roots and in bamboo stumps. When found in the latter habitat,

they may be associated with albipes. All known adults are reared and nothing

is known about their behavior. 7

WILSONI SUBGROUP

PUPA. Trumpet, Largely dark brown, cylindrical. Abdomen. Hair

1-II moderately developed, moderately long, dendritic or with 1 or more

stalks from which 18-34 fine branches arise; 5-V, VI long, usually extending

G eee alveolus of hair 4 of second following segment; 5-VI usually double

=3D).

LARVA. Thorax. Hair 1-M, T 1-5b, moderately long to long, sub-

equal in length to or much longer than 3-M and 2-T. Abdomen. Hair 6-1, IT

with branches finer than 6-III, tapering to fine apex and barbed from near base

to near apex; posterior comb scales narrow at apex and with a single long,

strong spine.

TAXONOMIC DISCUSSION. The monotypic wilsoni subgroup, known

only from the Malay Peninsula, is well differentiated from the other subgroups

in all stages. The immature stages are normally found in bamboo internodes.

Zavortink: Orthopodomyia in Southeast Asia 11

ORTHOPODOMYIA WILSONI MACDONALD

(Figures lb, c, g, 6, 9)

Orthopodomyia wilsoni Macdonald 1958, Proc. R. ent. Soc. Lond. (B) 27: 121

(o*, ss L*); Zavortink 1968, Contr. Amer. ent. Inst. 3(2): 130 (o*,

2, P*, LF).

FEMALE. (Figs. lb, c, g) Head. Dorsolateral scales brown anteri-

orly, yellowish posteriorly; labial light scales cream-colored, band about

0.75 distance from base; palpus 0. 42 length of proboscis, light scales white,

in 2 patches beyond the base. Thorax. Pyrvabristles absent; pleural scales

largely cream-colored to yellow; ssp scale patch weakly developed, a simple

usually short row. Legs. Fore and mid tarsomeres 2 and 3 entirely dark

scaled or light scaled only at base; hind tarsomere 1 light or dark scaled at

apex, hind tarsomere 2 conspicuously white ringed at both ends, hind tarso-

mere 3 usually like 2, but sometimes entirely white scaled, hind tarsomere 4

all white. Wing. Vein Sc without presectoral light patch; 1A dark at base.

Abdomen. Light scales white or more frequently yellow or golden; additional

light scales usually in large to very large basal middorsal patch on tergum II

or terga II and III, and sometimes scattered basally or in narrow basal band

on remaining terga; sterna II-VII dark scaled with basolateral light patch

which becomes smaller on distal segments.

MALE. Head. Proboscis with additional light scales in preapical dor-

sal patch or oblique band; palpus 0.84 length of proboscis. Abdomen. Terga

more frequently and broadly banded and subdorsal median light patches larger.

Terminalia, (Fig. 6) Tergum VII lobe with apex truncate or emarginate and

serrate; basimere without strong, elongate specialized bristles distad of basal

mesal lobe; basal mesal lobe with 4 stout bristles and 1 finer; aedeagus moder-

ately sclerotized, pyriform in dorsal aspect, with a pair of separated ventral

teeth near apex, and without a conspicuous projection between or below ven-

tral parameres.

PUPA. (Fig. 9) Cephalothorvax. Hairs 1-5-C moderately developed,

1-3 usually single or double (1-3b), hair 4 usually single (1-4b), hair 5 usual-

ly single or double (1-3b); 7-C very strongly developed, usually double (2, 3b);

8,9-C moderately developed, usually single (single, double). Abdomen. Hair

2-II-IV fine, 2-II mesad of 5-II; 1-III usually 5, 6b(4-7); 5-IV usually 3, 4b;

5-V usually 2, 3b. :

LARVA. (Fig. 6) Head. Integument largely straw-colored to tan; hair

13-C moderately strong, usually with 6, 7b(5-8). Antenna. Moderately long.

Abdomen. Hair 6-I 7, 8b, 6-II usually 5-7b(5-8); 1-IIl moderately long, usual-

ly 2, 3b(1-3); 1-V very long, single; 1-VI equal to or longer than 4-VI; seg-

ment VI without sclerotized plate; siphon index usually 7. 0-12.0; hair 1-S lo-

cated 0.30-0.37 from base of siphon and usually 8-10b(6-10).

TYPE DATA. Orthopodomyia wilsoni Macdonald, holotype male with

associated larval and pupal skins (0348/4) in British Museum; type locality:

Ulu Gombak Forest Reserve, Selangor (Macdonald), MALAYSIA.

DISTRIBUTION. Specimens examined: MALAYSIA, Selangor, 10

males, 7 females, 14 pupae, 14 larvae, 14 rearings. THAILAND, Nakhon Si

Thammarat, Phangnga, Ranong, 2 males, 2 females, 3 pupae, 21 larvae, 3

rearings. Records from the literature: MALAYSIA, Selangor (Macdonald

1958). THAILAND (Zavortink 1968).

TAXONOMIC DISCUSSION. O. wilsoni is one of the most distinctive

of the Southeast Asian species. The adult has a poorly developed subspiracu-

lar scale patch and an unusually large basal light patch on abdominal tergum

II or terga II and III. In addition, the abdominal light scaling is more golden

or yellow in wilsoni than in any other species. The adult is also distinct from

all others except lanyuensis in banding of the hind tarsomeres. The terminalia

12 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

of the male differ from those of albipes, the only other Southeast Asian species

with a distinctly pyriform aedeagus, by lacking strong, elongate specialized

bristles on the basimere distad of the basal mesal lobe. The pupa has a dark

cylindrical trumpet and cephalothoracic setae 1-6, 8,9 relatively weakly devel-

oped, short and few branched. The shape of the larval comb scales is unlike

that of any other species. The larva also differs from all others except szam-

ensis by having a long thin siphon.

Most adults of wilsoni have conspicuous, subequal white bands at both

ends of hind tarsomeres 2 and 3. I have seen a single reared specimen from

the Isthmus of Kra which has hind tarsomere 3 entirely white scaled on both

legs. The form of larval hairs 1-M, T,I-IV and 13-T,II-V is quite variable;

these hairs may be long and relatively few branched to short and relatively

many branched. Hairy forms are unknown for the pupa.

O. wilsoni is apparently a relictual species; it has no close relatives,

but its affinities probably lie with @lbipes.

BIOLOGY. The immature stages occur in cracked or punctured bam-

boo internodes. There is one reared record of larvae having been collected

in the concave top of the fungus Polystictus xanthopus. Although Macdonald

(1958) did not recover this species from the same internodes as albipes in

Malaysia, the two species are frequently associated in collections from penin-

sular Thailand. Habits of the adults are largely unknown, but a male has been

collected resting on vegetation in a forest. Resting adults of both anopheloides

and andamanensis were also taken at the same time and place.

ALBIPES SUBGROUP

PUPA. Trumpet. Largely brown to dark brown, broad, attaining

maximum width in basal 0.33. Abdomen. Hair 1-II moderately developed,

short, usually with a central stalk from which 19-35 very fine dendritic or

forked branches arise, sometimes without stalk and with as few as 2 simple

branches; 5-V, VI long, extending to alveolus of hair 4 of second following

segment; 5-VI usually double (2-5b).

LARVA. Thorax. Hair 1-M, T 1-4b, usually long, much exceeding

3-M and 2-T. Abdomen. Hair 6-1,II with branches finer than 6-III, tapering

to fine apex and barbed from near base to near apex; posterior comb scales

narrow at apex and with a single strong but relatively short spine.

TAXONOMIC DISCUSSION. The albipes subgroup consists of a single

species found from eastern India to northern Borneo. The subgroup is quite

well marked in all stages. Bamboo internodes are the preferred breeding

Sites. |

ORTHOPODOMYIA ALBIPES LEICESTER

(Figures ld, 2, 4, 10)

Orthopodomyia albipes Leicester 1904, In Theobald, Entomologist 37:237 (¢,

¢); Barraud 1934, Fauna Brit. India, Diptera 5: 102 (¢, ¢, 3 Knight &

Mattingly 1950, Proc. ent. Soc. Wash. 52: 16 (o*, 9, P*, L*); Zavortink

1968, Contr. Amer. ent. Inst. 3(2): 133 (o*, 9, P*, L*).

FEMALE. (Figs. 1d, 2) Head. Dorsolateral scales brown anteriorly,

white posteriorly; labial light scales white, band at about middle; palpus 0.58

length of proboscis, light scales white, in 3 patches beyond the base. Thorax.

Pra bristles absent; pleural scales largely white, sometimes cream-colored;

ssp scale patch well developed, a simple moderately long row. Legs. Fore

and mid tarsomeres 2 and 3 usually light scaled at base only; hind tarsomere

Zavortink: Orthopodomyia in Southeast Asia 13

1 dark scaled at apex, hind tarsomere 2 usually with more extensive white

scaling at base than at apex (at least on outer surface), hind tarsomere 3 en-

tirely white or rarely with dark patch basad of middle, hind tarsomere 4 en-

tirely white. Wing. Vein Sc often with presectoral light patch; 1A white

scaled at base. Abdomen. Light scales white or rarely some yellow; terga

with additional light scales sometimes scattered basally or in basal bands;

sternum II mostly light scaled, II-VI dark scaled with light scales in baso-

lateral patch which becomes smaller on distal segments, in midventral apical

patch on segments II or III-V or VI, and sometimes in narrow basal band.

MALE. Head. Proboscis with additional light scales in preapical band

which extends further basally along ventral surface; palpus 0.75 length of pro-

boscis. Abdomen. As for female. Terminalia. (Fig. 4) Tergum VIII lobe

with apex rounded or truncate and serrate; basimere with moderately to strong-

ly developed elongate curved or sinuous bristles distad of basal mesal lobe;

basal mesal lobe with 4 or 5 stout bristles and 1 or 2 finer ones; aedeagus

moderately sclerotized, pyriform in dorsal aspect, with a pair of separated

ventral teeth near apex and without a conspicuous projection between or below

ventral parameres.

PUPA. wee 10) Cephalothorax. Hair 1-C moderately developed,

usually 2-4b(2-5); 2-C strongly developed, usually 2, 3b(1-3); 3-C very strong-

ly developed, usually 4-6b(4-9); 4-C strongly developed, usually 3, 4b(2-4);

5-C very strongly developed, usually 4, 5b(3-6); 7-C very strongly developed,

usually 3, 4b(3-5); 8, 9-C moderately developed, 8 usually single or double, 9

usually single. Abdomen. Hair 2-II-IV fine, 2-II mesad of 5-II; 1-III usually

6-8b(4-8); 5-IV usually 3-5b; 5-V usually 3b(2-4).

LARVA. (Fig. 4) Head, Integument largely straw-colored; hair 13-C

moderately developed, usually 4-6b. Antenna. Moderately long. Abdomen.

Hair 6-I, Il 5-7b; 1-III long, usually 2, 3b(1-4); 1-V very long, usually single

(single, double); 1-VI equal to or longer than 4-VI; segment VI without sclerot-

ized plate; siphon index usually 4. 0-6. 0(3.6-7.5); hair 1-S located 0.32-0. 40

from base of siphon and usually 9-11b(6- 14).

TYPE DATA. Orthopodomyia albipes Leicester, lectotype male with

terminalia slide in British Museum; type locality: Kuala Lumpur, Selangor,

MALAYSIA.

DISTRIBUTION. Specimens examined: INDIA, Bihar, 1 male, 1 fe-

male. MALAYSIA, Kedah, Kelantan, Pahang, Perak, Selangor, 246 males,

294 females, 266 pupae, 384 larvae, 265 rearings. EAST MALAYSIA, Sabah,

Sarawak, 2 males, 3 females. SINGAPORE, 1 larva. THAILAND, Chiang

Mai,. Kanchanaburi, Lampang, Nakhon Ratchasima, Nakhon Si Thammarat,

Narathiwat, pee ht Ranong, Surat Thani, Tak, 87 males, 66 females, 147

pupae, 372 larvae, 142 rearings. Records from the literature: INDIA,

Darjeeling District (Barraud 1934). MALAYSIA, Perak (Edwards 1928), Sabah

(Zavortink ey Sarawak (Barraud 1934), Selangor (Leicester 1904,

Macdonald 1958). THAILAND, Kanchanaburi, Ranong (Zavortink 1968). VIET

NAM (Borel 1930).

TAXONOMIC DISCUSSION. O. albipes, like the other bamboo inter-

node-breeding species, wilsoni, is also one of the most differentiated inSouth-

east Asia. The adult is distinctive in having hind tarsomere 2 usually more

extensively light scaled basally than apically and in possessing a midventral

apical light patch on abdominal sterna II or III to V or VI. Segment 4 of the

palpus of the female is elongated in albipes. In all but a very few specimens,

the added length separates the light scales at the apex of the segment from

those at the base, so that the palpus has 3 light patches beyond the base in-

stead of the 2 found in all other species. The male terminalia are character-

ized by the combination of a pyriform aedeagus and strong, elongate special-

ized bristles on the basimere distad of the basal mesal lobe. The pupa is

easily separated from the other species by the shape of the trumpet. The lar-

va differs from other species in shape of the comb scales and, to a lesser ex-

tent, in shape of the siphon.

14 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

I have seen specimens of albipes from West Malaysia and Thailand

which have a conspicuous white ring at both ends of hind tarsomere 2. In

other respects, including larval and pupal characters, these individuals are

typical albipes. Hind tarsomere 3, normally entirely white, sometimes has

a dark patch basad of the middle. The distimere of the male terminalia fre-

quently bears 2 claws. Larval hairs 1-M, T,III,IV and 13-T,II-V are quite

variable in length and number of branches. Hairy forms of the pupa are un-

known.

O. albipes is one of the more widely distributed Southeast Asian spe-

cies. Although it has no close relatives, it is probably derived from the same

phyllad as wilsonz. :

BIOLOGY. This species breeds in cracked or perforated bamboo

internodes and, less frequently, bamboo stumps. It is sometimes associated

with wilsoni in bamboo internodes and has been taken with siamensis in bam-

boo stumps. The immatures are encountered far more frequently than those

of the other Southeast Asian species. Females are known to bite man

(Macdonald 1958: 124).

ANOPHE LOIDES SUBGROUP

PUPA. Trumpet. Largely brown to dark brown, broadening gradually

from base. Abdomen. Hair 1-II moderately to strongly developed, moderate-

ly long to long, and with 8-35 mostly simple branches, some much finer than

others, arising at base; 5-V, VI short, extending to alveolus of hair 4 of first

following segment, in non-hairy forms, but long, extending to alveolus of hair

4 of second following segment, in hairy forms; 5-VI usually 3-7b(2-8).

LARVA. Thorax. Hair 1-M, T 1-4b, long to very long, much surpas-

Sing 3-M and 2-T. Abdomen. Hair 6-I, II with branches finer than 6-III, ta-

pering to a fine apex and barbed from near base to near apex; posterior comb

scales expanded at apex and with fan-like fringe of several elongate spines.

DISTRIBUTION. Species of this subgroup are found from India, south-

ern China, and southern Japan, south to Ceylon, Java, and northern Queens -

land. :

TAXONOMIC DISCUSSION. The anopheloides subgroup consists of the

3 species andamanensis, anopheloides, and papuensis. Only the first 2 of

these occur in the Southeast Asian area.

O. andamanenstis and anopheloides are very Similar to each other in

all stages and often cannot be separated. The male terminalia and immature

stages of the pair are well differentiated from the other species in the South-

east Asian area, but the adults are frequently indistinguishable from those of

Ssiamensis, a species in the flavicosta subgroup.

The anopheloides subgroup is undoubtedly the most recent in the albipes

group. The species are poorly marked and widespread.

BIOLOGY. The preimaginal stages of species in this subgroup are

commonly found in tree-holes, bamboo stumps, and artificial containers.

Adults have been collected resting on tree trunks and forest vegetation. Fe-

males of andamanensis apparently bite man.

ORTHOPODOMYIA ANDAMANENSIS BARRAUD

(Figures 1f, 8, 10)

Orthopodomyia anopheloides yar. andamanensis Barraud 1934, Fauna Brit.

India, Diptera 5: 102 (¢, 9).

Orthopodomyia andamanensis Barraud: Knight & Mattingly 1950, Proc. ent.

Soc. Wash. 52: 15 (o'*, 9, P*, L*); Zavortink 1968, Contr. Amer. ent.

Inst. 3(2): 141 (o*, 9, P*¥, L*).

Zavortink: Orthopodomyia in Southeast Asia 15

FEMALE. (Fig. 1f) Head, Dorsolateral scales brown anteriorly,

white posteriorly; labial light scales usually white, band at or distad of middle

or sometimes absent, preapical patch sometimes either absent or completed

ventrally, and sometimes with additional light scales scattered near base; pal-

pus 0.43 length of proboscis, light scales white, in 2 patches beyond the base.

Thorax. Pra bristles absent; pleural scales largely white or cream-colored;

ssp scale patch well developed, a simple moderately long row. Legs. Fore

tarsomeres 2 and 3 entirely dark scaled or light scaled at base or at bothends,

mid tarsomeres 2 and 3 light scaled at both ends; hind tarsomere 1 usually

white scaled at apex, hind tarsomere 2 with at most a narrow basal white ring

and with a narrow to moderately broad white ring at apex, hind tarsomere 3

entirely white or rarely with subbasal dark patch or ring, hind tarsomere 4

all white. Wing. Vein Sc without presectoral light patch; 1A sometimes white

scaled at base. Abdomen. Light scales white or cream-colored, rarely yel-

lowish; sometimes terga with additional light scales scattered basally or in

basal bands; sterna II- VII dark scaled with basolateral light patch which be-

comes smaller on distal segments.

MALE. Head. Proboscis with additional light scales in a moderately

broad to broad preapical band which extends farther basally along ventral sur-

face; palpus 0. 80 length of proboscis. Abdomen. Terga and sterna sometimes

with light basal bands. Terminalia. (Fig. 8) Tergum VIII lobe emarginate or

truncate, serrate; basimere without strong specialized bristles distad of basal

meSal lobe; basal mesal lobe with 4 or 5 stout bristles and 1-3 finer ones;

aedeagus weakly sclerotized, approximately fusiform in dorsal aspect, with-

out ventral teeth, but with a basal projection extending conspicuously ventrad

below ventral parameres.

PUPA. (Fig. 10) The hairs in the hairy form have more branches (in

italics in the following description) and are generally more strongly developed

than in the non-hairy form. Cephalothorax. Hair 1-C moderately developed,

usually 2,3b, up to 5b; 2,3-C moderately developed, usually 3-5b(2-6), up to

10b; 4-C moderately to strongly developed, usually 2-4b, up to 8b; 5-C strong-

ly to very strongly developed, 2,3b, up to 6b, 7-C very strongly developed, 2,

3b, up to 6b;8, 9-C moderately to strongly developed, usually 2,3b(2-5), up to

8b. Abdomen, Hair 1-II moderately developed, usually 16-25b, up to 35d,

2-TI-IV fine, 2-II mesad of 5-II; 1-II usually 12-14b(10-17); 5-IV 5, 6b, up to

8b; 5-V usually 4, “sh up to 7b; 5-VI usually 6, 7b(4-8).

LARVA. (Fig. 8) Head. Integument brown to dark brown; 13-C mod-

erately developed, 4-6b(4-7). Antenna. Short. Abdomen. Hair 6-I usually

— 9,10b(7-10), 6-II usually 9-11b(7-12); 1-III, V very long, single; 1-VI shorter

than 4-VI; segment VI without sclerotized plate; siphon index usually 4. 0-5. 6;

hair 1-S located 0.37-0.50 from base of siphon and usually 11, 12b(10-14).

TYPE DATA. Orthopodomyia anopheloides var. andamanensis Barraud

lectotype male with terminalia slide in British Museum; type locality:

ANDAMAN ISLANDS (Covell).

DISTRIBUTION. Specimens examined: ANDAMAN ISLANDS, 1 male,

1 female. INDONESIA, Java, 6 males, 10 females, 1 pupa; Kalimanian, 2

males, 1 female, 1 pupa, 1 larva, 1 rearing; Sumatra, 5 males, 7 females, 1

pupa, 1 larva. MALAYSIA, Perak, Selangor, 5 males, 8 females, 8 pupae, 8

larvae, 8 rearings. PHILIPPINES, Palawan, 1 male, 1 female, 2 pupae, 2

larvae, 2rearings. SINGAPORE, 2 males, 1 female, 1 pupa, 1 larva.

THAILAND, Chanthaburi, Chiang Mai, Chiang Rai, Khon Kaen, Lampang,

Nakhon Nayok, Nakhon Ratchasima, Ranong, Tak, Trang, 35 males, 15 fe-

males, 7 pupae, 15 larvae, 6 rearings. VIET NAM, 1 male, 1 female. Re-

cords from the literature: ANDAMAN ISLANDS (Barraud 1934). INDIA,

Darjeeling District (Barraud 1934). INDONESIA, Celebes (Knight & Mattingly

1950), Java, Kalimantan (Bonne-Wepster 1954), Sumatra (Brug & Bonne-

Wepster 1947). MALAYSIA, Perak (as albipes, Leicester 1908), Selangor

(Macdonald 1958). PHILIPPINES, Palawan (Knight & Mattingly 1950).

16 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

SINGAPORE (Barraud 1934). THAILAND, Chiang Mat, Chiang Rai, Khon Kaen,

Nakhon Nayok, Nakhon Ratchasima (Zavortink 1968), Lampang (Thurman 1959).

VIET NAM (Zavortink 1968).

TAXONOMIC DISCUSSION. The adult of andamanensis is apparently

indistinguishable from that of szamensis. It differs from the majority of those

of anopheloides by having hind tarsomeres 3 and 4 entirely white scaled. From

those rare specimens ofanopheloideswith hind tarsomeres 3 and 4 entirely

white scaled, it sometimes differs by having a narrower white ring at the apex

of hind tarsomere 2. The aedeagus of the male terminalia of both andamanensis

and anopheloides is quite different from that of the other species. It lacks

ventral teeth but has a basal projection between or below the ventral para-

meres. In andamanensis this basal projection is usually directed more ven-

trally than in anopheloides. Non-hairy pupae of andamanensis and anopheloides

differ from other Southeast Asian species by having hair 5-VI short and multi-

ple. The pupa of andamanensis is separated from that of anopheloides by hav-

ing a greater number of branches in hairs 1-III and 9-C. Larvae of the 2 spe-

cies have characteristically shaped posterior comb scales, long thoracic hair

1-M, T, and branches of abdominal hair 6-I, II finer than hair 6-IIl. The lar-

va of andamanensis usually has a greater number of branches in hair 6-I, II

than anopheloides. ,

I have seen 3 male specimens from Thailand that have a dark ring in

the basal portion of hind tarsomere 3; they agree with andamanensis in all

other characters and are probably that species. Hairy forms are known for

the pupa only. |

O. andamanensis and anopheloides are very Similar in all stages, are

sympatric over a broad area and are found in the same breeding sites. It is

possible that further study will show them to be only forms of a single species.

O. andamanensis occurs from east India to Java, Celebes and Palawan.

I have not seen the material upon which the Celebes record is based. It is

possible that the species there is the closely related papuensis rather than

andamanensis.

BIOLOGY. The immature stages of andamanensis have been taken

from tree-holes, bamboo stumps, and artificial containers; they are often as-

sociated with those of a@nopheloides. Females of the Celebes population are

known to bite man (Knight & Mattingly 1960: 16). Adults are sometimes taken

resting in the forest; such collections frequently contain anopheloides also.

ORTHOPODOMYIA ANOPHELOIDES (GILES)

(Figures le, 7, 11, 12)

aay en oe Giles 1903, Jn Wyville Thomson, J. trop. Med. 6: 315

a, 9*, L).

Orthopodomyia albipes var. nigritarsis Leicester 1908, Cul. Malaya :177 (¢,

?); Edwards 1913, Bull. ent. Res. 4: 239 (synonymy).

Orthopodomyia maculata Theobald 1910, Rec. Indian Mus. 4: 29 (o); Edwards

1913, Bull. ent. Res. 4:239 (synonymy); Barraud 1934, Fauna Brit.

India, Diptera 5: 101 (A, L; as variety of anopheloides); Knight &

Mattingly 1950, Proc. ent. Soc. Wash. 52: 9 (o*, P, L; as subspecies

of anopheloides); Thurman 1959, Univ. Md. Agric. Exp. Sta. Bull.

Pacif. Sci. Congr. 9:55 (as variety of anopheloides); Zavortink 1968,

Contr. Amer. ent. Inst. 3(2): 146 (Synonymy).

Orthopodomyia maculipes Theobald 1910, Mon. Cul. 5:470 (?*); Barraud, 1934,

Fauna Brit. India, Diptera 5: 101 (A, L; as variety of anopheloides);

Knight & Mattingly 1950, Proc. ent. Soc. Wash. 52: 10 (o*, 9, P*, L;

elevated to specific rank); Zavortink 1968, Contr. Amer. ent. Inst.

3(2): 146 (synonymy).

Zavortink: Orthopodomyia in Southeast Asia 17

Orthopodomyia manganus Baisas 1946, Mon. Bull. Bur. Hlth. Philipp.,

Manila 22: 35 (o*, 9, P*, L*); Knight & Mattingly 1950, Proc. ent. Soc.

Wash. 52: 7 (Synonymy).

Orthopodomyia (Orthopodomyia) nipponica LaCasse & Yamaguti 1948, Mosq.

Fauna Japan and Korea 2: 264 (o'*, 9*, P*, L*); Knight & Mattingly 1950,

Proc. ent. Soc. Wash. 52: 7 (synonymy).

Orthopodomyia (Orthopodomyia) lemmonae Thurman 1959, Univ. Md. Agric.

Exp. Sta. Bull. A-100: 58 (L*); Zavortink 1968, Contr. Amer. ent.

Inst. 3(2): 147 (synonymy).

Orthopodomyia anopheloides (Giles): Edwards 1913, Bull. ent. Res. 4: 239

(taxonomy); Barraud 1927, Indian J. med. Res. 14: 527 (c'*, 2);

Barraud 1932, Indian J. med. Res. 19: 1014 (P*, L*); Barraud 1934,

Fauna Brit. India, Diptera 5: 98 (o, 9*, P*, L*); Knight & Mattingly

1950, Proc. ent. Soc. Wash. 52: 7 (o*, 9, P*, L); Delfinado 1966, Mem.

Amer. ent. Inst. 7: 66 (o*, 9, P*, L*); Zavortink 1968, Contr. Amer.

ent. Inst. 3(2):146 (o*, 9*, P*, L*).

Orthopodomyia mcgregori of Knight & Chamberlain 1948, Proc. helm. Soc.

Wash. 15: 10 (P*; misidentification); Knight & Mattingly 1950, Proc.

ent. Soc. Wash. 52: 13 (¢, P, L; misidentification).

FEMALE. (Fig. le) Head. Dorsolateral scales brown anteriorly,

white posteriorly; labial light scales usually white, band at or distad of mid-

dle or sometimes absent, preapical patch sometimes either absent or com-

pleted ventrally, and sometimes with additional light scales scattered near

base; palpus 0.48 length of proboscis, light scales white, in 2 patches beyond

the base. Thovax. Pra bristles absent; pleural scales largely white or cream-

colored, rarely yellow-tinged; ssp scale patch well developed, a simple mod-

erately long row. Legs. Fore and mid tarsomeres 2 and 3 usually light scaled

at both ends; hind tarsomere 1 usually white scaled at apex, sometimes with

broad white ring, hind tarsomere 2 dark or light scaled basally and with a

moderately broad to very broad apical white ring, hind tarsomeres 3 and 4

rarely all white, more commonly 3 or 3 and 4 with dark patch or ring distad

of middle. Wing. Vein Sc without presectoral light patch; 1A sometimes

white scaled at base. Abdomen. Light scales white or cream-colored, rare-

ly yellowish; sometimes terga with additional light scales scattered basally or

in basal band; sterna II- VII dark scaled with light scales in basolateral patch

which becomes smaller on distal segments and sometimes in basal band.

MALE. Head, Proboscis with additional light scales in preapical dor-

sal patch or narrow to broad band which extends further basally along ventral

surface; palpus 0.78 length of proboscis. Abdomen. Terga and sterna more

frequently and extensively banded than in female. Terminalia, (Fig. 7) Ter-

gum VIII lobe emarginate or truncate and usually serrate; basimere usually

without strong specialized bristles distad of basal mesal lobe; basal mesal

lobe with 4-6 stout bristles and 1-3 finer ones; aedeagus weakly to moderately

sclerotized, approximately fusiform in dorsal aspect, without ventral teeth,

but with a basal projection extending conspicuously cephalad between ventral

parameres.

PUPA. (Fig. 11) As in andamanensis the hairs in the hairy form have

more branches (in italics in the following description) and are generally more

strongly developed than in the non- oki form. Cephalothorax. Hair 1-C

moderately developed, 2-3b, up to 4b; 2-C moderately developed, 1-3b, up to

7b; 3-C moderately developed, 3-5b, up to 10b; 4-C moderately to very strong-

ly developed, 2,3b, up to 6b; 5-C very strongly developed, 2-3b, up to 6b;

1-C very strongly developed, usually double, up to 6b; 8,9-C moderately to

strongly developed, usually single (single, double), 1-3b. Abdomen. Hair

1-II moderately developed, usually 12-19b . a up to 34b; 2-II fine and me-

sad of 5-II; 2-III, IV fine; 1-III usually 5-7b(3-11); 5-IV 3, 4b, up to 8b; 5-V

usually 3-4b(2-4), up to 7b; 5-VI usually 3-5b(2-6).

18 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

LARVA. (Figs. 7, 12) Head, Integument largely brown to dark brown;

hair 13-C moderately developed, 4-6b(4-7). Antenna. Short. Abdomen.

Hair 6-I usually 5-8b, 6-II usually 4-8b; 1-III, V very long and single in non-

hairy form, long and double in hairy form; 1-VI shorter than 4-VI in non-hairy

form, equal to or longer than 4-VI in hairy form, segment VI without sclerot-

ized plate; siphon index usually 3. 5-4. 8(3. 2-6. 4}. hair 1-S located 0. 36-0.50

from base of siphon and usually 10-12b(9-13).

TYPE DATA. Mansonia anopheloides Giles, lectotype male with ter-

minalia slide in British Museum; type locality: Dehra Dun, Uttar Pradesh,

INDIA. Orthopodomyia albipes var. nigritarsis Leicester, lectotype female

(76) in the British Museum; type locality: Pangkor Laut, Pevak (Daniels),

MALAYSIA. Orthopodomyia maculata Theobald, holotype male in Indian Mu-

seum; type locality: Maddathorai, Tvavancore (Annandale), INDIA. Orthopo-

domyia maculipes Theobald, lectotype female in British Museum; type local-

ity: ANDAMAN ISLANDS (Lowis and White). Orthopodomyia manganus

Baisas, holotype male with associated larval and pupal.skins (THIII-5) lost;

type locality: Llavac, Laguna, Luzon (Sunico), PHILIPPINES. Orthopodomyia

(Orthopodomyia) nipponica LaCasse & Yamaguti, lectotype female with asso-

ciated pupal skin in U.S. National Museum; type locality: Kyoto, Honshu,

JAPAN. Orthopodomyia (Orthopodomyia) lemmonae Thurman, holotype larva

(M398-1) in U.S. National Museum; type locality: Doi Chom Cheng of Doi

Sutep Range, Chiang Mai (Thurman), THAILAND. |

DISTRIBUTION. Specimens examined: ANDAMAN ISLANDS, 4 fe-

males. CEYLON, 4 males, 8 females. CHINA, Chekiang, 1 male, 1 female.

INDIA, Assam, Himachal Pradesh, Mysore, Uttar Pradesh, 5 males, 7 fe-

males, 5 pupae, 19 larvae, 8 rearing. INDONESIA, Java, 1 male, 1 female,

2 pupae, 5 larvae; Sumatra, 4 males, 1 female, 2 pupae. JAPAN, Honshu,

13 males, 5 females, 2 pupae, 22 larvae, 2 rearings. MALAYSIA, Perak,

Selangor, 13 males, 11 females, 19 pupae, 20 larvae, 19 rearings. EAST

MALAYSIA, Sabah, 9 males, 8 females, 13 pupae, 13 larvae, 14 rearings.

NEPAL, 1 larva. PAKISTAN, East Pakistan, 1 larva. PHILIPPINES, Leyfe,

Luzon, Mindoro, Negros, Palawan, 29 males, 25 females, 23 pupae, 56 lar-

vae, 25 rearings. RYUKYU ISLANDS, Iriomote Island, Ishigaki Island,

Okinawa, 30 males, 35 females, 28 pupae, 49 larvae, 28 rearings.

SINGAPORE, 3 males, 2 females, 1 pupa, 7 larvae. TAIWAN, 5 males, 11

females, 11 larvae. THAILAND, Chanthaburi, Chiang Mai, Kanchanaburi,

Khon Kaen, Nakhon Nayok, Nakhon Ratchasima, Nakhon Si Thammarat,

Phangnga, Prachuap Khiri Khan, Ranong, Surat Thani, Tak, Trang, 61 males,

58 females, 52 pupae, 156 larvae, 49 rearings. VIET NAM, 5 males, 3 fe-

males, 3 pupae, 12 larvae, 3 rearings. Records from the literature:

ANDAMAN ISLANDS (as maculipes, Theobald 1910b). CEYLON (as maculata,

Theobald 1910b). CHINA, ‘Chekiang (Wu 1936), Hainan (as maculata, Chu

1957), Hunan (Chang 1957), Yunnan (Chow 1949). INDIA, Assam, Eastern

Himalayas, North Bengal, South Bombay, Western Himalayas (Barraud 1927),

Malabar, North Kanara (Barraud 1932), Travancore (as maculata, Theobald

1910a), United Provinces (Giles 1903). INDONESIA, Java (as maculata,

Barraud 1934), Kalimantan (as maculipes, Haga 1924), Sumatra (as maculipes,

Brug & Edwards 1931). JAPAN, Honshu (as nipponica, LaCasse & Yamaguti

ta MALAYSIA, Pevak (as nigritarsis, Leicester 1908), Sabah (Zavortink

1968), Selangor (partly as andamanensis and maculipes, Macdonald 1958).

NEPAL, PAKISTAN, East Pakistan (Zavortink 1968). PHILIPPINES, Leyte,

Negros, Mindoro (Zavortink 1968), Luzon (as manganus, Baisas 1946),

Palawan (as mcgregori, Knight & Chamberlain 1948 and as maculipes, Knight

& Mattingly 1950). RYUKYU ISLANDS, Iviomote, Ishigaki (Bohart 1959),

Okinawa (Gentry 1957). SINGAPORE (as maculipes, Edwards 1926). TAIWAN

(Chow 1950). THAILAND, Chanthaburi, Khon Kaen, Nakhon Nayok, Nakhon

Ratchasima, Trang (Zavortink 1968), Chiang Mai (partly as lemmonae and

co Thurman 1959), Nakhon Si Thammarat(as maculipes, Edwards

Zavortink: Orthopodomyia in Southeast Asia 19

TAXONOMIC DISCUSSION. O. anopheloides and andamanensis are a

pair of very Similar species. The characters which separate them from the

other Southeast Asian species are discussed under andamanensis. The adult

of anopheloides is told from that of andamanensis by the usual presence of a

dark patch or ring distad of the middle of hind tarsomere 3 or hind tarsomeres

3 and 4 and by the usually broader apical white ring on hind tarsomere 2. Male

terminalia of anopheloides usually have the basal projection of the aedeagus

directed more cephalad than in andamanensis. The pupa of anopheloides dif-

fers from that of andamanensis by having fewer branches in hairs 1-III and

9-C. The larva of anopheloides is distinguished from that of andamanensis

by having fewer branches in hair 6-I, II.

O. anopheloides is the most variable and widespread of the Southeast

Asian species of Orthopodomyia. As aresult, it has several synonyms.

Leicester was apparently unaware of Giles‘ earlier description when he pro-

posed nigritarsis in 1908. O. maculata Theobald,1910a and maculipes Theobald,

1910b were based on specimens with different types of leg banding. O. manganus

Baisas,1946 and nipponica LaCasse & Yamaguti,1948 were considered to be

distinct from anopheloides largely on the basis of markings of the proboscis.

O. lemmonae Thurman,1959 was based on young fourth instar larvae of the

hairy form. Large series of anopheloides with associated larval and pupal

skins are now available from many parts of the range of the species; study of

these series has indicated that only one variable species is involved.

While most adults of anopheloides have a dark patch or ring on hind

tarsomeres 3 and 4, a few have a patch or ring on only hind tarsomere 3, and

even fewer have hind tarsomeres 3 and 4 entirely white scaled. The chaeto-

taxy of both the larva and pupa of this species is very variable; hairy forms,

individuals in which many of the setae are elongate, thickened and more highly

branched, are known for both stages.

O. anopheloides is the dominant species of Orthopodomyia in the Ori-

ental region. It is found throughout nearly the entire Southeast Asian region.

BIOLOGY. The immature stages of anopheloides have been found in

tree-holes, bamboo stumps, artificial containers and rock-holes. Adults have

been found resting on tree trunks and in vegetation. The aquatic stages may

occur with those of andamanensis and madrensis and adults may rest in the

same areas as those of wilsoni and andamanensis.

HYBRIDS

ANOPHELOIDES X ALBIPES

(Figure 12)

I have seen 4 males with associated larval and pupal skins from north-

ern Thailand, Chiang Rai, that I think are hybrids between anopheloides and

albipes. The adults resemble albipes, but differ from that species and all

other Orthopodomyia by having a patch of scales on each side of the clypeus.

The larvae and pupae are also similar to albipes, but have some less conspic-

uous features of anopheloides. The aedeagus of the male terminalia (Fig. 12)

is more or less intermediate between that of anopheloides and albipes.

20 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

NOMEN DUBIUM

ORTHOPODOMYIA MCGREGORI (BANKS)

Kerteszia mcgregori Banks 1909, Philipp. J. Sci. 4: 548 (o); Edwards 1932,

In Wytsman, Genera Insect., fasc. 194: 108 (reduced to synonymy with

albipes); Baisas 1946, Mon. Bull. Bur. Hlth. Philipp., Manila 22: 34,

35 (suggested leaving status uncertain until topotypic material was

available); Knight & Chamberlain 1948, Proc. helm. Soc. Wash. 15:10

(resurrected from synonymy with albipes); Zavortink 1968, Contr.

Amer. ent. Inst. 3(2): 154 (changed to status of nomen dubium).

TYPE DATA. Kerteszia mcgregori Banks, holotype male destroyed;

type locality: Basilan Island (Mcgregor), PHILIPPINES.

TAXONOMIC DISCUSSION. I am following Baisas (1946: 34, 35) in

considering mcgregori to be anomen dubium. Although Banks' description of

mcgregori is unusually detailed for descriptions of that time, it remains un-

certain to what taxon the name applies. However, the species is not albipes,

as thought by Edwards (1932: 108). The mcgregori of Knight & Chamberlain

(1948: 10) and Knight & Mattingly (1950: 13) is anopheloides.

ACKNOWLEDGEMENTS

I am grateful to Dr. Botha de Meillon, Responsible Investigator,

Southeast Asia Mosquito Project, Dr. Alan Stone, Agricultural Research

Service, U.S. Department of Agriculture and Dr. John N. Belkin, Depart-

ment of Zoology, University of California at Los Angeles, for their counsel

in the preparation of this paper. I thank Dr. Wallace A. Steffan, Bernice P.

Bishop Museum, Dr. Peter F. Mattingly, British Museum (Natural History),

Dr. William W. Macdonald, Liverpool School of Tropical Medicine, Dr. J.C.

Lien, Taiwan Malaria Research Institute, and Dr. Richard M. Bohart,

University of California at Davis, for loans of specimens. The illustrations

were prepared by Mrs. Elaine Hodges, Mr. L. M. Druckenbrod and illustra-

tors of the 406th General Medical Laboratory, Japan. The manuscript was

typed for offset reproduction by Miss Helle Starcke. |

LITERATURE CITED

BAISAS, F.E.

1946. Notes on Philippine mosquitoes, VII. Species found in the

jungles of Llavac. Mon. Bull. Bur. Hlth. Philipp. , Manila

22: 27-49.

BANKS, C.S.

1909. Four new Culicidae from the Philippines. Philipp. J. Sci.

4: 545-551.

BARRAUD, P.J. :

1927. A revision of the culicine mosquitoes of India. Part XIX.

The Indian species of Aedomyia and Orthopodomyia with de-

scriptions of two new species. Indian J. med. Res. 14:

523-532.

1932. The early stages of some Indian mosquitoes: Orthopodomyia.

Indian J. med. Res. 19: 1013-1017.

Zavortink: Orthopodomyia in Southeast Asia 21

BARRAUD, P.J.

1934. Family Culicidae. Tribes Megarhinini and Culicini. The

fauna of British India, including Ceylon and Burma. Diptera,

vol. 5. London, Taylor & Francis. 463 pp.

BELKIN, J.N.

196 The mosquitoes of the South Pacific (Diptera, Culicidae).

vol. 1. Berkeley, Univ. Calif. Press. 608 pp.

BOHART, R. M.

1959. A survey of the mosquitoes of the southern Ryukyus.

Mosquito News 19: 194-197.

BONNE-WEPSTER, J.

1954. Synopsis of a hundred common non-anopheline mosquitoes of

the Greater and Lesser Sundas, the Moluccas and New

Guinea. Spec. Publ. R. trop. Inst. Amsterdam 111. 147 pp.

BOREL, E.

1930. Les moustiques de la Cochinchine et du Sud-Annam. Monogr.

Coll. Soc. Path. exot. 3. 423 pp.

BRAM, R.A.

1967. Contributions to the mosquito fauna of Southeast Asia. II.

The genus Culex in Thailand (Diptera: Culicidae). Contr.

Amer. ent. Inst. 2(1). 296 pp.

BRUG, S.L. & J. BONNE-WEPSTER

1947. The geographical distribution of the mosquitoes of the Malay

Archipelago. Chron. nat. 103: 179-197.

BRUG, S.L. & F.W. EDWARDS .

1931. Fauna Sumatrensis. Culicidae (Diptera). Tijdschr. Ent.

74: 251-261.

CHANG, T.H.

1957. A preliminary record on the tree-hole species of mosquitoes

of Changsha, Hunan. Actaent. sin. 7: 213-222.

CHOW, C.Y.

1949. Culicine mosquitoes collected in western Yunnan, China,

during 1940-1942 (Diptera, Culicidae). Proc. ent. Soc.

Wash. 51; 127-132.

1950. Collection of culicine mosquitoes (Diptera, Culicidae) in

Taiwan (Formosa), China, with description of a new species.

Quart. J. Taiwan Mus. 3: 281-287.

Cau, FF. 1;

1957. Collection of megarhine and culicine mosquitoes from Hainan

Island, South China, with description of a new species.

Acta Zool. sin. 9: 145-163.

DELFINADO, M.D.

1966. The culicine mosquitoes of the Philippines, tribe Culicini

(Diptera, Culicidae). Mem. Amer. ent. Inst. 7. 252 pp.

22 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

DELFINADO, M.D. |

1967. Contributions to the mosquito fauna of Southeast Asia. I.

The genus Aedes, subgenus Neomacleaya Theobald in

Thailand. Contr. Amer. ent. Inst. 1(8). 74 pp.

1968. Contributions to the mosquito fauna of Southeast Asia. III.

The genus Aedes, subgenus Neomacleaya Theobald in South-

east Asia. Contr. Amer. ent. Inst. 2(4). 76 pp.

EDWARDS, F.W.

1913. Bei eynonynay in Oriental Culicidae. Bull. ent. Res. 4:

1926. Mosquito notes. -- Vi. Bull. ent. Res. 17: 101-131.

1928. Diptera Nematocera from the Federated Malay States muse-

ums. J. F.M.S. Mus. 14: 1-139. )

1932. Genera Insectorum. Diptera. Family Culicidae. Fasc. 194.

Brussels, Desmet-Verteneuil. 258 pp.

GENTRY, J.W.

1957. New mosquito distribution records from Okinawa. Mosquito

News 17: 82.

GILES, G.M.

1903. In F. Wyville Thomson, Notes on the Culicidae of Dehra Dun.

J. trop. Med. 6: 314-315.

HAGA, J.

1924. Aanteekening omtrent muskieten (II). Geneesk. Tijdschr.

Ned.-Ind. 64: 815-834. :

KNIGHT, K.L.

1968. Contributions to the mosquito fauna of Southeast Asia. IV.

Species of the subgroup Chrysolineatus of group D, genus

Aedes, subgenus Finlaya Theobald. Contr. Amer. ent.

Inst. 2(5). 45-pp.

KNIGHT, K.L. & R.W. CHAMBERLAIN

1948. A new nomenclature for the chaetotaxy of the mosquito pupa,

based on a comparative study of the genera (Diptera: Culici-

dae). Proc. helm. Soc. Wash. 15: 1-10.

KNIGHT, K.L. & P. F. MATTINGLY

1950. The Orthopodomyia anopheloides subgroup of mosquitoes

(Diptera, Culicidae). Proc. ent. Soc. Wash. 52: 1-20.

LACASSE, W.J. & 8. YAMAGUTI

1948. Mosquito fauna of Japan and Korea. Part 2. Kyoto, Off.

Surgeon, 8th U.S. Army. 273 pp.

LEICESTER, G. F.

1904. In F.V. Theobald, New Culicidae from the Federated Malay

States. Entomologist 37: 236-239.

1908. The Culicidae of Malaya. Stud. Inst. med. Res. F.M.S.

3(3): 18-261.

Zavortink: Orthopodomyia in Southeast Asia 23

LIEN, J.C.

1968. New species of mosquitoes from Taiwan (Diptera: Culicidae).

Part Il. New species of Tripteriodes, Orthopodomyia,

Culiseta and Uvanotaenia. Trop. Med. 10: 1-20.

MACDONALD, W. W.

1958. Description of a new species of Orthopodomyia Theobald

from Malaya, with a note on the genus in that countr

eer: Culicidae). Proc. R. ent. Soc. Lond. (B) 27:

-126.

THEOBALD, F.V.

1904. New Culicidae from the Federated Malay States. Entomolo-

gist 37:236- 239.

1910a. Second report on the collection of Culicidae in the Indian

Museum, Calcutta, with descriptions of new genera and

species. Rec. Indian Mus. 4: 1-33.

1910b. A monograph of the Culicidae or mosquitoes. vol. 5.

London, British Museum. 646 pp.

THURMAN, E.B.

1959. A contribution to a revision of the Culicidae of northern

a. Univ. Maryland Agr. Expt. Sta. Bull. A-100.

182 pp.

1963. The mosquito fauna of Thailand (Diptera: Culicidae). Proc.

IX Pacif. Sci. Congr. 9: 47-57.

WU, S.C.

1936. Further notes on the mosquitoes of Hangchow, Chekiang,

with description of one new species. 1935 Yb. Bur. ent.

Hangchow 5: 46-53.

ZAVORTINK, T. J.

1968. Mosquito studies (Diptera, Culicidae). VII. A prodrome

oF ee genus Orthopodomyia. Contr. Amer. ent. Inst. 3(2).

pp.

damanens

: as

madrens

= ZA: ~ 255

SS ee as “A A

Ly 5 Hy = AS

Y tribes £4 A ey

by) ie r

Uf fee L.

BS OAS

Yise/2 29 ©

> LSS

SOS

IT layUY

2 =

= v

dorsal ventral

Philippines

madrensis

ventral

Thailand

albipes

Yere SS

1} */ he

1011 4]

dorsal __— ventral b

Thailand

A VAM-T siamensis

ventral

dorsal

Malaysia

wilsoni

Thailand

anopheloides

Thastand andamanensis

Thailand Malaysia

siamensis wi/soni

spt PrN.

“ail

Thailand Phillippines

albipes andamanensis

Thailand Philippines

anopheloides madrensis

Thailand

pupa

(hairy form)

Thailand

(non-hairy form)

larva

Taiwan

anopheloides Frey clenen

(hairy form)

f _-

aedeagus

HYBRID

dorsal ventral anopheloides X albipes

Zavortink: Orthopodomyia in Southeast Asia 37

INDEX

Names of valid taxa included in the present study are in roman type.

Synonyms, valid extralimital species, misidentifications and nomina dubia

are in italic type. Italic numerals are used for the principal text reference,

roman numerals for secondary text references, and roman numerals in paren-

theses for figures.

albipes Leicester

albipes group

albipes subgroup

andamanensis Barraud

anopheloides (Giles)

anopheloides subgroup

anopheloides subgroup of

Knight & Mattingly

anopheloides x albipes

flavicosta Barraud

flavicosta subgroup

flavithorax Barraud

lanyuensis Lien

lemmonae Thurman

maculata Theobald

maculipes Theobald

madrensis Baisas

manganus Baisas

mcgregori (Banks) nomen dubium

mcgregorvi of authors

nigritarsis Leicester

nipponica LaCasse & Yamaguti

papuensis Zavortink

siamensis Zavortink

wilsoni Macdonald

wilsoni subgroup

hy By 8, 10, 12 18 Ae 49, 20,

Re eae vi

, 3, 6, 8

, £5, 8, 205 318, 2S.) $65 PY, 19 Ay @,

$. 4. 6, 6,’ & WRI te 17. 18,

19, 20 (1, 7, 11, 12)

3, 6, 10, 14

19 (12)

9, 6.44

3. 4, 5. 6,7, 8, 9, 40,11

i? 7849

16, 18, 19

9.4.5, 6.7, 8) 9: 10; 10H, 3, 12

17, 18, 19

16, 18, 19

17, 18, 19

3, 14, 16

3, 4, 5, 6, 9, 10, 14, 16 (5, 9)

$. 4,8. 6, 88) 4249-18 19 11° 6, 6)

3, 10

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. X.

‘CONTENTS

INTRODUCTION (on. 6) 0 eve 5s os Ore Gut oe ogee wn eens eee

GORE CHISEIL BOLE eB cae lbw ee we Ze Ae ea

KEYS TO THE WORLD SUBGENERA OF CULISETA

MALES AND PRVIAIES .<: 4°. Gos) 3 a. a aa PS

FOURTH STAGE LARVAE 2ee ea te Sere Ses

PUntenUS CUSEIe FO 6g oie ie 8 ok wt ee eS ees

niveitaeniata (Theobald). ........2ee.eee- ae anak

Subgenus Climacura Howard, Dyar & Knab ........

marchettei oe Jeffery RRUCMICK oa 28 ge oe

ACKNOWLEDGEMENTS. pe AO ea ehwce i ee eee

PCE eer Ee bee i Pre aS abe ew el eee

Eis aS hia eee ew Ge Se ew os 5 Pee wert

THE GENUS CULISETA FELT IN SOUTHEAST ASIA‘

N. V. Dobrotwor sky”

INTRODUCTION

The genus Culiseta was erected by Felt (1904) for a mosquito he had

previously described under the name Culex absobrinus at the same time he

erected Culicella for Culex dyari Coquillett. Until recently mosquitoes of

Felt's genera have been known under the generic name of 7Theobaldia. This

had been proposed by Neveu- Lemaire (1902) for Culex annulatus Schrank, a

species subsequently placed by Blanchard (1905) in another new genus

Theobaldinella. A related genus Pseudotheobaldia was described by Theobald

neg for niveitaeniata. However, Stone et al. (1959) pointed out that Fischer

1885) had already used the generic name Theobaldia and that the next available

name was Culiseta Felt (1904). The name Culicella has been retained as a

subgenus but the others have become synonyms. Edwards (1921) in a revision

of the mosquitoes of the Palaearctic region divided the genus into three sub-

genera Theobaldia, Culicella and Allotheobaldia. These were defined mainly

on larval structure. Edwards expressed the opinion that Leptosomatomyia

fraseri Edwards should probably be included in this genus and in 1930 he in-

troduced for it the subgeneric name Theomyia. Later Culex melanurus

Coquillett was also transferred to the genus with the subgeneric name

Climacura Howard, Dyar & Knab (1915). These five subgenera were listed in

the Genera Insectorum (Edwards 1932), and with two new ones, Austrotheobaldia

(1954) and Neotheobaldia (1958) introduced by Dobrotworsky for Australian spe-

cies make up a total of seven recognized by Stone et al. (1959). Edward's as-

sessment of the importance of larval characters for the recognition of subgen-

era has been widely accepted. Maslov (1964), however, believes that more at-

tention should be paid to adult structure. With this approach, he has limited

the genus Culiseta to include only the four subgenera Culiseta, Culicella,

Climacura and Neotheobaldia;the remaining three, Allotheobaldia, Austro-

theobaldia and Theomyia he regards as full genera.

In my opinion Maslov's treatment of the genus is not justified. No

doubt there are different degrees of relationships between the various subgen-

era and some are clearly more specialized than others but there are advan-

tages in employing larger generic concepts. As Edwards (1932) has pointed

out, this means that "the wider relationships of the species are more clearly

indicated" and that "limits can more readily be assigned to the genera than in

the case of more numerous and smaller groups."

In this review I shall follow Belkin (1962) and treat the tribe Culisetini

(Stone 1957) as monotypic with the single genus Culiseta. The genus contains

mosquitoes with generalised characters which place them amongst the nearest

living representatives of the primitive stock, or stocks, of the Culicinae

(Edwards 1932; Belkin 1962; Marks 1968). It includes seven subgenera, which

are based mainly on larval characters, 35 species and seven subspecies.

5 beeen gs

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

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

of the Surgeon General, Washington, D.C.

Department of Zoology, University of Melbourne, Parkville, Vic. 3052,

Australia.

40 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

GENUS CULISETA FELT

Theobaldia Neveu- Lemaire 1902, C.R. Soc. Biol., Paris 54: 1331 (non

Fischer 1885). Orthotype: Culex annulatus Schrank.

Culiseta Felt 1904, Bull. N.Y. St. Mus., no. 79: 391c. Orthotype: Culex

absobrinus Felt.

Theobaldinella Blanchard 1905, Les Moustiques :390. Diatype: Culex

annulatus Schrank.

Pseudotheobaldia. Theobald 1907, Mon. Cul. 4: 271. Haplotype: nzveziaeniata

Theobald.

FEMALE. Medium sized to large. Head. Eyes almost touching; ver-

tex clothed mainly with narrow, curved decumbent and upright forked scales;

lateral scales broad, flat; antenna shorter than proboscis; palpus always short,

5-segmented; segment 5 short or minute; proboscis moderately long, not swol-

len at apex. Thorax. Anterior pronotum small, widely separated, usually

with bristles and scales; posterior pronotum with several bristles and some-

times with scales; scutum clothed with narrow scales; acrostichal and dorso-

central bristles numerous but not very long; scutellum usually with narrow

scales, rarely with broad; pleural scales varied in shape and distribution;

spiracular bristles present, usually numerous and pale in colour; in some

species these reduced in number to a few or a Single one. Legs. Bristles

well developed on femora and tibiae; tarsi all dark, banded, or pale apically;

claws of all legs small, subequal, simple; pulvillae absent. Wing. Frequently

with dark pattern of scales; wing membrane with distinct microtrichia; cell Rg

longer than vein R9,3; vein 1A somewhat sinuous; base of Sc ventrally with

patch of hairs and sometimes with a few scales as well (except fraseri which

have no hairs ventrally on base of Sc); alula and squama with marginal row of

hairlike scales or bristles. Abdomen. Terga, except tergum I, and sterna

almost completely scaled; tergum I always with numerous long bristles; abdo-

men blunt tipped, segment VII not retractile; cercus and post- genital plate

short and broad; three large, equal spermathecae.

MALE. In general, similar to female. Head. Antenna strongly plu-

mose; palpus variable in length, usually as long as proboscis (about a half

length of proboscis in fyaseri and two-thirds of it in longiareolata); last two

segments of palpus sometimes turned upwards or more or less swollen.

Terminalia. Apical margin of tergum VII with (in longiareolaia) or without

stout spines; lobes of tergum IX with several setae; basimere usually rather

long, more or less conical; basal mesal lobe present; subapical lobe usually

absent; distimere simple, usually with single, simple or bifid terminal ap-

pendage, longiareolata. alone has two terminal appendages; phallosome simple;

aedeagus conical, rarely with small teeth at top and only in marchettet with

prominent teeth; paraproct strongly sclerotized with one or more teeth; clasp-

ette absent.

PUPA. Cephalothorax. Hairs 8-C and 9-C caudad of trumpet; respi- |

ratory trumpet short with large opening. Abdomen. Hair 1-1 a well developed

dendritic. Paddle. Usually more or less finely serrate and with one or two

hairs at tip of midrib, in fyaseri these hairs are absent.

LARVA. Head. Usually wider than long; antenna varied in length;

shaft spiculate; hair 1-A varied in position, always well developed and branch-

ed; mouth brushes numerous, all filamentous or inner serrated distally; men-

tum broad, triangular with numerous teeth; hairs 5-7-C well developed, 5-C

frequently single. Thorax. Hairs 1-3-P and 9-12-P,M, T on large common

tubercles; all other large hairs on more or less distinct separate basal tuber-

cles or plates. Abdomen. Hairs 6-I-VI of similar length; hair 7-1 long, mark-

edly different from 7-II-VI; 2-VII and 4-VIII usually single or 2-branched; 1,

3,5-VI usually multibranched; comb scales in triangular patch or a single

row, no sclerotized plate; siphon varied in length; acus present; pecten usually

Dobrotworsky: Culiseta in Southeast Asia 41

of strong teeth (only in subgenus Neotheobaldia pecten teeth in form of hairs);

hair 1-S at base of siphon ane in subgenus Austrotheobaldia 1-S half-way

along siphon); accessory hairs la-S present or absent; valves small; saddle

almost always complete (only in subgenus Allotheobaldia saddle incomplete);

3-X usually single; ventral brush of 5 or more pairs of branched hairs on grid

and frequently 1-7 precratal hairs; anal papillae usually narrow, pointed.

EGGS. In most species eggs rounded at one end and tapered at the

other (in some species both ends pointed); some species have eggs with short

transparent stem; eggs laid usually in raft on water surface or singly on ground

above water level.

DISTRIBUTION. The subgenus Culisefa is confined to the Holarctic,

Culicella occurs in the Holarctic and Australian regions and Neotheobaldia is

restricted to Australia. Of the three monotypic subgenera, 'Theomyia is re-

stricted to the tropics of Africa, Austrotheobaldia is found in eastern Australia

and Tasmania, while Allotheobaldia is distributed across the southern part of

the Palaearctic from the Azores to Central Asia through the Ethiopian region

into India and Pakistan.

The subgenus Climacura has a remarkable distribution; one species oc-

curs in the eastern and central parts of the United States, one in West Malaysia,

one in eastern Australia and two in New Zealand. As Marks (1968) has pointed

out, the distribution pattern of Culiseta is a relict one, but there is no unanim-

ity of opinion about the place of origin of the genus.

Maslov (1967) has postulated a northern origin which is centered in

"Bering Land" where, during the end of the Mesozoic and early Tertiary, the

climate was temperate. From this centre protoculisetines dispersed to the

southwest and to the east and southeast across the tropics to Australia. How-

ever, Maslov's basic assumption that the subgenus Culiseta is the most primi-

tive one does not seem to be justified. Surtees (1959) who made an extensive

study of the structural and functional adaptations of the mouthparts of mosquito

larvae concluded that filter feeding is primitive. This method is found in the

members of the genus Culiseta, but not in the subgenus Culiseta. Here the

larvae are browsers, and their mouth brushes are modified and adapted for

scraping particles from submerged surfaces and on this basis the subgenus

cannot be regarded as primitive. Marks (1968) has argued that the least spe-

cialised of the Australian representatives of the Culicella group are more

primitive than the northern ones. On this basis and on Brundin's (1966) con-

clusion about an austral centre of origin of the family Chironomidae, Marks

has postulated a southern centre of origin for Culiseia. However, Darlington

(1970) has convincingly shown that the idea that primitive groups mark places

of origin is erroneous. Further, it may be doubted whether a parallel can be

drawn between chironomids and culicids. Brundin believes that the chirono-

mids "originated in cool running waters;" their present distribution is mark-

edly amphitropical (Brundin 1967). Culicid larvae which, in contrast to chiro-

nomids, are air-breathers, mostly favour still water and the greatest number

of present day genera and species are tropical.

Belkin (1962) believes that intercontinental areas of the Old and New

World, particularly those in the tropics, have been the places of origin and

evolution of new types of mosquitoes. On the available evidence, it seems

likely that Culiseia evolved in the tropics (Dobrotworsky 1965) and subsequently

spread to the northern and southern temperate regions, being progressively

displaced by later evolving elements.

There is no doubt that mosquitoes from tropical groups have dispersed

te southern Australia and Tasmania and become adapted to cold climatic con-

ditions. Anopheline mosquitoes, for example, entered Australia from the

north, probably during the Pleistocene (Mackerras 1950), but A. stigmaticus

Skuse now breeds in cold water (129-139 C) in mountainous areas. Again, al-

though species of Aedes of the subgenus Finlaya belong to the northern ele-

ment of the Australian fauna (Mackerras 1950), vrubrithorax (Macquart) is able

Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

“ALOYELIOGL] Ul UOISSTUSUBI} TeyJUOUTTIedxe Aq °D

‘OJINDSOU UT SNITA JO UOTJVdT[AT}NU SuTyerjsuoWsp Aq “gq

“SUdTITOOdS yYUSNeD pTIM Wor ATaAODNI AG °Y

rita nee asd PATER Pres ERE cs

9G6T WqTANA Es | bobo el STIN SOM 010109 AMISUO]

ve babbes lersdas i Pivives (Geis:

nt bb ree UE ES: Petes Hes Pee ie

YysnoT[s ‘DADULOUL

Alia

tt ee kee:

uMOJSOWeL

batt ait Peter

ede: =| sa

T= Sane

snit Tepe Sotded be vig e.

SEPET SoADSY BY uowUIey

896 J SoA00Y

8961 e0meT

6S6T SSeH 4 UeploH

ASPET SeaAvoy Y uoOWWeH

«| <| <| al <

4 2 6) BS

Bevel sere 7 UOWUIE CREWE

"Te 19 UOWUILTY LY

Bes &

1961 ‘Te 30 SOkEH {GG6T “Te 10 SUIISSTS

PI6T ASUTUM

oF GOOF [ee Seh8H sob 11%

J8 USPIOH -PG6T “TS6T “Te 39 UleTroquieyoO

SHSNUIAOGUV AHL GNV VLYSITNO AO SHIOddS

~ ee Baaiaae

Dobrotworsky: Culiseta in Southeast Asia 43

to complete its development through to the adult stage at temperatures as low

as 10°C. The present pattern of distribution of Culiseia is entirely consistent

with a tropical origin.

BIOLOGY. Some species are restricted to treeless plains, others to

forests or mountains, but a fair number do not show preference to a certain

type of country. The majority of species breed in ground pools, bogs, ponds,

marshes, edges of streams, rarely in treeholes. Some of the Australian spe-

cies breed underground in the burrows of land crayfishes (Engaeus spp. ).

Only one species, fvaseri is restricted to treeholes. A few breed under do-

mestic conditions in artificial containers. In the northern areas, most species

overwinter as hibernating adults, but some species do so as well in the larval

stage; in the southern areas breeding may be almost continuous during the

winter. Australian species overwinter mostly in the larval stage; New Zealand

species both as adults and larvae. Biting activity in most species begins after

sunset, but some are active during the day. Several species attack man as

well as domestic animals. In nature they feed on wild mammals but some

feed on birds and may prefer avian hosts.

MEDICAL IMPORTANCE. Several species of Culiseta have been found

to be naturally infected with arthropod-borne viruses (arboviruses). The table

shows that members of the genus are deeply involved in this field and suggests

that they certainly deserve further examination in this respect. The species

have mostly been incriminated through the isolation of viruses from wild

caught specimens but wherever transmission has been attempted it appears to

have been successful. It is of further interest to note that melanura was con-

sidered to be the primary vector during the great epidemic and epizootic of

eastern equine encephalomyelitis which occurred in New Jersey, USA in 1959.

Blanchard (1905) suggested that longiareolata might be involved in the trans-

mission of the bacillus of Malta fever. Maslov (1967) reports the successful

laboratory infection of bergrothi with the microfilariae of Dirofilaria immitis

in the Amur region of the USSR.

KEYS TO THE WORLD SUBGENERA OF CULISETA

MALES AND FEMALES

1. No hairs at base of subcosta on underside of wing;

male palpus barely half as long as proboscis and :

_ with only a few long bristles atits tip. ......... Theomyia

Hairs present at base of subcosta on underside of

wing; male palpus as least two-thirds the length

Gf PPODOSCIE. hie 2 Go iieecw iets Eigen ay age Whe sO em ae ge een ae 2

2(1). Postspiracular area with a few fine scales .... Austrotheobaldia

Postspiracular area bare or witha few hairs .......... 3

3(2). Tibia lined with white; male palpus shorter than

proboscis; tergum IX with a pair of long pro-

COBROS 5 Vo hee Re ee ee es -..- Allotheobaldia

Tibia not lined with white; male palpus at least

as JONG AS PrOBOSCI6 05. ke ee eae ae gk aii Sw ad Genet aki ee 4

4(3). Base of subcosta on underside of wing with scanty

hairs and not more than 5 spiracular bristles. ......... 5)

Spiracular bristles and hairs on underside of

WINE. TUINOPOUS nie cd 0a desde & eel wae ee. 8 wees 6

44, Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

5(4). Anterior pronotum without scales or with very

long hairlike scales; posterior pronotum with

bristles and fine hairlike scales. ........2.-. Climacura

Australia only. Culicella.

Anterior and posterior pronotum with pale

curved scales and bristles... .... 2c cece Neotheobaldia

Australia only. . Culicella

6(4). Cross-veins approximated, usually m-cu in

line with r-m but sometimes m-cu is well

before r-m; last segment of male palpus more

or less swollen, not turned upwards; basimere

of male terminalia with subapical lobe or ous

OG TIAPS Th 106 GAACE te Pe e o BE S R ge aes Culiseta

Cross-vein m-cu always well before r-m; last

segment of male palpus may be more or less

swollen; if not, often turned upwards; basimere

of male terminalia without subapical lobe

48 Re ee ar iS Eo 68 A RR eee a not Australia... 3°. “Cilitela

FOURTH STAGE LARVAE

1. Siphon with hair 1-S halfway along cgen ee aa a es

Pino With Nai t-9 Al DHSS i Ne PD a Oe

2(1). Siphon with pair of basal hairs only. .......... Se aes 3

Siphon with hairs la-S along ventral side of |

Siphon in addition to basal pair .......2.2-s+ee-. Climacura

3(2). Pecten of spine-like scales ........2-6.e06-. Mg REP i 4

Pecten of hair-like scales only ........... Neotheobaldia

4(3). Siphon long and slender; index 6-8 .........4.ee-8 wieale 5

Siphon short; index’2.5 "S375 vee a ete. RD rer ee aT oats 6

5(4). Antenna long; hair 1-A well beyond middle ........ Culicella

Antenna short; hair 1-A near middle; siphon

heavily sclerotized, almost black; spines of

pecten bifid from base .........se.2- iodine ted 7 § Theomyia

6(4). Siphon with pecten of spines only; saddle small

not forming complete ring ..%... 4.462022 se-e A llotheobaldia

Siphon with row of hairs beyond pecten; saddle |

forming complete ring ........ A Wis Sue eee ee 8 Culiseta

SUBGENUS CULISETA FELT

Culiseta Felt 1904, Bull. N.Y. St. Mus. no. 79: 391c. Orthotype: Culex

absobrinus Felt.

FEMALE. Head, Palpus with segment 5 sometimes distinctly white.

Thorax. Anterior pronotum with bristles and a few scales; posterior prono-

tum with posterior bristles and with broad scales or some narrow ones; spi-

racular bristles usually more than 7 in number, a few scales may also ‘be pres-

ent at base of these; scutellum usually with narrow scales but niveitaeniata

has broad scales. Legs. Tarsi all dark or with white basal bands. Wing.

Dobrotworsky: Culiseta in Southeast Asia 45

Wings often spotted; base of Sc ventrally with large patch of hairs; cross-

veins usually approximated. Abdomen. Tergum I with patch of scales and

long bristles; remaining terga with more or less distinct basal pale bands.

MALE. In general similar to female. Head. Palpus about as long as

proboscis or longer than it; usually with very long bristles; last segment more

or less swollen but not turned upwards. Terminalia. Lobes of tergum IX usu-

ally with long setae; basimere with basal mesal lobe relatively small, conical,

bearing numerous setae on sides and a few apical spines; subapical lobe usu-

ally more or less distinct or in place of it there is a patch of hairs; distimere

with single, usually bifid terminal appendage; 2 plates of aedeagus usually

separate, strongly sclerotized.

PUPA. Tip of midrib of paddle with single hair.

LARVA. Head. Head relatively not very large; antenna short or of

moderate length; hair 1-A near middle, relatively short; hairs 2-6A usually

short and on tip of antenna; some of mouth brushes serrated distally. Abdomen.

Comb scales in triangular patch; siphon relatively short with index 2.5-3.5;

pecten of strong teeth, with a row of hairs beyond; saddle forming complete

ring; hair 2-X multibranched; ventral brush 4-X usually with a few tufts pierc-

ing saddle.

EGG. Eggs laid in rafts on water surface.

CULISETA (CULISETA) NIVEITAENIATA (THEOBALD)

(Figures 1, 2, 3, 4)

Pseudotheobaldia niveitaeniata Theobald 1907, Mon. Cul. 4: 272 (o*).

Theobaldia niveitaeniata (Theobald), Barraud 1924, Indian J. med. Res. 12:

141; 1934, Faun. Brit. India, Diptera 5: 91 (o*,9, L*); Edwards 1932,

in Gen. Insect., Fasc. 194: 104; Stackelberg 1937, Faune de 1'URSS,

Ins., Dipt. 3(4); Qutubbudin 1952, Proc. R. ent. Soc. Lond. 21: 39 (L).

Theobaldia kanayamensis Yamada of Liu & Feng (mec Yamada) 1956, Acta ent.

sinica 6(3): 335. Synonymized by Maslov (1964, 1967).

Culiseta (Culiseta) niveitaeniata (Theobald), Stone, Knight & Starcke 1959,

Synop. Cat. Mosq. World 6: 219; Maslov 1964, Rev. d'Ent. de 1'URSS

43(1): 206; 1967, Opred. Faune SSSR 93: 145 (o*,?, L*).

Theobaldia (Theobaldia) sinensis Meng & Wu 1962, Acta ent. sinica 11(4): 383

(¢*, L*), Synonymized by Maslov (1964, 1967).

Culiseta (Culiseta) lishanensis. Lien 1968, Trop. Med. 10(1): 6 (o*, 9*, P*, L*).

New synonymy.

FEMALE. Head, (Figs. 1, 2) Vertex with pale decumbent scales;

upright forked scales mostly dark; lateral scales flat, broad, pale; palpus

dark brown, about 1/4 length of proboscis; some pale scales dorsally on seg-

ment 3; segment 5 small, but distinct, nude, pale; pedicel with broad white

scales mesially. Thovax. (Figs. 1, 2) Anterior pronotum with broad pale

scales; posterior pronotum with relatively narrow white scales dorsally and

broad white scales on lower part; scutum clothed with narrow, curved golden

scales; small patch of narrow white scales in front of wing root; scutellum

with broad creamy-white scales; 6-8 spiracular bristles and sometimes 2-4

scales at base of bristles; pleural bristles pale, scales broad, white. Legs.

(Fig. 2) Dark brown; coxa and trochanter with patch of broad pale scales;

posterior surface of femur of fore and mid leg pale from base almost to tip;

anteriorly fore femur with subapical white spot, hind femur white except for a

subapical black ring and a dark line along dorsal side and a white kneespot and

an apical tibial ring on all legs; tarsi dark, first tarsomere slightly paler ven-

trally at base in some specimens; tarsal claws as in Figure 1. Wing. (Fig. 1)

Wing membrane with darker patches in regions of cross-veins and base of fork

46 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

cells; veins with narrow dark scales; cross-veins r-m and m-cu closely

approximated. Abdomen. (Figs. 1,2) Terga brownish-black; tergum I with

patch of white scales medially; II-VI. with whitish basal bands, which on terga

IV and V narrows in middle; bands broken in middle on terga VI and VII.

MALE. In general similar to female. Head, (Fig. 2) Palpus dark

brown, paler at joints, about as long as proboscis; segments 3 and 4 with long

bristles, segment 5 widened and spatulate with a few strong, rather short

bristles at tip; tarsal claws as in Figure 1. Terminalia. (Fig. 3) Tergum

VUI without short stout spines on apical margin; each lobe of tergum IX with

8-13 setae; basimere narrow, about 3 times as long as broad at base and with-

out scales; basal mesal lobe conical, relatively small, with 2 strong bristles,

sometimes bent almost at right angles, and about 20 small fine setae; subapi-

cal lobe prominent with about 8 bristles, 4-7 of these stronger and with bent

tip; distimere about half as long as basimere, terminal appendage relatively

short, bifid; aedeagus simple with narrow pointed tip, sclerotized on ventral

side; paraproct with 1 long and 3-4 short teeth apically and 3-4 cercal hairs.

PUPA. (Fig. 3). Cephalothorax. Respiratory trumpets about 2 1/2

times as long as pinna. Hair 8-C single; 9-C single or 2-branched. Abdomen.

Hairs 1,5-IV-VII single; 5-IV-VI very long, on VII shorter and weaker ; 1-IV-V

as long as 5, shorter and weaker on VI, VU; 6-I-VI single. Paddle. Oval,

with posterior margin spiculated; hair 1-P single or with 2 branches.

LARVA. (Fig. 4) Head. Antenna short, about 1/2 length of head, with

Sparse, minute spiculation; hair 1-A inserted about at middle of shaft, with

, 4-6 slightly plumose branches, about 1/3-1/2 as long as shaft; hair 4-C small,

fine, with 2-4 simple branches; 5-7-C plumose, 5-C 6-7b, 6-C with 3 unequal

branches, median longest and strongest, 7-C 8-12b, 8-C single, 9-C 2-5b, |

10-C with forked tip; median mouth brush hairs serrated distally; mentum with

12-13 lateral teeth on each side. Thorax. Prothoracic hairs more or less

plumose; pleural groups of hairs well developed; hairs 1-3-P arising from a

sclerotized plate, 1-P long, 3-5b, 2-P single, long, 3-P shorter than 2-P,

6-9b, 4-P 6-10b, 5,6-P single, 7-P 5-9b, 8-P 4-5b, 9-P 4-7b, 10,12-P single,

long, 11-P very short, multibranched; 5-7-M single, long, 8-M 6-10b, plu-

mose, on sclerotized plate; 7-T 13-15b, plumose, 9-T plumose, 8-12b, 13-T

4b. Abdomen. Hair 6-I,II 4-6b, 6-III-IV 2b; I-VI 3b, 2,4-VII single; 3-VHI

7-12b, plumose, 5-VIII 4b; comb of about 40 scales; siphon index about 3.0,

hair 1-S inserted at base 5-10b, plumose; pecten with 12-20 spines followed

by an even row of 12-15 hairs extending to near apex of siphon; anal segment

completely ringed by saddle; hair 1-X 2-4b, 2-X 7-8b; 3-X 1, 2b; ventral brush

consisting of 15 tufts 1-3 of which precratal, 1 may be inserted in the saddle;

anal papillae slender, tapering, more than twice as long as saddle

EGG. Unknown.

TYPE DATA. Pseudotheobaldia niveitaeniata Theobald 2 male cotypes

in the British Museum. Type locality: Dehra Dun, United Provinces, INDIA.

Culiseta lishanensis Lien, holotype male and allotype female with associated

larval and pupal skins; paratypes 3 males and 6 females in Provincial Malaria

Research Institute, Taipeh, Taiwan; 1 male and 1 female in USNM. Type

locality: Lishan, Taichung Hsien, TAIWAN.

DISTRIBUTION. Specimens examined: INDIA, Punjab, Kasauli 3

males, 4 females; Almorva, Kausani 1 female, 4 rearings. TAIWAN 6 males,

5 females, 6 rearings. Records from literature: INDIA, Punjab, Dehra Dun, ©

Murree, Theog on Hindustan- Tibet road 2.439 m; Uttar Pradesh, Naini Tal,

Muktesar. TIBET, Yatung, near Sikkim border 3.658 m. (Barraud 1934).

CHINA, North East, Central and South (Maslov 1967).

TAXONOMIC DISCUSSION. In China, this species was misidentified by

Liu & Feng (1956) as Theobaldia kanayamensis Yamada. Later it was de-

scribed as a new species, Theobaldia sinensis, by Meng & Wu (1962) but

Maslov (1964, 1967) has pointed out that there are no consistent differences

between their species and niveitaeniata, which is more variable than Meng &

Dobrotworsky: Culiseta in Southeast Asia 47

Wu realized. Lien (1968) described lishanensis from Taiwan as very closely

related to niveitaeniaia but distinct from it. I have examined the paratypes

and also specimens collected at Alishan, Taiwan, and have found that there

are no Significant differences between them and niveitaeniaia. from India. It

should be noted that Lien's description and drawings (his Figure 1E) of the

terminalia of lishanensis are inaccurate; the terminalia does not differ from

that of niveitaeniata. On these grounds I have relegated lishanensis to the

synonymy. ;

BIOLOGY. The larvae are found in mountainous areas at an elevation

of 1600 m. to 3658 m. They live in clear or polluted water in a wide variety

of habitats such as ground and rock pools, pits, ditches, seepages, shallow

wells and artificial containers. The larvae were usually present during the

cooler part of the year from November to May and generally disappear during

the rest of the year. According to Meng & Wu (1962) the larvae can survive in

frozen pools. The adults I have seen, were collected in March and April. In

China niveitaeniata bites cattle, water buffalo and man (Meng & Wu 1962).

SUBGENUS CLIMACURA HOWARD, DYAR & KNAB

Climacura Howard, Dyar & Knab 1915, Mosq. N. and C. Amer. 3: 452.

Orthotype: Culex melanurus Coquillett.

FEMALE. Head. Palpus with segment 5 small, round or elongate.

Thorax. Anterior pronotum with bristles only; posterior pronotum with poste-

rior bristles and very narrow scales; 1-7 spiracular bristles; scutellum with

narrow scales. Legs. Tarsi all dark or with distinct basal bands on hind

legs. Wing. Wing with or without spots; base of Sc with reduced number of

ventral hairs; cross-vein m-cu well before r-m. Abdomen. Tergum I with

setae only; terga II- VIII usually dark scaled; in marchetiei terga V- VII have

basal pale bands.

MALE. In general similar to female. Head, Palpus as long as pro-

boscis or longer, last segment slender. Terminalia. Lobes of tergum IX with

long setae; basimere with relatively small conical basal mesal lobe bearing

bristles, mostly medium to small but also a few strong ones; subapical lobe

absent; distimere with single terminal appendage; aedeagus usually simple and

weakly sclerotized except in marchettei where it has strongly sclerotized lat-

eral walls and 3-4 prominent apical teeth.

PUPA. Tip of midrib of paddle with 2 hairs.

LARVA. Head, Head very large; antenna long and slender; hair 1-A,

a large tuft well beyond middle of antenna; 2-A and 3-A very long and usually

slightly removed from tip of antenna. Abdomen. Comb scales in a single row;

siphon long, index 4.0-7.5; basal hair 1-S small, single or with 2-5 branches;

ventral hairs la-S extending nearly to end of siphon; saddle forming complete

psa hair 2-X multibranched; precratal tufts of ventral brush 4-X, if present,

small. :

EGG. Eggs laid in rafts on water surface.

CULISETA (CLIMACURA) MARCHETTEI GARCIA, JEFFERY & RUDNICK

(Figures 5, 6, 7, 8)

Culiseta marchettei Garcia, Jeffery & Rudnick 1969, J. med. Ent. 6(3): 252

(#*,9, L*,P*); 1968, Med. J. Malaya 23(1): 29.

FEMALE. Head. (Figs. 5, 6) Vertex with decumbent pale scales; up-

right forked scales blackish; lateral scales broad, pale; palpus about 1/5 length

48 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

of proboscis, brown; without scales. Thorax. Anterior pronotum without

scales; posterior pronotum with fine black hair-like scales; 3-4 spiracular

bristles; scutum and scutellum with narrow dark scales; pleuron with a few

scales. Legs. (Fig. 6) Dark scaled; mid legs with pale scaling on posterior

surface of femur and with lighter scales along tibia and at base of tarsomeres

1-3; hind femur with pale scaled posterior surface; tibia with pale scaled apex,

tarsomeres 1-4 with basal pale bands, tarsomere 5 mostly white scaled; tarsal

claws as in Figure 5. Wing. (Fig. 5) Membrane of wing clear; scales on

veins uniformly dark; cross-veins without scales. Abdomen. Tergum I with-

out scales, II-VI brown scaled, III and IV with lateral basal patches of white

scales, V-VII with white basal bands, VII mostly pale scaled; sterna mostly

pale scaled. |

MALE. In general similar to female. Head. (Figs. 5, 6) Palpus

longer than proboscis, dark scaled with some pale scaling laterally on seg-

ments 2 and 3; distal part of segment 3 and segments 4 and 5 with long hairs;

segment 5 slender. Legs. Tarsal claws as in Figure 5. Terminalia. (Fig. 7)

Tergum VIII with some short, stout bristles at apical margin; each lobe of

tergum IX with 2-4 setae; basimere long and conical, without scales; basal

mesal lobe conical with 1-2 strong and some medium and small bristles; di-

stimere simple, about half as long as basimere, terminal appendage broad,

blunt, spoon-shaped; aedeagus more or less cylindrical, with lateral walls

strongly sclerotized.and with 3-4 prominent teeth; paraproct with 3-4 strong

teeth apically and 3-4 cercal setae. |

PUPA. (Fig. 7) Cephalothorax. Respiratory trumpets short with

opening almost to base. Hair 8-C single; 9-C 2-3b. Abdomen. Hair 1-I0

strong, 5-6b; 1-IV strong, 3-4b; 5-IV,V strong, 4-5b; 6-I-IV single or 2b,

6-V 2b, 6-V1 4b, 6-VII 3-4b. Paddle. Oval; posterior margin with minute

spicules; hair 1-P 2-3b; accessory hair single, short.

LARVA. (Fig. 8) Head. Head broader than long; antenna long, only

slightly shorter than the length of head, with sparse minute spiculation; hair

1-A inserted on distal 1/7 of shaft with 20-25 plumose branches, more than

1/2 length of shaft; hair 4-C small, fine, 2b, 5-C about as long as 4-C about

10b, 6-C long, single, 7-C 5-9b, 8-C about 7b, 9-C minute about 8b, 10-C

single, 11-C 4-6b, 13-C well developed 5-7b; all mouth brush hairs filiform;

mentum with 7 teeth on each side. Thorax. Hairs 1-3-P arising from sclero-

tized base, 1, 2-P long, single, 3-P 3b, 4-P 5-9b, 5,6-P single, 7-P 5-6b

plumose, 8-P 4-6b plumose, 9,10,12-P single; 5,6,7-M single, 8-M about 6b

plumose, 9-M about 10b plumose; 7-T 6-10b plumose, 9-T 7b plumose. AD-

domen. Hairs 6-I,II 4-6b plumose, 6-III-VI 3-5b; 1-VII 9-12b, 3-VIII 8-10b,

o- VII 12-16b, 2, 4-VIII single; comb with a single row of 13-20 spindle shaped,

finely fringed scales; siphon index 4.0-5.7, black on apical quarter; pecten

with 9-11 spines; hair 1-S inserted at base, small, 3-5b, la-S in single row

of 13-18 hairs (usually 6b) distal to pecten; abal segment completely ringed by

saddle; hair 1-X small 5-7b, 2-X 3-4b, 3-X 2b; ventral brush consisting of

16-18 tufts, 1 or 2 precratal; gills narrow, more than twice as long as saddle,

with constriction near base.

EGG. Elongate-oval, grey with posterior end narrower and darker;

deposited in raspberry- shaped clusters of up to 149 eggs.

TYPE DATA. Holotype male, allotype female and paratypes: 4 males

and 2 females with associated larval and pupal skins in U.S. National Museum.

Type locality: Tanjong Rabok, Kuala Langat Forest Reserve, Selangor, WEST

MALAYSIA.

DISTRIBUTION. Specimens examined: WEST MALAYSIA, Selangor,

Tanjong Rabok, Kuala Langat Forest Reserve 1 paratype male and 1 paratype

female, 3 larval and 3 pupal skins, 8 whole larvae. Records from literature:

Selangor, Batang Berjuntai; Pahang, highway to Pekan, 32 km. from Kuantan

(Garcia et al. 1968).

Dobrotworsky: Culiseia in Southeast Asia 49

TAXONOMIC DISCUSSION. The species of the subgenus Climacuva

from North America, Australia and New Zealand show a great degree of simi-

larity in the larval stages and in the terminalia of the male. In all these spe-

cies the aedeagus has a simple structure and weak sclerotization. However,

marchettei shows some specializations in the larva and the male genitalia.

The siphon in marchettei is more heavily pigmented apically and the aedeagus

is more sclerotized with prominent teéth apically.

BIOLOGY. Only adults have been collected, all of them in fresh-water

peat-swamp forests. Although no larvae have been found in nature they have

been successfully reared, from eggs, in the laboratory. Adults were obtained

when larvae were reared in covered pans in an air-conditioned room at a tem-

perature of 20- 25°C. (Garcia et al. 1968, 1969). It is probable that like some

Australian species of Culiseta, marchettei breeds in subterranean waters.

Adults have been recorded from traps baited with pig-tail monkey and

chicken. However, freshly engorged mosquitoes were recovered only from

the chicken-baited traps. Presumably this species prefers avian hosts.

ACKNOWLEDGEMENTS

I am grateful to Drs. Alan Stone and Botha de Meillon for the loan of

type material now in the U.S. National Museum and for many helpful sugges-

tions during the preparation of this paper; to Dr. F. H. Drummond, Depart-

ment of Zoology, University of Melbourne for assistance in the preparation of

the manuscript. The loan of collections of niveitaeniata from India by Dr. P.

F. Mattingly, Department of Entomology, British Museum (Natural History)

and lishanensis from Taiwan by Capt. R. H. Watten, Department of U. S.

Naval Medical Research Unit No. 2 is acknowledged with sincere appreciation.

The illustrations prepared by Miss T. L. Ford of the Southeast Asia Mosquito

Project and artists of the U.S. Army 406th General Medical Laboratory, Tokyo,

were also much appreciated.

LITERATURE CITED

BARRAUD, P. J.

1934. The fauna of British India, including Ceylon and Burma. Diptera

V. Family Culicidae, tribes Megarhinini and Culicini. Taylor

and Francis, London. 463 pp. 7

BELKIN, J. N.

1962. The mosquitoes of the South Pacific. Univ. Calif. Press,

Berkeley 1: 282.

BLANCHARD, R.

1905. Les Moustiques. Paris, de Rudeval. 673 pp.

BRUNDIN, L.

1966. Transantarctic relationships and their significance as evidenced

by chironomid midges with a monograph of the subfamilies

Podonominae and Aphroteniinae and the austral Heptagyiae.

K. svenska Vetensk. Akad. Handl. 11(1):1-472.

1967. Insects and the problem of austral disjunctive distribution.

Ann. Rev. Ent. 12: 149-168.

50 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

CHAMBERLAIN, R. W., RUBIN, H., KISSLING, R. E. & M. E. EIDSON

1951. Recovery of virus of eastern equine encephalomyelitis from a

Sea Culiseta melanura (Coquillett). Proc. Soc. exp. Biol.

7: 396-397.

CHAMBERLAIN, R. W., SIKES, R. K., NELSON, D. B. & W. D. SUDIA

1954. Studies on the North American arthropod- borne encephalitides.

VI. Quantitative determinations of virus-vector relationships.

Amer. J. Hyg. 60: 278-285.

DANIELOVA, V., MINAR, J. & J. RYBA

1970. Isolation of Tahyna virus from mosquitoes Culiseta annulata

(Schr'k) 1776. Folia parasitol. 17(3): 281-284.

DARLINGTON, P. J.

1970. A practical criticism of Hennig- Brundin ''Phylogenetic system-

atics'' and Antarctic biogeography. Syst. Zool. 19: 1-18.

DOBROTWORSKY, N. V.

1954. The genus Theobaldia (Diptera, Culicidae) in Victoria. Proc.

Linn. Soc. N.S.W. 79: 65-78.

1958. Designations of the type species of the subgenus Neotheobaldia

Dobrotworsky (Genus Theobaldia Neveu- Lemaire 1902). Proc.

ent. Soc. Wash. 60: 186.

1965. The mosquitoes of Victoria (Diptera, Culicidae). Melb. Univ.

Press, Melbourne. 237 pp.

EDWARDS, F. W.

1921. A revision of the mosquitoes of the Palaearctic region. Bull.

ent. Res. 12: 263-351.

1930. Mosquito Notes. IX. Bull. ent. Res. 21: 287-306.

1932. in Wytsman, Genera Insectorum. Diptera. Family Culicidae.

Fasc. 194, Desmet-Verteneuil, Brussels. 258 pp.

FELT, E. P.

1904. Mosquitoes or Culicidae of New York State. Bull. N.Y. St. Mus.

79: 391c.

FISCHER, P.

1885. Manuel de Conchyliologie et de RG ane oat Conchyliologique.

p. 744.

GARCIA, R., JEFFERY, J. & A. Rudnick

1968. "A first report of the genus Culiseta in Malaysia. Med. J.

Malaya 23(1): 29.

1969. Culiseta marchettei, a new species of the subgenus Climacura

Howard, Dyar and Knab from Malaysia with notes on its biology

(Diptera: Culicidae). J. med. Ent. 6(3): 251-256.

HAMMON, W. McD. & W. C. REEVES

1943a. Laboratory transmission of western equine encephalomyelitis

virus by mosquitoes of the genera Culex and Cae J. eXp.

Med. 78: 425-434.

Dobrotworsky: Culiseta in Southeast Asia 51

HAMMON, W. McD. & W. C. REEVES

1943b. Laboratory transmission of St. Louis encephalitis virus by

three genera of mosquitoes. J. exp. Med. 78: 241-253.

HAMMON, W. McD., REEVES, W. C., BRENNER, S. N. & B. BROOKMAN

1945. Human encephalitis in the Yakima Valley, Washington, 1942,

with forty-nine virus isolations (Western equine and St. Louis)

from mosquitoes. J. Amer. med. Assoc. 128: 1133-1139.

HAYES, R. O., DANIELS, J. B. & R. A. MACCREADY

1961. Western encephalitis virus in Massachusetts. Proc. Soc. exp.

Biol., N.Y. 108(3): 805-808.

HAYES, R. O., BEADLE, L. D., HESS, A. D., SUSSMAN, O. & M. J.

BONESE

1962. Entomological aspects of the 1959 outbreak of eastern encephali-

tis in New Jersey. Amer. J. trop. Med. 11(1): 115-121.

HOLDEN, P., MILLER, B. J. & D. M. JOBBINS

1954. Isolation of eastern equine encephalomyelitis virus from mosqui-

toes (Culiseta melanura) collected in New Jersey 1953. Proc.

Soc. exp. Biol., N.Y. 87(2): 457-459.

HOLDEN, P. & A. D. HESS

1959. Cache Valley virus, a previously undescribed mosquito- borne

agent. Science 130(3383): 1187-1188.

HOWARD, L. O., DYAR, H. G. & F. KNAB

1915. The mosquitoes of North and Central America and the West Indies.

vol. 3, 523 pp. Washington, D.C.

HURLBUT, H.S.

1956. West Nile virus infection in arthropods. Amer. J. trop. Med.

5(1): 76-85.

KISSLING, R. E., CHAMBERLAIN, R. W., NELSON, D. B. & D. D. STAMM

1955. Studies on the North American arthropod-borne encephalitides.

ee Equine encephalitis studies in Louisiana. Amer. J. Hyg.

62: 233-254.

LAMOTTE, L. C.

1968. In Garcia, R., Jeffery, J. & A. Rudnick.

LIEN, J. C.

1968. New species of mosquitoes from Taiwan (Diptera: Culicidae).

Part Il. New species of Tripteroides, Orthopodomyia, Culiseta

and Uvanotaenia. Trop. Med. 10(1): 1-20.

LIU, C. W. &L. C. FENG

1956. On a species of Theobaldia, Theobaldia kanayamensis newly dis-

covered from Peking. Acta ent. sinica 6(3): 335-341.

MARKS, E. N.

1968. Northern records of the genus Culisefa Felt in Australia, with

the description of a new species (Diptera, Culicidae). J. Aust.

ent. Soc. 7: 43-56.

52 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

MASLOV, A. V. |

1964. The taxonomy of bloodsucking mosquitoes of the Culiseia group

(Diptera, Culicidae). Rev. Ent. U.R.S.S. 43(1): 193-217.

1967. Blood sucking mosquitoes of the subtribe Culisetina (Diptera,

ie of the world fauna. Akad. Nauk S.S.S.R., Opred.

93: 1-182.

MACKERRAS, I. M.

1950. The zoogeography of the Diptera. Aust. J. Sci. 12(5): 157-161.

MENG, C. H. & C. Y. WU |

1962. A new species of Theobaldia (Diptera, Culicidae). Theobaldia

(Theobaldia) sinensis, sp. nov. Acta ent. sinica 11(4): 382-387.

NEVEU- LEMAIRE, M.

1902. Sur la classification des Culicides. C.R. Soc. Biol. Paris

54: 1329-1332.

REEVES, W. C.

1968. In Garcia, R., Jeffery, J. & A. Rudnick.

ROSS, R. W., AUSTIN, F. J., MILES, J. A. R. & T. MAGUIRE

1963. An arbovirus isolated in New Zealand. Aust. J. Sci. 26(1): 20-21.

SPALATIN, J.,; BARTON, A. N., McLINTOCK, J. & R. CONNELL

1963. Isolation of a western equine encephalitis (WEE) virus from mos-

age in Saskatchewan, 1962. Canad. J. comp. Med. 27(12):

83-289.

STONE, A.

1957. Corrections in the taxonomy and nomenclature of mosquitoes

(Diptera: Culicidae). Proc. ent. Soc. Wash. 58: 333-344.

STONE, A., KNIGHT, K. L. & H. STARCKE

1959. A synoptic catalog of the mosquitoes of the world (Diptera:

Culicidae). Ent. Soc. Amer. (The Thomas Say Found.) vol. 6:

1-358.

SURTEES, G.

| 1959. Functional and morphological adaptations of the larval mouth

parts in the subfamily Culicinae (Diptera) with a review of some

a studies by Montchadsky. Proc. R. ent. Soc. Lond. 34:

THEOBALD, F. V. :

1907. A monograph of the Culicidae or mosquitoes. IV. London. 639 pp.

WHITNEY, E.

1964. Flanders strain, an arbovirus newly isolated from mosquitoes

and birds of New York State. Amer. J. trop. Med. 13(1): 123-

131.

fore claw o7

m™

mid claw o7

yy 0.1mm

\ = |

hind claw o&%

mid claw 9

| Sr

/ 2 hind claw Q

C. (Culiseta) niveitaeniata

= =

anter

/taeniata

(Culiseta) nive

~~ Tac

“iNaul b

elses Cn ee ee

aedeagus

rei apex

sternal

1.0mm

LE Se RTT

mi! AA

a A

i \)

veal in .! a (i a

h We \ee Nigh +I NY

I-T iy

i fu

hy vy iG. | | \ Ba

ae | t

NEVA \ |

a aN TNS ee Re ae

fie

‘faenlata

/ve/

C. (Culiseta) n

Fig. 5

fore claw o7

mid claw o7

hind claw of

fore claw 9

| ee.

this \

AT\

Why a

af wy

C. (Climacura) marchettei Sy

Fong

“tea

\S)

ble Sly i

(Di ye pe gvet Ant

' Ari V Lech ot

IN} 7A rsa

Ava)

1) Mate

AON |

vt ‘ : f ( YAN PID 6

UA (Fight lias (hd WAAR es

SE SN ee q Nery Ae 0 ee I nS |

C. (Climacura) marchette/

C. (Climacura) marchettei SATS TE nea ;

Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971 61

INDEX

Valid names are printed in roman type, synonyms are italicized. Ital-

icized page numbers are those which begin the primary treatment of that spe-

cies. Numbers in parenthesis refer to the figures illustrating some portion of

that species.

Allotheobaldia

annulata

Anopheles stigmaticus

Austrotheobaldia

bergrothi

Climacura

Culex absobrinus

Culex annulatus

Culex dyari

Culex melanurus

Culicella

Culiseta

Culiseta lishanensis

Dirofilaria immitis

Engaeus

Finlaya

fraseri

incidens

inornata

Leptosomatomyia fraseri

longiareolata

marchettei

melanura

Neotheobaldia

niveitaeniata

Pseudotheobaldia

rubrithorax

Theobaldia

Theobaldia kanayamensis

Theobaldia sinensis

Theobaldinella

Theomyia

tonnoiri’

39, 41,

44, 47, 49

44.

41, 43, 44, 49

49 (5, 6, 7, 8)

45, 46, AT (1, :2..3, 4)

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. XI.

CONTENTS

ABSTRACT... ... ewe eee ee a ee ee 62

INTRODUCTION AND HISTORY OF PREVIOUS CLASSIFICATIONS ... 63 |

A CONSIDERATION OF THE PROPOSED RECLASSI FICATION oe fae 66

OUTLINE OF THE PROPOSED SCHEME OF RECLASSIFICATION ... 67

KEY TO THE SUBGENERA 2.4 665 20S es > ous 6 Ge as a Oe

| Subcenus 1. NEOCULEX Dyar 2.5 cise ss ie en 0 ew eee 8c 69

Key to the groups of subgenus Neoculex .. 2... eee sees 69

Subgenus II. MAILLOTIA (Theobald). ......-.2-+-eeees 71

Key to the groups of subgenus Maillotia .......2...ee-s 71

_ Subgenus Il. EUMELANOMYIA Theobald ... 2.0502 s secs 72

Key to the groups of subgenus Eumelanomyia ......+.-++-s 73

Key to the subgroups of group mochthogenes ......+.- 74

POCRNOW DEDGEMENTS:. 6 kw ee we eS oe es ein ce, ee 15

MOPERONGbe COED 0a ¥ coccij we 8 ee oes Se 76

PIGURES So ee ee ee ek Sk ek ae Spee eee ee 80

TD a a es 8 ae 83

ABSTRACT

A reclassification of Neoculex proposed here is based on the

comparative morphology of the male terminalia as well as on other corre-

lated external characters of the adults. In this scheme, three subgenera:

Neoculex, Maillotia and Eumelanomyia instead of two, Neoculex and Moch-

thogenes, as proposed by Edwards (1932), are recognized. The subgenera

Maillotia and Eumelanomyia are resurrected from synonymy with Neoculex

and Mochthogenes which Edwards (1930) treated as a full subgenus is sup-

pressed by synonymizing it with Eumelanomyia. The species tricuspis

Edwards 1930 is transferred to Culiciomyia and sumatranus Brug 1931 and

caeruleus King & Hoogstraal 1947 to Lophoceraomyia. }

Over 80 species previously assigned to Neoculex and Moch- _

thogenes in the synoptic catalog of Stone et al. (1959) and Stone (1961, 1963,

1967, and 1970) are placed in various species groups of the three subgenera.

Keys to the subgenera, groups and subgroups are provided and each category

is briefly defined with regard to systematics and zoogeography.

A PROPOSED RECLASSIFICATION OF NEOCULEX DYAR

BASED PRINCIPALLY ON THE MALE TERMINALIA4

Sunthorn Sirivanakarn2

INTRODUCTION AND HISTORY OF PREVIOUS CLASSIFICATIONS

The subgenus Neoculex Dyar 1905 in the broad sense of Edwards (1932,

1941) has remained perhaps one of the most poorly understood subgenera of

Culex. The subgenus, as defined by Edwards, with the material at his dispos-

al, is a heterogeneous array of several distinct lineages of 70 or more species

known at the present time. These species are mostly restricted to certain

zoogeographical regions. The one exception is territans (Walker, 1856) from

North America which is also known to occur in some European countries. Re-

cords of species from different areas are as follows: 33 from the Ethiopian

region, 8 from the Mediterranean subregion, 9 from the Oriental region, 9

from the Australasian region (Australia and New Guinea), 5 from the South

Pacific and 6 from the Nearctic region. |

No attempt has yet been made to revise Neoculex on a world basis, but

there have been a good number of taxonomic papers dealing with local species

in several regional works (see references below). Edwards' subgeneric inter-

pretation and his internal classification have been largely followed by a few

critical comments. Mattingly & Marks (1955) and Belkin (1962) pointed out the

weaknesses regarding the relationships between Edwards' species groups, but

these were limited to brief statements. A critical examination of Edwards'

scheme is made here to set a stage for the further development of a phyloge-

netic classification. The main purpose of the present attempt is to lay out

certain basic and significant features not used by Edwards in his interpretation

of Neoculex. The basis of this discussion includes the study of his work and

the re-examination of almost all species which he used in devising his scheme.

Edwards' interpretation of Neoculex was based on many superficial

characters which greatly overlap with those of other subgenera, particularly

Mochthogenes, Lophoceraomyia, Culiciomyia and to some extent even with

those of other Culex subgenera. He apparently defined all species involved on

the basis of the simple phallosome of the male terminalia which he did not de-

scribe in detail. This has resulted in some incorrect subgeneric assignments

of certain species to the subgenus. Edwards’ description of the male phallo-

some is brief and also appears to conceal a number of significant features with

regard to its varied shape and the relative position of the tergal bridge which

connects the two lateral plates. This point will be considered and illustrated

below in my interpretation of various species groups. It suffices to mention

that the shape of the phallosome is quite constant in certain lineages and ap-

pears to be strongly differentiated. The other characters which appear to be

more or less consistently correlated with the differences in shape of the phallo-

some, but were not considered by Edwards are: texture of the spicules of the

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

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

of the Surgeon General, Washington, D.C. 20314.

2 Southeast Asia Mosquito Project, Department of Entomology, Smithsonian

Institution, Washington, D.C. 20560.

64 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

proctiger crown, number of rodlike setae in the proximal division of the sub-

apical lobe and the presence or absence of scale patches on the pleura. A brief

summary of Edwards’ scheme is as follows:

In grouping species in the subgenus Neoculex, Edwards (1932) suppres-

sed 3 genera of Theobald (1907, 1910) including Maillotia, Eumelanomyia and

Protomelanoconion by reducing them to synonymy with the genus Neoculex Dyar

(1905). The genus Neoculex, as originally conceived by Dyar, was based on C,

territans (Walker 1856) from North America, a form which shows a great deal

of superficial resemblance to some members of the pipiens group of subgenus

Culex. In clarifying his classification, Edwards separated 3 groups in the sub-

genus, namely: group A. Neoculex s. str. or apicalis group, group B. Eu-

melanomyia or albiventris group and group C. Protomelanoconion or uniformis

group. These groups are distinguished by the relative length of male palpi,

texture of decumbent scales on vertex and presence or absence of apical bands

on abdominal terga. Later, in his work on the Ethiopian species, he (1941)

split group A. into 3 groups by incorporating features of the female buccopha-

ryngeal armature, color pattern of pleural integument and certain conspicuous

ornamentation. This subdivision resulted in: group A. pulchrithovax, group B.

Neoculex s. sty. and group C. vima group. The original groups B. and C. be-

came groups D. and E. The first and second treatments are essentially simi-

lar in outline and scope and in 1947, King & Hoogstraal followed this scheme

by recognizing another additional group F. pedicellus from New Guinea. A

critical study of Edwards' system indicates that he lumped a number of unre-

lated forms in Neoculex s. sty. group and his group characters largely over-

lapped and remained confused. The relationships between these groups are not

at all clear and the entire treatment appears to suffer from the lack of unifor-

mity in most parts. A broader examination of his system reveals the difficul-

ties he experienced in ranking certain species groups. This is quite obvious

from his treatment of Mochthogenes as a subgenus separated from the Proto-

melanoconion or uniformis group of Neoculex based on the relative length of

the male palpi which are as short as in the female of all Mochthogenes species

but are longer in the Protomelanoconion species. Although such treatment is

convenient in practice since species of Mochthogenes share these characters,

the relationship with Protomelanoconion on the basis of the male terminalia

and many other characters do not seem to warrant its separation. The two

groups, as recently pointed out by Bram (1969), are also so similar in the lar-

val stage to be treated in the same subgenus.

In my current study of Neoculex in Southeast Asia and other adjacent

areas, the difficulty presented by Edwards' classification are illustrated by

the following examples. Four species, namely tenuipalpis Barraud 1924,

hayashii Yamada 1917, hackevi Edwards 1923 and Rkiriensis Klein & Sirivanakarn

1969, all with extremely similar male terminalia and several external charac-

ters, but with palpi of different lengths, will, according to Edwards, have to

be placed in either Neoculex or Mochthogenes. Similarly tricuspis Edwards

1930, at present in Neoculex, should in fact be reassigned to Culiciomyia;

sumatranus Brug 1931 and caeruleus King & Hoogstraal 1947 on the other hand

rightly belong to Lophoceraomyia.

In developing the following scheme of reclassification, I have been for-

tunate in having the opportunity to re-examine several authenticated specimens,

including the types and identified specimens of species used by Edwards in his

revision of the world fauna, by Mattingly (1953) in his work on the Mediterra-

nean species and other type specimens from Several areas, at the British

Museum. In addition, while undertaking the revision of the Indomalayan spe-

cies at Southeast Asia Mosquito Project, I have also seen specimens of the

North American species and others from several areas in the reference collec-

tions of the United States National Museum.

The history of the classification of Neoculex and Mochthogenes is sum-

marized in table I below.

— 65

Sirivanakarn: Reclassification of Neoculex

sidjpdinuaj ‘14049040

eos ‘s~suaAjsva 1h] DU

‘snsonaidsuoour ‘snzpurdun

‘szsuasunjsury :sdnoisqns)

dnois SaUuasoyjyIOU °F

dno13 wozuor0unj awojord °¢

(oUtA *snjourgna :Sdnoisqns)

Gnois DWIA-SNJOULQNA °Z

dno1is wdwmounjawmna °T

mtmounaung snuesqng

dnois 2d140aS °¢

Gno1s Szsuajsoy °Z

dnoi3 xnsoypAryoindg *T

nLjOTILDWY Snuasqng

dno1z3 snzk7sissvrd °¢

dnois muor0unjawmopnasd °Z

GnNOId SUDJIAAA] *T

xajnooan snhuesqng

NOILVOMISSVTIO LNaSddd

sauasoylyIOW snuesqng

(LPG, Teetyss00H 7% Suty)

Gnois snjjaI1pad °q

dnois szwmsofiun 10

UOLUDIOUDNLBWOJOAT °F

dno1s s21juaarq]y IO

MIKWUMOUDIAWNA °C

dnois DWu21y °D

dnois s27097dp IO

"AJS *S XaINIOAN °A

ANOS XDAOYJAYIINT °VW

xaynd0aN snuesqng

(L¥61) IVVULSDOOH F ONDI

(TP61T) SAUVMCH

‘squasoyly20w snuesqns

dnois s2zmsofiun 10

UOLUOIOUNLIWOJOAd °D

dnois $247Uua0a7q]y0 IO

DINMOUNIWNA °|

dnois s27091dpv IO

°IIS *S xa7ndI0aN “VW

xajno0an snuesqns

(ze61) SGUVMAH

ee ee perc pf Sn Sp oS ee

SANADOHLHOOW GNV XHTNOOAN AO NOILVOLMIISSVIO FHL AO AMOLSIA “1 a IaVL

66 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

A CONSIDERATION OF THE PROPOSED RECLASSI FICATION

The adults of all species, previously assigned to Neoculex and

Mochthogenes and to be considered at present are generally distinguished

from other Culex subgenera as follows: From Lophoceraomyia by the absence

of scale tufts on the male antennal flagellum; from Culiciomyia by the absence

of a row of lanceolate scales on the ventral surface of segment 3 of the male

palpus; from Acalleomyia by the narrow scutellar scales; from Barraudius

and Lasiosiphon by having tarsomere 1 of the hind tarsus about as long as the

tibia and by the absence of scales on the basimere of the male terminalia; from

Culex and Lutzia by the absence of basal sternal processes on the proctiger of

the male terminalia ; and from all New World subgenera including Melanoconion,

Aedinus, Isostomyia, Carrollia, Mochlostyvax, Microculex,and Micraedes by

the absence of a basal hook on the lateral plate of the male phallosome.

Taxonomic Characters. As indicated earlier in the introduction, basis

to the present interpretation of various species groups and their reclassifica-

tion is the comparative morphology of the phallosome and other correlated

characters in the proctiger crown and subapical lobe of the male terminalia.

The shape of the phallosome is not only of taxonomic significance in this inter-

pretation, but also provides, in addition to other conspicuous characters, a

clear-cut separation of all species involved from other closely related subgen-

era of Culex. It appears that there are at least 3 basic types of male phallo-

some present among species of Neoculex as interpreted by Edwards. These

are: Type I tubular, elongate with the axis of the two lateral plates more or

less parallel and with the tergal bridge located above the middle or near the

apex of the lateral plate; Type II more or less globular or subspherical with

tergal bridge as in Ior at the middle of the lateral plate; Type III slightly

modified from type II in being oval with tergal bridge at or slightly below the

midpoint of the lateral plate. In the proctiger crown, there are two kinds of

spicules; one of these is flat and blunt, the other fine and pointed. In certain

lineages, the crown may consist of flat or fine spicules only, or a mixture of

both. In the subapical lobe, there is a great deal of variation in the develop-

ment of parts and the number of specialized setae, however, the features

which appear to be correlated with different types of phallosome are the pres-

ence of 2 or 3 rodlike setae in the proximal division and the presence or ab-

sence of a leaflet (foliform seta) in the distal division. |

In the external morphology, a number of conspicuous characters which

are of practical value in separating species groups at various levels are: (1)

resence or absence of pleural scaling, (2) color and texture of scutal scales,

3) relative length of male palpus and (4) presence or absence of pale abdomi-

nal banding. These and a few other features may be correlated with the ter-

minalia to a certain extent and have been considered here in developing key

and group characters.

The female buccopharyngeal armature and the immature stages may

also prove to be useful in developing this classification but because of insuffi-

cient material, no attempt has been made here to incorporate them with the

present scheme. | |

Systematics. In the present interpretation, I believe it would be much

sounder, considering both the comparative male terminalia and, to Some ex-

tent, zoogeography to recognize at least 3 principal subgeneric categories

among 80 or more species known at the present time. In the general outline

presented below, I recognize Neoculex, Maillotia and Eumelanomyia as dis-

tinct subgenera based on differences in the shape of the male phallosome as

discussed above. The subgenera Maillotia and Eumelanomyia are resurrected

from synonymy under Neoculex and the subgenus Mochthogenes is downgraded

to a species group of Eumelanomyia.

These three subgenera are further subdivided into groups and subgroups

wherever it is appropriate to accomodate all species presently listed in

Sirivanakarn: Reclassification of Neoculex 67

Neoculex and Mochthogenes in Stone et al. (1959) and Stone (1961, 1963, 1967

and 1970), except caeruleus King & Hoogstraal 1947; sumatranus Brug 1931

and tricuspis Edwards 1930. The first two ofthese I am transferring to

Lophoceraomyia and tricuspis to Culiciomyia. The assignment of some spe-

cies to groups below the subgeneric level has been based only on the published

descriptions and may need future realignment in order to indicate a more ac-

curate affinity.

OUTLINE OF THE PROPOSED SCHEME OF RECLASSIFICATION

Subgenus I.

(1).

(2).

(3).

NEOCULEX Dyar 1905.

territans group with territans (Walker 1856); apicalis Adam 1903;

boharti Brookman & Reeves 1950; vreevesi Wirth 1948; avrizonensis

Bohart 1950; derivator Dyar & Knab 1906; deserticola Kirkpatrick

1924; judaicus Edwards 1926; impudicus Ficalbi 1889; rubensis

Sasa & Takahashi 1948 and martinii Medschid 1930.

pseudomelanoconia group with pseudomelanoconia (Theobald

1907); postspiraculosus Lee 1944; chaetoventralis (Theobald 1910);

douglasi Dobrotworsky 1956; latus Dobrotworsky 1956; fergusoni

(Taylor 1914); cheesmanae Mattingly & Marks 1955; dumbletoni

Belkin 1962; gaufini Belkin 1962 and millironi Belkin 1962.

crassistylus group with crassistylus Brug 1934; pedicellus King &

Hoogstraal 1947 and leonardi Belkin 1962.

Subgenus II. MAILLOTIA (Theobald 1907).

(1).

(2).

(3).

pulchrithorax group with pulchrithorax Edwards 1914.

hortensis group with hortensis Ficalbi 1889; arbieeni Salem 1938

and quettensis Mattingly 1955.

seyrigi group with seyrigi Edwards 1941; peringueyi Edwards 1924;

salisburensis Theobald 1901 and avianus de Meillon 1943.

Subgenus III]. EUMELANOMYIA (Theobald 1909).

(1).

(2).

eumelanomyia group with albiventris Edwards 1922; andersianus

Edwards 1941; acrostichalis Edwards 1941; vinckei Hamon,

Holstein & Rivola 1961(1962); kanyamwerima Someren 1951; kilara

Someren 1951; gavioui Bailly-Choumara & Rickenbach 1966.

rubinotus -vima group

(a). vubinotus subgroup with rubinotus Theobald 1901;

kingianus Edwards 1927; andreanus Edwards 1927;

pseudoandreanus Bailly-Choumara 1965 and simplici-

cornis Edwards 1930.

(b). vima subgroup with vima Theobald 1901; subrima

Edwards 1941; galliardi Edwards 1941; calabarensis

Edwards 1941; wigglesworthi Edwards 1941; insignis

(Carter 1911); sunyaniensis Edwards 1941; albertianus

Edwards 1941; wansoni Worlfs 1945; adamiHamon &

Mouchet 1955; Japlantei Hamon, Adam & Mouchet 1955;

Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

amaniensis Someren & Hamon 1964 and chauveti Brunhes

& Rambelo 1968.

(3). protomelanoconion group with brevipalpis (Giles 1902); stellatus

Someren 1947 and horridus Edwards 1922.

(4). mochthogenes group

_ (a). hinglungensis subgroup with hinglungensis Chu 1957;

culionicus Delfinado 1966; tricontus Delfinado 1966 and

cataractarum Edwards 1923.

_(b). uncinatus subgroup with uncinatus Delfinado 1966.

(c). inconspicuosus subgroup with inconspicuosus (Theobald

1908); simpliciforceps Edwards 1935; castor de Meillon

& Lavoipierre 1944; hamoni Brunhes et al. 1967;

mijanae Brunhes et al. 1967 and orstom Brunhes et al.

1967; perhaps also bokorensis Klein & Sirivanakarn 1969.

_(d). malayi subgroup with malayi (Leicester 1908); laureli

Baisas 1935 and yeageri Baisas 1935.

_(e). castrensis subgroup with castrensis Edwards 1922;

foliatus Brug 1932; latifoliatus Delfinado 1966; chiyutoi

ope ea 1935 and shrivastavii Wattal, Kalra & Krishnan

966

(f). femineus subgroup with femineus Edwards 1926.

(g). ogi subgroup with ofachati Klein & Sirivanakarn

69.

(h). tenuipalpis subgroup with tenuipalpis Barraud 1924;

hayashii Yamada 1917; hackeri Edwards 1923; pluvialis

Barraud 1924; kiriensis Klein & Sirivanakarn 1969; selai

Klein & Sirivanakarn 1969; campilunati Carter &

Wijesundara 1948; and perhaps also okinawae Bohart

1953; lini Lien 1968: Rkhazani Edwards 1922; and iphis

Barraud 1924,

KEY TO THE SUBGENERA

1. Phallosome elongate, more or less uniformly

cylindrical, H-shaped in tergal view with tergal

bridge above the midpoint or near apex of lat-

eral plate; proctiger with crown of flat and blunt

spicules; pleuron usually with broad scale patches

on propleuron, upper corner and lower border of

sternopleuron, anterior upper mesepimeron and

occasionally also on postspiracular area... Subgenus NEOCULEX

Phallosome short, stout, oval or subspherical

in shape with tergal bridge at or below the

midpoint of lateral plates; proctiger with

crown of flat and blunt spicules or fine pointed

Spines; pleural scaling present as above or

BDSCHL Bei ME nha eee a ar Beh apily Hs 2

Sirivanakarn: Reclassification of Neoculex 69

2(1). Proctiger with crown of flat and blunt spicules

entirely or with some coarse pointed spines

in addition; pleural scaling present; scutal

scales usually pale or sand-colored...... Subgenus MAILLOTIA

Proctiger usually with a crown of fine pointed

spines only, or sometimes with a few

coarse ones in addition; pleural scaling

entirely absent; scutal scales predominantly

GOV DP OWN, 2. wicae tncder nee) GUBUNG Bil ae oes Subgenus KUMELANOMYIA

Subgenus I. NEOCULEX Dyar

1905 Neoculex Dyar, Proc. ent. Soc. Wash. 7:45; Type species Culex

territans Walker 1856, original designation.

1932 Culex (Neoculex) in part of Edwards, Gen. Insect. Dipt. Fam. Culicidae,

Fasc. 194:193-195.

Culex (Neoculex) in part of Edwards (1941: 249-270); King & Hoogstraal

(1947: 65-69); Mattingly & Marks (1955: 166-175); Dobrotworsky (1956:

105-114); Belkin (1963: 238-247); and Dobrotworsky (1965: 193-202).

Subgeneric Characters. As diagnosed in the key to subgenera, with the

following additional features. Medium to large sized species, wing length over

3.0 mm. Head. Male palpi as long as or longer than proboscis; antenna

strongly plumose. Thorax. Scutal scales usually predominantly pale, some-

times with striking pattern of coloration or entirely dark; pleuron usually with

extensive broad scale patches on propleuron, upper corner and posterior lower

border of sternopleuron and anterior upper mesepimeron, sometimes also on

postspiracular area and prealar area; rarely absent entirely. Abdomen. Ter-

ga with or without apical or basal bands, sometimes with apicolateral pale

spots. Male Terminalia. (Fig. 1) Phallosome elongate, tubelike, H-shaped

in tergal view or slightly modified, tergal bridge usually near to or almost at

the apex of lateral plate, rarely at the middle, a few denticles present or ab-

sent; proctiger with crown of flat and blunt spicules only or also with a few

coarse pointed spines in addition; subapical lobe always with 2 long rodlike

setae in the proximal division; distal division with only narrow flattened setae,

broad leaflets absent.

Systematics. Species which are strictly or provisionally placed within

this subgenus are generally similar in the configuration of the phallosome,

proctiger and features of the subapical lobe as described above. They may be

well divided into 3 groups on the basis of slight differences in phallosome

structure, presence or absence of pleural scaling as in the following key.

KEY TO THE GROUPS OF SUBGENUS NEOCULEX

1. Phallosome uniformly tubular in shape with

tergal bridge between the midpoint and apex

of lateral plates; pleural scaling always

present; abdominal terga with apical pale

bands; scutal scales sand-colored......... territans group

Phallosome broad in apical half, narrow in

basal half; tergal bridge nearly at or below

the apex of lateral plates; pleural scaling

present or absent; abdominal terga with

apical or basal pale bands, apicolateral

spots or sometimes entirely dark; scutal

scales usually entirely dark or sometimes

partly golden brown...... Bg Rat Gar a Me ae Pe ater a” 2

70 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

2(1). Tergal bridge nearly at the apex of lateral

plates; distimere usually slender, sickle

shaped; proximal and distal divisions of

subapical lobe usually clearly separated

but not elongated; pleural scaling usually 3

DECC kk ee ee ee — pseudomelanoconia group

Tergal bridge at or just above the midpoin 7

of lateral plates; distimere modified from

above; proximal and distal divisions of

subapical lobe elongated into stemlike

lobes; pleural scaling entirely absent or

sometimes only a few scales present on

SEOTNOUICHTOR 4 eso es we, edo. ooo: VASSIStVIUS group

TERRITANS GROUP

This group includes 6 species from the Nearctic (mainly North America);

territans (also known in Europe), apicalis, boharti, reevesi, arizonensis and

derivator; 1 species from northern Palearctic of the Oriental region: rubensis

and 4 species from the Mediterranean, namely deserticola, judaicus,impudicus

and martinii. The group can be easily recognized by the predominantly pale

or sand-colored scutal scales, presence of broad scale patches on 3 or 4areas

pleura and presence of apical banding on abdominal terga, as indicated in

the key. ,

PSEUDOMELANOCONIA GROUP

This group contains 6 species from Australia: pseudomelanoconia,

chaetoventralis, douglasi, fergusoni, latus and postspivaculosus and 4 species

from the South Pacific: cheesmanae, dumbletoni, gaufiniand millironi. The

extent of the pleural scaling is variable, but the shape of the phallosome and

the characteristic crown of the proctiger are very constant in nearly all spe-

cies involved. According to Dobrotworsky (1956: 105) the members of this

group could be well divided into two subgroups, one involving fergusoni and

latus with apical abdominal banding or apicolateral abdominal spots and pre-

sence of pleural scaling, the other involving douglasi and pseudomelanoconia

with basal abdominal banding and reduced pleural scaling.

CRASSISTYLUS GROUP

Two members of this group: crassistylus and pedicellus are known

from New Guinea and the other one, leonardi, is from the South Pacific. They

are strikingly differentiated from the other two groups in the almost complete

absence of pleural scaling, modified shape of distimere, development of parts

of the subapical lobe and in having striations on the upper tergal surface of the

lateral plate of the phallosome. They are probably derived from members in

the pseudomelanoconia group.

Sirivanakarn: Reclassification of Neoculex 71

Subgenus II. MAILLOTIA (Theobald)

1907 Maillotia Theobald, Mon. Cul. 4:274; Haplotype: pilifeva (presently known

as hortensis).

Culex (Neoculex) in part of Edwards (1932: 193); Edwards (1941: 249);

Mattingly (1955: 376-389).

Subgeneric Characters. As given in the key to the subgenera with the

following additional description. Very similar in general external features to

the territans group of subgenus Neoculex, but pleural scaling is more exten-

sive, sometimes scale patch also present on prosternum and scutal scales

rather coarse. Male Terminalia. (Fig. 2) Phallosome short, broad, oval,

subspherical or cup-shaped from tergal view, tergal bridge at or just above

the midpoint of lateral plates, denticles not developed or sometimes only afew

~ ones present on apex; proctiger heavily sclerotized with a relatively large

crown of flat and blunt spicules arranged in comblike fashion or sometimes

mixed with coarse pointed spines in the form of a tuft; proximal division of

subapical lobe with 2 or sometimes 3 rodlike setae, distal division with few

short narrow setae or none; distimere more or less modified.

Systematics. This is perhaps the most primitive of the three subgen-

era. It is rather heterogeneous consisting of species which are perhaps better

placed with either Neoculex or Eumelanomyia. However, as they show the

type of phallosome, and other features, somewhat intermediate between the

other two subgenera, I think it is probably better to consider them as belong~-

ing to a subgenus of their own. As they are either exclusively Ethiopian or

Mediterranean, it seems better to treat them this way.

KEY TO THE GROUPS OF SUBGENUS MAILLOTIA

1. Head, scutum and pleuron with a pattern of

silvery white scale lines contrasting sharply

with dark scaled background; apex of proc-

tiger of male terminalia with a heavily

sclerotized plate bearing a small crown of

COArsespicules: 2 6h CE ALE a ROARS pulchrithorax group

Head, scutum and pleuron without above

ornamentation; apex of proctiger of male

terminalia without heavily sclerotized

' plate, but with crown of flat and blunt

spicules or of coarse pointed spineS = .....4+e.se2e-. 2

2(1). Proximal part of subapical lobe with 2

rodlike setae; lateral plate of phallosome

without denticles: Gio 1860s) tie IEA, git hortensis group

Proximal part of subapical lobe with 3

rodlike setae; apex of lateral plate of

phallosome with some denticles ........... seyrigi group

PULCHRITHORAX GROUP

This group, as keyed above, corresponds to Edwards' group A (1941:

249) with only one species, namely, the Ethiopian C, pulchrithorax. Its out-

standing ornamentation on the dorsum of head, scutum, pronotum and upper

pleura is very distinctive as described and illustrated by Edwards (1941: 254).

The female buccopharyngeal armature is, however, rather similar to species

in the seyvigi group.

a2 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

HORTENSIS GROUP

This group contains 3 or perhaps more species from the Mediterranean.

Their external characters are more or less similar to the territans group of

Neoculex, but with propleural scale patch extended to prosternum in some

species. At present, 3 forms, namely hortensis, arbieeni and quettensis are

grouped together here. They appear to show the characters of the subgenus

better than the other two groups.

SEYRIGI GROUP

I tentatively place 4 forms, namely seyrigi, peringueyi, salisburensis

and avianus, all from the Ethiopian region, in this group, based on the char-

acters given in the key. They are similar to members of the hortensis group

in external features, but the male phallosome and other features of the male

terminalia resemble members in the subgenus Eumelanomyia.

Subgenus III. EUMELANOMYIA Theobald

1909 Eumelanomyia Theobald, Mon. Cul. 5:240; Haplotype: znconspicuosus

(presently known as albiventris).

1910 Protomelanoconion Theobald, Mon. Cul. 5:462; Haplotype: fusca (pres-

ently known as horvridus).

1930 Culex (Mochthogenes) Edwards, Bull. ent. Res. 21:306; Type:C. malaya

Leicester 1908.

Culex (Neoculex) in part of Edwards (1932: 193-195); Edwards (1951: 249-

269); Barraud (1934: 347-352); Bohart & Ingram (1946); Bohart (1953:187);

Delfinado (1966: 124-128); Bram (1967: 23-32).

Culex (Mochthogenes) of Edwards (1932:195; 1941:277-279); Barraud

ioe eecaan © Baisas (1935:175-177); Delfinado (1966:128-135); Bram

967:33-42).

Subgeneric Characters. As indicated in the key to subgenera, with the

following additional description. Small to medium sized, wing length usually

not more than 3.0 mm., dark brown to black species. Head. Male palpus

from 0. 2 to longer than the length of proboscis; antennal flagellomeres 1-10

usually with a single whorl of long hairs each, sometimes also with a much

smaller whorl] of short but conspicuous hairs in addition (Fig. 3). Thorax.

Scutal scales narrow and usually entirely dark, rarely pale; pleural scaling

absent or only a few scales present on upper corner of sternopleuron. Abdo-

men. Terga entirely dark or with apical bands, apicolateral pale spots, rarely

with basal bands. Male Terminalia. (Fig. 3) Phallosome small, generally

broad, oval or subspherical in shape; tergal bridge at or below the midpoint of

lateral plates; usually with several denticles over the upper tergal surface,

rarely bare; proctiger with small dark crown of fine spinelike spicules, some

coarse ones present or absent; proximal division of subapical lobe always with

3 rodlike setae; distal division usually with at least a broad leaflet; distimere

slender, sickle-shaped or else modified.

Systematics. Members of this subgenus can be easily distinguished

from the other two subgenera by smaller size, dark scaled scutum and the ab-

sence of scale patches on the pleura. The phallosome is variable among dif-

ferent species but all are remarkably constant with regard to shape and posi-

tion of tergal bridge and do not appear to overlap in these characters with spe- _

cies in Neoculex. The fine texture of the spicules of the proctiger crown and

the presence of 3 rodlike setae in the proximal division of the subapical lobe

are also characteristics of this subgenus.

Sirivanakarn: Reclassification of Neoculex 73

This subgenus contains the majority of species from the Ethiopian

region and the Indomalayan part of the Oriental region. The number of spe-

cies involved is the largest of the 3 subgenera recognized here. They are di-

vided into 4 major groups as follows:

KEY TO THE GROUPS OF SUBGENUS EUMELANOMYIA

1. Male palpus as long as or longer than

proboscis; flagellomeres 1 to 10 of male

antenna each with a single large whorl

GE 40g DRIP S666 aac kn cary aun Pike dem epee kee Se ee ee 2

Male palpus from 0. 2 to 0. 75 the length of

proboscis; flagellomeres 1 to 10 of male

antenna each with a smaller whorl in ad-

dition to a large normal whorl (Fig. 3)% .........64.. 3

2(1). Decumbent scales in center of vertex broad;

acrostichal bristles absent; tergal bridge

of phallosome present or absent ...... | eumelanomyia group

Decumbent scales in center of vertex narrow; |

acrostichal bristles present; tergal bridge

\ of phallosome present. ...... pg 1. rubinotus -rima group

3(1). Male palpus about 0.75 the length of pro-

boscis; acrostichal bristles absent; lower

anterior mesepimeral bristle absent . . . pvotomelanoconion group

Male palpus usually about 0. 2 the length

of proboscis, sometimes longer to about

0.75; acrostichal bristles present; lower

anterior mesepimeral bristle usually

present mochthogenes group

EUMELANOMYIA GROUP

This group corresponds to Edwards' group B (1932) and group D. (1941)

or albiventris group. Five species, all from the Ethiopian region are placed

here: albiventris, andersianus, vinckei, acrostichalis and kanyamwerima and

perhaps also kilara and garioui. The group is characterized as in the key and

may be further subdivided into two subgroups on the basis of presence or ab-

sence of a tergal bridge of the phallosome. Certain members of this group ap-

pear to show affinity to the subgenus Culiciomyia. |

RUBINOTUS -RIMA GROUP

This group corresponds to the 7ima group or group C and Neoculex s.

sty. group, in part, of Edwards (1941). It could be subdivided into 2 sub-

groups: 1) rubinotus subgroup with rubinotus, kingianus, andreanus,, Pseudo-

andreanus from the Ethiopian region and simplicicornis from the Indomalayan

3 As illustrated, the term "normal whorl" as used here refers to the large

tuft of 10 to over 20 long hairs arising from a series of tubercles encircling

the middle part of each flagellomere, whereas the term "small or minor

whorl" refers to a much smaller tuft with 4-8 short hairs arising near the

junction of the flagellar segments.

14 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

area (Borneo) by having abdominal terga entirely dark and by having the pleural

integument uniformly dark brown; and 2) vima subgroup with vima, subrima,

galliardi, calabarensis, wigglesworthi, insignis, albertianus, wansoni and

perhaps also sunyaniensis and others as listed by having apical band or apico-

lateral spots on abdominal terga and by having a pattern of dark and pale areas

on the pleura. The members in the rubinotus subgroup show strong affinity to

the subgenus Lophocevraomyia on the basis of several characters indicating

that the latter subgenus is probably derived from them.

PROTOMELANOCONION GROUP

This group includes brevipalpis, a dominant form from several areas

in the Oriental region, horridus from the Ethiopian region and stellatus from

Seychelles. It is closely related to the mochthogenes group, but with longer

male palpi and differing in other constant features as indicated in the key.

-_MOCHTHOGENES GROUP

This group is dominant in the Indomalayan areas and other southern

parts of the Oriental region in which it is represented by many distinct line-

ages (see list above). It is perhaps represented by a single lineage (incon-

spicuosus subgroup) in the Ethiopian region. In the South Pacific, it is repre-

sented by a single species (femineus). I recognize 8 subgroups in this group.

They are separated as follows:

KEY TO THE SUBGROUPS OF GROUP MOCHTHOGENES

1. Decumbent scales in the center of vertex

entirely or predominantly broad along

MteriGn GCUlar TING Se Sn See ee Oe ee eels 8 2

Decumbent scales in the center of vertex

entirely narrow ee a ee SES a ee a eee 9)

2(1). Distimere of male terminalia simple,

SICMICG SHAOGO es eee ees ees pers 3

Distimere of male terminalia strongl

PIOGITLS CERO MT GOOVE a I Og gs a eae 4

3(2). Lateral plate of phallosome without large

internal process; minor flagellar whorls

of short hairs of male antenna present. .. hinglungensis Subgroup

Lateral plate of phallosome with a large

internal process; minor flagellar whorls

of short hairs of male antenna absent. ..... wuncinatus subgroup

4(2). Distimere sharply angled at the middle of

dorsal curvature, its basal half thick,

distal half narrow and tapered to a curved

BOING Sy A a ae aan a 2 ee « = ainconspicuosus subgroup

Distimere divided into a short dorsal and

a long ventraldrm<.. 3 we. ws BO ears Ue malayi subgroup

Sirivanakarn: Reclassification of Neoculex 15

5(1). Male antennal flagellar whorls with rather

weak and relatively few hairs; size minute

or very small, wing length usually not

OXCCCCING 3: O IIs: ay. eel we ocd eer ee dod ra Taba cass shige 6

Male antennal flagellar whorls with strong

and numerous hairs; size relatively large,

wing length usually.3. 0 mim orimore.'. oe lh thal dil FAs 7

6(5). Abdominal terga entirely dark; phallosome

of male terminalia short and oval in shape;

basimere small, slender and conical in

SNOQC. ae is (nia: hs sac Ca aA OAS ARR Road aS mere castrensis subgroup

phallosome of male terminalia tubular

in shape; basimere swollen and broadly

OV at UD SIAC a. oo a:. wicnccetan cu. ak he ce een ae ale femineus subgroup

7(5) Male phallosome heavily sclerotized and

dark, lateral plate rodlike and pointed

with some heavy lateral teeth ......... otachati subgroup

Male phallosome weakly sclerotized and

pale yellow or brown, lateral plate oval

or subspherical in shape; teeth confined

to-inner: terval, surface: veils sie) vais bey. ener’ tenuipalpis subgroup

As indicated in the above key, the mochthogenes group is rather com-

plex as it contains several lineages, most of which can be easily recognized

by the short male palpus more or less similar to the female. Only the tenui-

palpis subgroup, as far as known, consists of some members with male palpi

longer than those of the females. These are fenuipalpis, hayashii and okinawae

which Edwards (1932:194-195) and Bohart (1953:187) placed with the proto-

melanoconion group of Neoculex s. lat. The femineus subgroup is also rather

anomalous in male terminalia but since it shows several characters common

to most mochthogenes members, I place it here for the present.

Culex gamma Seguy (1924, Encycl. ent., :47) was described from larva

only and cannot definitely be placed with any subgenus according to the present

scheme,

ACKNOWLEDGEMENTS

I wish to thank Dr. Botha de Meillon, Project Leader, Southeast Asia

Mosquito Project, Smithsonian Institution and Dr. Alan Stone, Agriculture

Research Service, U.S. Department of Agriculture for their many helpful sug-

gestions in improving the manuscript. I also wish to thank Dr. Stone for the

loan of specimens from the United States National Museum and several papers

on Neoculex in his private file for study and consultation. I am indebted to Dr.

P. F. Mattingly for his courtesy extended to me while studying the specimens

at the British Museum. Dr. John N. Belkin, Department of Zoology, Univer-

sity of California, Los Angeles, kindly read the manuscript and I wish to thank

him for his comments and advice. I thank Mr. Vichai Malikul for his assist-

ance with the illustrations.

76 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

REFERENCES CITED

BAILLY-CHOUMARA, H.

1965. Contribution a l'etude des Culex (Neoculex) (Diptera, Culicidae)

de la region ethiopienne. Bull. Soc. Path. exot. 58:660-664.

BAILLY-CHOUMARA, H. & A. RICKENBACH

1966. Contribution a l'etude des Culex (Diptera, Culicidae) de la region

ethiopienne. Description de Culex (Neoculex) garioui sp. n.

eerie nouveau du Cameroun. Bull. Soc. Path. exot. 59:

~ 8. ‘

BAISAS, F.E.

1935. Notes on Philippine mosquitoes, II. Genus Culex: Groups

oe Mochthogenes and Neoculex. Philip. J. Sci.

BARRAUD, P. J. |

1934. The fauna of British India including Ceylon and Burma. Diptera

V. Family Culicidae, tribes Megarhinini and Culicini. Taylor

& Francis, London. 463 pp.

BELKIN, J.N.

1962. The mosquitoes of the South Pacific. Univ. Calif. Press,

Berkeley, 2vols., 608 and 412 pp.

BOHART, R.M.

1948(1949). The subgenus Neoculex in America north of Mexico

(Diptera, Culicidae). Ann. ent. Soc. Amer. 41:330-345.

1953. A new species of Culex and notes on other species of mosquitoes

ee eae (Diptera, Culicidae). Proc. ent. Soc. Wash. 55:

83-188.

BRAM, R.A.

1967. Contributions to the mosquito fauna of Southeast Asia (Diptera,

Culicidae). II. The genus Culex in Thailand. Contr. Amer.

ent. Inst. 2:1- 296.

1969. Relationships of adult and larval anatomy in the supra- specific

classification of the genus Culex in Southeast Asia (Diptera:

Culicidae). Mosq. Syst. Newsletter 1:9-12.

BRUG, S.L.

1934. ee on Dutch East Indian mosquitoes. Bull. ent. Res. 25:501-

519.

BRUNHES, J.

1968. Contribution a l'etude des Culicides de Madagascar. Synonymie

entre Culex (N.) seyrigi Edwards 1941 et Culex (N.) robici

Doucet 1960(1950). Description de la nymphe et de la femelle

Jae as rey (N.) seyrigi Edwards. Cah. ORSTOM, ser. Ent.

med. 6:15-18. |

BRUNHES, J. et al.

1967. Contribution a l'etudes des Culex de la region ethiopienne appar-

tenant au Sous-genre Mochthogenes (Diptera, Culicidae) avec

description des males de cing nouvelles especes. Cah. ORSTOM,

ser. Ent. med. 5:43-52.

Sirivanakarn: Reclassification of Neoculex 17

BRUNHES, J. & J. RAMBELO

1968. Contribution a l'etude des Culicides de Madagascar. Description

des adultes, nymphe et larve de Culex (Neoculex) chauveti sp. n.

Cah. ORSTOM, ser. Ent. med. 6:113-118.

CARPENTER, S.J. & W.J. LaCASSE

1955. Mosquitoes of North America (North of Mexico). Univ. Calif.

Press vl + 360 pp.

CERVONE, L.

1957. Sulla presenza di Culex (Neoculex) martinit Medschid in Provincia

di Latina e contributo alla conoscenza delle specie. Riv. Paras-

sit. 18:235- 248.

DELFINADO, M.D.

1966. The Culicine mosquitoes of the Philippines, tribe Culicini

(Diptera, Culicidae). Mem. Amer. ent. Inst. 7:1-252.

de MEILLON, B.

1943. New records and new species of Nematocera Pinter) from the

Ethiopian region. J. ent. Soc. S. Afr. vi:90-113.

DOBROTWORSKY, N.V.

1956. Notes on Australian mosquitoes (Diptera, Culicidae) I. Some

stacey Fe the subgenus Neoculex. Proc. Linn. Soc. N.S.W.

81:105- ‘

1965. The mosquitoes of Victoria (Diptera, Culicidae). Melbourne

Univ. Press, 1 vol. pp. 193-202.

DYAR, H.G. |

1905. Remarks on genitalic genera in the Culicidae. Bull. 97, N.Y.

State Mus. p. 48.

EDWARDS, F. W.

1930. Mosquito notes - IX. Bull. ent. Res. 21:305.

1932. in Wytsman, Genera Insectorum. Diptera. Family Culicidae.

Fasc. 194, Desmet-Verteneuil, Brussels. 258 pp.

1941. Mosquitoes of the Ethiopian Region. III. Culicine adults and

pupae. Brit. Mus. (Nat. Hist), London. 499 pp.

HAMON, J., ADAM, J.P. & J. MOUCHET

1955. Contribution a l'etude des Neoculex (Diptera, Culicidae) de la

region ethiopienne. 3. Description de Neoculex laplantei Sp. nN.

Bull. Soc. Path. exot. 48:862- 866.

HAMON, J., HOLSTEIN, M. & E. RIVOLA

1957. Description d'un nouveau moustique du Congo Belge: Culex

(Neoculex) vinckei sp. n. Bull. Soc. Path. exot. 50:681-689.

HAMON, J. & J. MOUCHET

1955. Contribution a l'etude des Neoculex (Diptera, Culicidae) de la

region ethiopienne. 2. Description de Neoculex adami sp. n.

Bull. Soc. Path. exot. 48:860-862.

78 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

HAMON, J. & A. RICKENBACH

1955. Contribution a l'etude des Neoculex (Diptera, Culicidae) de la

region ethiopienne. 1. Corrections de quelques descriptions de

terminalia males donnees par Edwards, avec etude d'une nou-

velle variete. Bull. Soc. Path. exot. 48:845-859.

KING, W.V. & H. HOOGSTRAAL

1947. Two new species of Culex (Neoculex) from New Guinea (Diptera,

Culicidae). Proc. ent. Soc. Wash. 49:65-69.

KLEIN, J.M. & S. SIRIVANAKARN

1969. Four new species of Culex, subgenus Mochthogenes from South-

oe Asia (Diptera, Culicidae). Proc. ent. Soc. Wash. 71:582-

LaCASSE, W.J. &S. YAMAGUTI

1950. Mosquito fauna of Japan and Korea. Off. Surgeon, 8th U.S. Army

Kyoto, Honshu. 3rd Ed., 268 pp.

LIEN, J.C.

1968. New species of mosquitoes from Taiwan (Diptera, Culicidae)

Part V. Three new subspecies of Aedes and seven new species

of Culex. Trop, Med. Nagasaki, Japan 10:217- 262.

MATTINGLY, P. F.

1955. Le sous-genre Neoculex (Diptera, Culicidae) dans la sous-region

mediterraneene. I. Espece, sous-espece et synonymies nou-

velles. Ann. Parasit. hum. comp. 30:375-388.

MATTINGLY, P.J. & E.N. MARKS

1955. Some Australian mosquitoes (Diptera, Culicidae) of the subgenera

Sanaa and Neoculex. Proc. Linn. Soc. N.S.W. 80:1 163-

76.

SOMEREN, E.C.C. van

1947. The OR aL of a new mosquito from the Seychelles. E. Afr.

med. J 9

1951, ode from Kenya and Uganda. Proc. R. ent. Soc. Lond.

B) 20:1-9.

SOMEREN, E.C.C. van & J. HAMON

1964. Ethiopian Culicidae (Diptera). A new species of Culex from

Tanganyika, the description of the pupae of Aedes usambara

Mattingly and the early stages of Evretmapodites tonsus Edwards.

J. ent. Soc. S. Afr. 27:78-85.

STONE, A.

1961. A synoptic catalog of the mosquitoes of the world, Supplement I

(Diptera: Culicidae). Proc. ent. Soc. Wash. 63:29-52,

1963. A synoptic catalog of the mosquitoes of the world, Supplement II

(Diptera: Culicidae). Proc. ent. Soc. Wash. 65:117-140.

1967. A synoptic catalog of the mosquitoes of the world, Supplement III

(Diptera: Culicidae). Proc. ent. Soc. Wash. 69:197- 224.

1970. A synoptic catalog of the mosquitoes of the world, Supplement IV

(Diptera: Culicidae). Proc. ent. Soc. Wash. 72:137-171.

Sirivanakarn: Reclassification of Neoculex 19

STONE, A., KNIGHT, K.L. & H. STARCKE

1959. A synoptic catalog of the mosquitoes of the world (Diptera:

Culicidae). Ent. Soc. Amer. (Thomas Say Found.) Washington,

D.C. 358 pp.

THEOBALD, F.V.

1907. A monograph of the Culicidae or mosquitoes. IV. London.

639 pp.

1910. A monograph of the Culicidae or mosquitoes. V. London.

646 pp.

Territans group

phallosome

proctiger

—_J

subapical lobe

| of basimere

C. territans

Pseudomelanoconia group

Sang

Crassistylus group

—"

C.leonardi (after Belkin 1962 )

Fig.2

Hortensis group proctiger

phallosome subapical lobe

of basimere

C. hortensis

aig

C. arbieeni

Pulchrithorax group

C. pulchrithorax (after Edwards 1941)

Fig. 3

Eumelanomyia group

phallosome

subapical lobe

of basimere

C. albiventris (after Edwards 1941)

Cor

Rubinotus-rima group

proctiger

Sn.

C. kingianus

( after Edwards 1941 )

Protomelanoconia group

a i,

x Ay is

RYE Rad

SA Va

Vole 1 2

Ni lf

as. ES

~ it

Bivins

Ai;

Z A

4 y,) B4

ry ‘A ‘a

i] iy aKa a4,”

if Anant a AM

AA A

/ AANA Ac?

Ara” arn

My FAs

a ha

C. brevipalpis

Sirivanakarn: Reclassification of Neoculex 83

INDEX

Valid names are in roman type, synonyms are italicized. Italicized

page numbers indicate the primary treatment of the subgenus. Numbers in

parentheses refer to the figures of the male terminalia of the species in ques-

tion.

Acalleomyia

acrostichalis

adami

Aedinus

albertianus

albiventris

amaniensis

ander sianus

andreanus

apicalis

ar bieeni

arizonensis

avianus

Barraudius

boharti

bokorensis

brevipalpis

caeruleus

calabarensis

campilunati

Carrollia

castrensis

castrensis subgroup

castor

cataractarum

chaetoventralis

chauveti

cheesmanae

chiyutoi

crassistylus

crassistylus subgroup

Culex |

Culiciomyia

culionicus

derivator

deserticola

douglasi

dumbletoni

Eumelanomyia

Eumelanomyia subgenus

Eumelanomyia group

femineus

femineus subgroup

fergusoni

foliatus

fusca

galliardi

gamma

garioul

gaufini

hackeri

hamoni

65, 67, 73 (3)

68, 75

67, 70

64, 66, 67, 73

65, 66, 67, 69, 72, 73

14, 15

84 Contrib. Amer. Ent. Inst., vol. 7, no. 3, 1971

hayashii

hinglungensis

hinglungensis subgroup

horridus

hortensis

hortensis group

inconspicuosa

inconspicuosus

inconspicuosus subgroup

impudicus

insignis

iphis

Isostomyia

judaicus

kanyamwerima

khazani

kilara

kingianus

kiriensis

laplantei

Lasiosiphon

latifoliatus

latus |

laureli

leonardi

lini

Lophoceraomyia

Lutzia

Maillotia, subgenus

malayi

malayi subgroup

martinii

Melanoconion

Micraedes

Microculex

mijanae

millironi

Mochlostyrax

Mochthogenes

-mochthogenes group

Neoculex, subgenus

okinawae

orstom

otachati subgroup

pedicellus

peringueyi

pilifera

pipiens group

pluvialis

postspiraculosus

protomelanoconion

protomelanoconion group

pseudoandreanus

pseudomelanoconia

pseudomelanoconia group

pulchrithorax

pulchrithorax group

reevesi

rima

68, 75

72, 74

72 (2)

67, 11,

68, 74

73 (3)

67, 68, 72, 73, 74, 75

72, 73, 74, 75

71 (2)

13, 74

Sirivanakarn: Reclassification of Neoculex

rima subgroup

rubensis

rubinotus

rubinotus subgroup

rubinotus-rima group

salisburensis

selai

seyrigi

seyrigi group

shrivastavil

simplicicornis

simpliciforceps

stellatus

subrima

sumatranus

sunyaniensis

tenuipalpis

tenuipalpis subgroup

territans

territans group

tricontus

tricuspis

uncinatus

uncinatus subgroup

uniformis group

vinckei

wansoni

wigglesworthi

yeagerl

67,

68, 75

65, 67, 69, 70, 72 (1)

fi? £5

ae!

ai .

a - ‘ | =) . , 4 | i x P

ec. a ee a, ix - pea a tee ok ie ee ie

: e = _ ’ Aid " ee : Lear,

Berta oy tae

—/.

—_

No.

(Continued from inside front cover)

Parts of Volume 4, with prices

Mosquito studies (Diptera, Culicidae), 1968.

X. Peters, T. Michael. Dixinae originally described from North America. 7 pages.

XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes

originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay.

29 pages. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XII. Berlin, O. G. W. A revision of the Neotropical subgenus Howardina of Aedes.

Carlson, Robert W. and Fred B. Knight, 1969.

Biology, taxonomy, and evolution of four sympatric Agrilus beetles (Coleoptera:

Buprestidae). 104 pages, 47 figures. Price: $3.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XIII. Barr, Ralph A. and Sylvia Barr. Pupae of the genus Psorophora in America

north of Mexico. 19 pages, 11 figures.

XIV. Berlin, O. G. W. Redescription of Toxorhynchites (T.) gigantulus from the

Philippines. 7 pages, 2 figures.

XV. Zavortink, Thomas J. A new species of treehole breeding Anopheles from

the southwestern United States. 12 pages, 4 figures.

Price: $1.25, postpaid.

Matthews, Robert W., 1970. A revision of the genus Spathius in America north of Mexico

(Hymenoptera, Braconidae). 86 pages, 31 figures. Price: $3.50, postpaid.

postpaid.

Parts of Volume 5, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XVI. Zavortink, Thomas J. A new species of treehole breeding Aedes (Ochlerotatus)

from southern California. 6 pages, 2 figures.

XVII. Adames, Abdiel J. and Charles L. Hogue. Two new species of Deinocerites from

Costa Rica. 6 pages, 6 figures.

XVIII. Berlin, O. G. W. The subgenus Micraedes of Culex. 30 pages, 12 figures, 1 map.

Price: $2.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XIX. Zavortink, Thomas J. The treehole Anopheles of the New World. 35 pages, 12

figures. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XX. Schick, Robert X. The Terrens Group of Aedes (Finlaya). 158 pages, 60 figures.

Price: $5.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

V. Reinert, John F. Genus Aedes, subgenus Diceromyia Theobald in southeast Asia.

43 pages, 15 figures. Price: $1.50, postpaid.

Howden, Anne T. The Tanymecini of the West Indies (Coleoptera: Curculionidae). 74 pages,

76 figures. Price: $2.75, postpaid.

Kethley, John B., 1970. A revision of the family Syringophilidae (Prostigmata: Acarina).

76 pages, 38 figures. Price: $2.75, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

VI. Mattingly, P. F. The genus Heinzmannia Ludlow in southeast Asia. 104 pages,

ol figures. Price: $4.00, postpaid.

Parts of Volume 6, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXI. Belkin, John N., Sandra J. Heinemann, and William A. Page. The Culicidae of

Jamaica. 458 pages, 126figures. Price: $15.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

Vil. Tyson, W. H. Genus Aedeomyia Theobald in southeast Asia. 27 pages, 6 figures.

Vil. Tyson, W. H. Genus Aedes, subgenus Mucidus Theobald in southeast Asia. 54 pages

21 figures. Price: $3.50, postpaid.

Parts of Volume 7, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XX. Zavortink, Thomas J. A new subgenus and species of Aedes from Arizona.

11 pages, 2 figures.

XXIII. Schick, Robert X. Additions and corrections to the revision of the Aedes

terrens group. 27 pages, 8 figures. Price: $1.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1971.

XXIV. Adames, Abdiel J. A revision of the crabhole mosquitoes of the genus

Deinocerites. 154 pages, 58 figures. Price: $6.00, postpaid.

(Continued from inside back cover)

No. 3. Contributions to the mosquito fauna of southeast Asia, 1971.

IX. Zavortink, Thomas J. The genus Orthopodomyia Theobald in Southeast Asia.

37 pages, 12 figures.

X. Dobrotworsky, N. V. The genus Culiseta Felt in Southeast Asia. 24 pages, 8 figures.

XI. Sirivanakarn, Sunthorn. A proposed reclassification of Neoculex Dyar based

principally on the male terminalia. 24 pages, 3 figures. Price: $3.50, postpaid.

Contributions

of the

American Entomological Institute

Volume 7, Number 4, 1971

ot be

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA, XII.

ILLUSTRATED KEYS TO THE GENERA OF MOSQUITOES

(DIPTERA, CULICIDAE)

Peter F. Mattingly

Ja HSOWN; Ay

SEP 16 1971

LiZRARIED

CONTRIBUTIONS

of the

AMERICAN ENTOMOLOGICAL INSTITUTE

The Contributions are for larger papers on insects. Each paper is a separate number, with separate

pagination and index. Separate numbers aggregating about 500 pages constitute a volume. Issues appear

irregularly, as suitable manuscripts are available.

Copies are sold separately or in subscriptions to complete volumes. Complete volumes are $12.00.

The price of separate numbers varies. Subscribers are billed for each volume with its beginning number,

and receive the parts as issued. Orders for separate numbers that total less than $8.00 must be accom-

panied by payment. Address orders or correspondence to the American Hate potonices Institute, 5950

Warren Road, Ann Arbor, Michigan 48105, U.S.A.

Parts of Volume 1, with prices

No. 1. Dasch, Clement E., 1964. The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae).

75 pages, 69 figures. Price: $2.25, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1965.

I, Belkin, John N., etal. A project for a systematic study of the mosquitoes of Middle America.

II. Belkin, John N., et al. Methods for the collection, rearing and prea voter of mosquitoes.

78 pages, 4 figures. Price: $2.25, postpaid.

No. 2a. Sameasno. 2, but in Spanish. Price: $2.25, postpaid.

No. 3. Matthews, Robert W., 1965. The biology of Heriades carinata Cresson. (Hymenoptera,

Megachilidae). 33 pages, 23 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1965.

II. Ramalingam, Shivaji, and John N. Belkin. Two new Aedes from Tonga and Samoa.

IV. Belkin, John N. The mosquitoes of the Robinson-Peabody Museum of Salem expedition to

the southwest Pacific, 1956.

34 pages, 3 figures. Price: $1.00, eee

No. 5. Mosquito studies (Diptera, Culicidae), 1965.

V. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from Middle America. 95 pages. Price: $2.75 postpaid.

No. 6. Mosquito studies (Diptera, Culicidae), 1966.

VI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from North America. 39 pages. Price: $1.00, postpaid.

No. 7. DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae (Homopotera,

Cicadellidae). A synopsis of the genus Ponana.

86 pages, 257 figures. Price: $2.50, postpaid.

No. 8. Contributions to the mosquito fauna of southeast Asia, 1967.

I. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald in Thailand.

55 pages, 20 figures. Price: $1.75, postpaid.

Parts of Volume 2, with prices

No. 1. Contributions to the mosquito fauna of southeast Asia, 1967.

Il. Bram, Ralph A. The genus Culex in Thailand (Diptera: Culicidae).

296 pages, 104 figures. Price: $9.00, postpaid.

No. 2. Schuh, Toby, 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region.

35 pages, 30 figures. Price: $1.25, postpaid.

No. 3. Snider, Richard J., 1967. The chaetotaxy of North American Lepidocyrtus s. str. (Collembola:

Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid. Sa

No. 4. Contributions to the mosquito fauna of southeast Asia, 1968.

Ii. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald of southeast

Asia. 76 pages, 26figures. Price: $2.25, postpaid.

No. 5. Contributions to the mosquito fauna of southeast Asia, 1968.

IV. Knight, Kenneth L. Species of the subgroup Chrysolineatus of group D, genus Aedes,

subgenus Finlaya Theobald. 45 pages, 12 figures. Price: $1.50, postpaid.

Parts of Volume 3, with prices

No. 1. Mosquito studies (Diptera, Culicidae), 1968.

VIl. Belkin, John N. The Culicidae of New Zealand.

182 pages, 30 figures. Price: $5.50, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1968.

VIII. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia.

221 pages, 41 figures. Price: $6.75, postpaid.

No. 3. Fleming, Richard C., 1968. Head musculature of sphinx moths (Lepidoptera: Sphingidae).

32 pages, 37 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1968.

IX. Belkin, John N. The type specimens of New World mosquitoes in European museums.

69 pages. Price: $2.00, postpaid.

(Continued on back cover)

Contributions

of the

American Entomological Institute

Volume 7, Number 4, 1971

-—

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA. XII.

ILLUSTRATED KEYS TO THE GENERA OF MOSQUITOES

(DIPTERA, CULICIDAE)

Peter F. Mattingly

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. XII.

CONTENTS

INTRODUCTION. ... .. cone: hee oo or

DISEASE RELATIONS AND BIONOMICS .........02cc0c 08.

GEOGRAPRICAL DiS TRIBUTION 2 Sock ee ee ae 8 ak ee ee

RYO FO CRE GEMMA L Sooo iw 8 ea ee Sarg ew ie

Fo Re a ie ae ee ee See te eee rae

De I os a a Se a eee cece, ae ee

Dil UA oe gee nce wa ey ce te a a a ee

TV, “FOURTH STAGE LARVAE 6. 6 eee we wee es ee

MOTES ON TH Ree ees ee ee oe Se ee ee

hese uber ove ae ns Oe Ture \ANGOY WEEK Soke Mabsek “LOO SMGhX Yitmged loom! Soom Desai hoiiet Ween Teen Sie Heed Pe Je hen Soles Stuy, MOkek emy Sioa Maes Iolani aeeeet Nennat |

Oo PF, FP CFO FE fF

ILLUSTRATED KEYS TO THE GENERA OF mosqutToEs *

Peter fF. Mattingly ”

INTRODUCTION

The suprageneric and generic classification adopted here follow closely

the Synoptic Catalog of the Mosquitoes of the World (Stone et al., 1959) and the

various supplements (Stone, 1961, 1963, 1967, 1970). Changes in generic no-

menclature arising from the publication of the Catalog include the substitution

of Mansonia for Taeniorhynchus and Culiseta for Theobaldia, bringing New and

Old World practice into line, the substitution of Toxorhynchites for Megarhinus

and Malaya for Harpagomyia, the suppression of the diaeresis in Aédes,

Aédeomyia (formerly Aédomyia) and Paraédes (Christophers, 1960b) and the

inclusion of the last named as a subgenus of Aedes (Mattingly, 1958). Theonly

new generic name to appear since the publication of the Catalog is Galindomyia

(Stone & Barreto, 1969). Mimomyia, previously treated as a subgenus of

Ficalbia, is here treated, in combination with subgenera Etorleptiomyia and

Ravenalites, as a separate genus. Ronderos & Bachmann (1963a) proposed to

treat Mansonia and Coquillettidia as separate genera and they have been fol-

lowed by Stone (1967, 1970) and others. I cannot accept this and they are here

retained in the single genus Mansonia.

It will be seen that the treatment adopted here, as always with

mosquitoes since the early days, is conservative. Inevitably, therefore, dif-

ficulties arise in connection with occasional aberrant species. In order to

avoid split, or unduly prolix, couplets I have preferred, in nearly every case,

to deal with these in the Notes to the Keys. The latter are consequently to be

regarded as very much a part of the keys themselves and should be constantly

borne in mind. A bibliography of key works is included for the benefit of

el who may wish to carry out identification at the subgeneric or species

evel.

DISEASE RELATIONS AND BIONOMICS

Disease Relations

Setting aside mechanical transmission and phoresy, mosquitoes serve

as vectors of three groups of human pathogens. These are Haemosporidia

(four species of human malaria parasite and an occasional simian malaria par-

asite, all belonging to the genus Plasmodium), two or more species of Filari-

oidea belonging to the genera Brugia and Wuchereria, and the arboviruses, of

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

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

of the Surgeon General.

2Department of Entomology, British Museum (Natural History), London SW 7,

England.

2 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

which more than 40 have beenrecovered from man and another 40 are believed

to infect man onthe evidence ofantibody surveys. The diseasesto which these

give rise are discussed as ecological systems by Mattingly (1969). Mono-

graphs or reviews dealing with particular groups of diseases include Edeson &

Wilson (1964), Garnham (1966), Reeves (1965), Russell et al. (1963), Taylor

(1967), Berge et al. (1970) and Ward & Scanlon (1970). For epidemiology and

epidemiological sampling methods see Detinova (1968), Muirhead- Thomson

(1968) and Pampana (1969). For control methods see Busvine (1966).

Human malaria is transmitted exclusively by Anophelesspp. Vectors

include An. labvanchiae Falleroni, sacharovi Favre, sergentii (Theobald),

superpicitus Grassi and pharoensis Theobald in parts of the Mediterranean

area, An. funestus Giles, moucheti Evans, nili (Theobald) and members of

the gambiae Giles complex in the Ethiopian Region, An. stephensi Liston,

fluviatilis James and pulcherrimus Theobald in western Asia, An. culicifacies

Giles in India and Ceylon, An. maculatus Theobald, sundaicus (Rodenwaldt)

and members of the An. barbirostris Van der Wulp, hyrcanus (Pallas),

umbrosus (Theobald), leucosphyrus Dunitz, minimus Theobald and annularis

Van der Wulp groups in southern Asia, members of the An. punctulatus com-

plex in Melanesia and An. pseudopunctipennis Theobald, bellator Dyar & Knab,

cruzii Dyar & Knab, darlingi Root, aquasalis Curry, albimanus Wiedemann,

albitarsis Lynch Arribalzaga and nuneztovari Gabaldon in Central and South

America. Other species may be locally important and in general a list such

as the above can present only a very crude picture since the importance ofa

particular vector in a given area will vary with the prevailing ecological con-

ditions (see, e.g. Reid in Ward & Scanlon, 1970). Complicating factors in-

clude the development of insecticide resistance among a number of important

vectors (World Health Organization, 1970) and the habit of some others of feed-

ing and resting outside houses where they fail to make contact with the insecti-

cide. (For behavioral aspects see Mattingly, 1962b). Vectors of wuchererian

filariasis include several of the major malaria vectors and, in addition, the

highly domestic Culex pipiens fatigans. Wiedemann throughout the tropics except

in the Pacific area where the nocturnal periodicity of the parasite is suppres-

sed and day-biting vectors, particularly Aedes Stegomyia) polynesiensis

Marks, take over. Vectors of brugian filariasis include Anopheles and

Mansonia spp. (See Wharton, 1962, Edeson & Wilson, 1964, Reid, 1968).

Viruses known or believed to infect man have been recovered from more

than 150 species of mosquitoes belonging to 14 different genera (Aedeomyia,

Aedes, Anopheles, Culex, Culiseta, Deinocerites, Eretmapodites, Haemagoagus,

Limatus, Mansonia, Psorophora, Sabethes, Trichoprosopon, Wyeomyia).

Bionomics

The egg stage, though of cardinal ecological importance, has been

somewhat neglected. The nearest to a general review will be found in

Mattingly (Riv. Parassit., in press). Oviposition sites (or larval breeding

places) may be roughly classified as follows:

A. Running water habitats (stream edges, mainly anophelines).

B. Still water habitats.

1. Ground water habitats

a. Permanent (anophelines and culicines other than Aedes and

aedine genera).

b. Temporary ie Aedes).

c. Specialized (crab holes, rock holes, wells, soak aways)

(various culicines, a few anophelines in wells).

2. Container habitats (artifacts, tree holes, cut, split or bored

bamboos, leaf axils, plant pitchers, flower petals, bracts,

spathes, fallen leaves or rachids, fruits and husks, snail shells,

cup fungi etc.) (Toxorhynchites, Anopheles subgenus Kerteszia,

sabethine genera, Some aedine genera, some subgenera of Aedes,

Mattingly: Illustrated Keys to the Genera of Mosquitoes 3

individual species and species groups of Anopheles and various

culicine genera).

Larval bionomics are reviewed in the various taxonomic monographs

listed on p. 37, notably Hopkins (1952), and by Laird (1956). Since the intro-

duction of residual insecticides a quarter of a century ago adult bionomics have

received increasing attention. Coverage in taxonomic monographs is not al-

ways adequate. Exceptions are Gillies & de Meillon (1968) and Reid (1968).

Among general reviews Bates (1949) and Muirhead- Thomson (1951) are clas-

Sics, the former recently republished. Horsfall (1955) is a compilation use-

ful mainly for references to earlier literature. Marshall (1938) has enjoyed

great popularity as an elementary introduction and has also been recently re-

published. Natvig (1948) contains much information regarding northern spe-

cies. Clements pe covers a much wider spectrum than the title might sug-

gest. Mattingly (Riv. Parassit., in press) summarizes a wide range of topics

and has an extensive and up-to-date bibliography. Publications on bionomics

of individual vectors include Christophers (1960a), Jachowski (1954) and de

Meillon et al. (1967). Wharton (1962) covers various aspects of Old World

Mansonia spp. Literature on Neotropical species is scattered. Galindo et al.

(1950) and Trapido & Galindo (1957) will serve as an introduction.

The role of diel rhythms in cyclical behavior has received much atten-

tion but there is no general review. Mattingly (Trans. 13th internat. Congr.

Ent., in press) reviews some ecological implications. Haddow (1955) discus-

ses techniques for recording and analyzing biting cycles. More fundamental

work has been concerned chiefly with the oviposition cycle. Gillett (1962) is

_akey paper for the modus operandi of the internal 'clock'. More recently this

type study has been extended to the diel activity rhythm (Taylor & Jones,

1969).

Interest in mosquito genetics stemmed mainly from the development of

insecticide resistance but many other aspects of mosquito biology are now in-

volved. The field as a whole is well covered by Wright & Pal (1967).

GEOGRAPHICAL DISTRIBUTION

Outline distributions are given in the Synoptic Catalog. For further de-

tails see the various regional monographs listed below. Mattingly (1962a) dis-

cusses mosquito Zoogeography in general. For zoogeography of the Pacific

area see Belkin (1962, 1968).

4 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

FAMILY CULICIDAE

KEYS TO THE GENERA

I, FEMALES

1. Proboscis long, strongly attenuated and recurved

with prominent setulae confined to base; poste-

rior edge of wing emarginated just beyond tip

of vein Cu2 (Fig. 1) ee Toxorhynchitinae) 5

Bate es eas eh erg eae lalle Merle tke Se Toxorhynchites Theobald

Fig. 1. Toxorhynchites splendens (Wiedemann),

a. female head, b. male head, c. wing.

PYODOECIS ANG Wine) GLher wise’). Ki es ee Sel ladial iw Aa le Slee 2

2. Abdomen with sterna (and usually also terga)

wholly or largely devoid of scales (Subfamily

AQGONCLINGSTY a Wie Vie We Sie Ae as BUG ie yal we Legh a Oe Se Se ; 3

Abdominal terga and sterna with dense, uniform

covering of scales (Subfamily Culicinae)..........eee8-6 5

3. Veins Cul and M, distad of the cross vein, wavy

(Fig. 2); all wing scales dark; Moluccas, Melanesia

and northern Australia only. .......... Bironella Theobald

Fig. 2. Bironella hollandi Taylor, wing.

These veins very rarely wavy; if so then wing with

conspicuous pattern of light and dark scales!*.......... 4

4. Scutellum trilobed with bristles in 3 distinct groups

ve 3a); posterior pronotal bristles present

Fig. 4); New World tropics only. ......... Chagasia Cruz

Fig. 3. Thorax in dorsal view, a. Chagasia

fajardoi (Lutz); b. Anopheles balabacensis

Baisas; c. Wyeomyia aporonoma Dyar & Knab;

d. Trichoprosopon digitatum (Rondani). apn

anterior pronotum, p postnotum, s scutum,

sc scutellum.

Fig. 4. Generalized mosquito thorax in lateral

view showing setae used in the keys. 1 me

lower mesepimeral, pa prealar, ppv posterior

pronotal, psp postspiracular, sp spiracular.

Scutellum smoothly rounded with bristles more or

less evenly distributed (Fig. 3b); posterior pronotal

bristles absent ...... 7 a eae eee Anopheles Meigen

5. Tip of proboscis swollen, upturned and hairy (Fig. | 2.3.5

O): Old World tronics Only 5 eka nese ee Malaya Leicester“ °?

Fig. 5. Malaya genurostris Leicester,

female head.

* Superscript, here andelsewhere, refersto ''Notes onthe Keys" p. 29.

10.

12.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 3)

Proboscis sometimes with tip swollen,

otherwise unmodified . <2... 6 u'0 ee ew sty: T5350 A a Eth 6

Scutum with double median longitudinal stripe

of broad, flat, usually white or silvery, scales;

spiracular bristles present, postspiraculars

absent (see Fig. 4); Southeast Asia and New

Guinea. Only 54: eae a eee Topomyia Leicester 2? 3:9

Squama and postnotum bare; vein 1A reaching

wing margin at most very slightly beyond

base of fork ol vein - Cad Pie, Ga) 46 54k ale Hepa sik eel Wie 8

Fig. 6. Hodgesia malayi Leicester, a. wing,

b. outstanding scales from outer half of wing

field. al alula, c costa, sc subcosta, sq

squama, Sz? stem vein.

Squama fringed at least in part (Fig. 14a, b, d)

or postnotum with bristles (Fig. 3c, d) or vein

1A reaching wing margin well beyond the base

of fork of vein Cu (Fig. 13) or with any combi-

nation of these. characters: 444 ce seek ae woke OS Wee ROR 10

. Outstanding scales on outer half of wing field with

emarginated tips (Fig. 6b); Old World tropics

ONG ai: ove a te 0 RA ee wee eae SWS SG Hodgesia Theobald2

. Pleuron with conspicuous vertical stripe of broad,

silvery scales extending from the prealar area

to mid coxa (Fig. 7); wing membrane with micro-

trichia visible under magnification of about 50x;

Southeast Asia ‘OMty ys yi4 ce GE re wis Zeugnomyia Leicester2

Fig. 7. Thorax in side view, Zeugnomyia

gracilis Leicester.

Pleural ornamentation otherwise; microtrichia

minute, visible only under high mangification

5! RES ey FOP ee OA Re Ate Uranotaenia Lynch Arribalzaga2

Spiracular area with scales or one or more bristles

Be By eee Re GS as GO Wee tit ee alae 11

Spiracular area Dareg ace Wis Bea SE REI Ce i Se og REM eM oe 18

At most 1 or 2 prealar bristles present (see Fig. 4);

stem vein and base of subcosta without bristles

(see Fig. 6a); Oriental and Australasian Regions

and far eastern Palaearctic only...... Tripteroides Giles?» ©

Prealar bristles more numerous; stem vein, at

least, with bristles or genera confined to New

Wor la. 6 ek. eC Re ae ae we oe a Bee, 12

Postspiracular bristles absent; abdomen blunt tipped;

prealar bristles relatively numerous; bristles

present dorsally on stem vein and, usually, also

6 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

ventrally at base of subcosta; Nearctic Region

aad Old. World only. 6 ee oe Culiseta Felt4

Without this combination of characters; New

World species only .°3 0. a eee Se ee eee 13

13. Postspiracular bristles present; postnotum bare;

abdomen pointed at tip. .... ee Psorophora Robineau- Desvoidy

Postspiracular bristles absent; postnotum usually

with a tuft of setulae (Fig. 3c, d); abdomen

Dh Ee eo a a ee a 8 14

14. Spiracular area with broad scales only, without

bristles; proboscis shorter than fore femur;

scutal scales with gold and purple metallic re-

flections; pleura with abundant golden and sil-

very scales; hind tarsus with only lclaw .... Limatus Theobald

Spiracular area with 1 or more small bristles

(Fig. 4); proboscis and thoracic ornamenta-

tion various; hind tarsus with 2 claws as

WSUgE St eee Se ae SR eee ee rs 15

15. Antenna at most about half the length of the pro-

boscis, usually less; proboscis long and slen-

der, at least one-sixth as long again as fore

femur (as in Fig. 16c); scutum covered with

broad, flat scales; scutellum with silvery

scales at least on part of mid lobe; one or

more tarsi nearly always with white markings

o.one Side of least ..4..4 26.47 es Phoniomyia Theobald’

16. Scutum covered with flat, metallic scales with

bright iridescent reflection; prealar bristles

absent (see Fig. 4); anterior pronotal lobes

large, almost touching in mid line; one or

more tarsi often with conspicuous "paddles" 8

of erect scales (Fig. 8a). . 2. 505. Sabethes Robineau-Desvoidy

Fig. 8. a. mid leg, Sabethes belisarioi Neiva; b,

c. base of wing, b. Wyeomyia moerbista (Dyar &

Knab); c. 'Trichoprosopon pallidiventer (Lutz).

Without this combination of characters; tarsi

never with “paddies s 99s. 6 Oo) Se a a 17

17. Anterior pronotal lobes large and closely approx-

imated (Fig. 3c); squama bare or with 1-3

bristles or hairlike scales arising from the

upper portion near the alula (Fig. 8b); clypeus

sometimes with scales, never with setulae ... Wyeomyia Theobald

Pronotal lobes smaller, well separated (Fig.

3d) or squama with bristles more numerous

or arising from lower portion (Fig. 8c) or

clypeus with conspicuous setulae or with

any combination of these characters... Trichoprosopon Theobald

18.

19.

20.

21.

22,

23.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 7

Antenna short, thick, tapering, basal

flagellomere with a prominent scale

tuft; mid and hind femora with large

tufts of sub-erect scales at apex (Fig.

Gay CoN eee 5 ea a Sea ee. ad Aedeomyia Theobald

Fig. 9. Aedeomyia catasticta Knab, a.

female antenna, b. male antenna, c. hind

femur.

Antenna and-femora otherwise = oc 6 6 aw Bee ee eS 19

Antenna with flagellomeres short and thick

and verticillary hairs unusually short;

vertex and occiput with numerous short

hairs (Fig. 10a, b); New Zealand only. 2... ..4 4 « Opifex Hutton

Fig. 10. Head of Opifex fuscus Hutton, a.

female, b. male.

Head quite otherwise. 6°62 Seb sar a a ee ee 20

Postnotum with a small patch of setulae (as in

Fig. 3c, d) or scutum covered with flat scales

with bright metallic reflection, usually both;

southern and eastern Asia only. ........ Heizmannia Ludlow?

Postnotum without setulae or distribution other- i

WESC SoC eke a. ole Boe ce ai we ew a ee a ee

Scutum covered with broad, flat scales with

bright metallic reflection; anterior pronotal

lobes unusually large; New World tropics

and subtropics only. <2. ee soe SS es se Haemagogus Williston

Scutal ornamentation otherwise; anterior pro-

notal lobes not unusually daree? 2 ee i. we RE, 22

Antenna unusually long, exceeding the proboscis

by about the length of the last 4 flagellomeres

‘or more; first flagellomere greatly elongated,

3 or more times as long as the terminal fla-

gellomere (Fig. lla); proboscis not noticeably

swollen apically; New World only ...... Deinocerites Theobald

Fig. 11. Antenna, palp and proboscis,

Deinocerites cancer Theobald, a. female,

b. male; Galindomyia leei Stone & Barreto,

c. female, d. male; Ficalbia minima

Theobald, e. female.

Without this combination of characters’. . . ... 2.6 ee «6 Soa 23

Antenna longer than proboscis with all flagello-

meres markedly elongated, apical flagellomere

at least half as long as the basal one (Fig. 11c);

proboscis distinctly swollen apically; scutum

without conspicuous ornamentation; postspiracu-

lar bristles absent; tarsi unbanded; New World

tropics only. °. «a ew ee ee ee Galindomyia Stone & Barreto! 9

24,

20.

26.

27.

28.

29.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

Without this combination of characters . . .... «cece ee ec ae 24

First fore and mid tarsomere distinctly longer

than the other 4 together; fourth tarsomere

shorter than fifth, only a little longer

than broad (Fig. 15b); (postspiracular bris-

tles absent; all claws simple) ...... Orthopodomyia Theobald

ee 5

Proportions of these tarsomeres otherwise ......+...2.-.

Postspiracular bristles present or fore tarsal

ClAWS SOOEIEG OP OU 2 eo aac ese, Sah Ia) WH Reid te Ligh tb kon ee 26

Postspiracular bristles absent; all tarsal claws

SUC faa ce wok Coie leh ive Stee Lee ieee eniets 30

Paratergite broad and bare; postnotum usually

with a group of setulae (as in Fig. 3c, d);

back of head, pleura and posterolateral

corners of abdominal terga with broad, sil-

very scales; tropical Africa only. ..... Eretmapodites Theobald

Without this combination of characters ........ce-2-eee2ceccse 27

Decumbent scales of vertex broad, flat; post-

spiracular and lower mesepimeral bristles

present and proboscis curved and laterally

compressed (Fig. 12a) or postspiraculars ab-

sent and palpus half length of proboscis or more; 11

southern Asia, Japan and Melanesia only... Avrmigeres Theobald

Fig. 12. Armigeres (Armigeres) subalbatus

(Coquillett); antenna, palp and proboscis,

a. female, b. male.

Proboscis otherwise; postspiracular bristles present ....... 28

Upper surface of wing with all or most scales very

broad, many often asymmetrical (Fig. 13); all

tarsal claws simple; decumbent scales of vertex 12

HATLOW . es eee es iat COS Mansonia Blanchard (part)

Fig. 13. Mansonia (Mansonioides) uniformis

(Theobald); wing.

Without this combination of characters ... .6 <6 6 8 6 6 wile See 29

Squama bare or with at most 6 short hairs (Fig. 14a);

alula with broad scales; posterior pronotum with-

out scales; pleural scales restricted to 4 small

white spots; those on sternopleuron and mesepi~

meron sometimes fused; hind tarsomeres I-IV

with conspicuous white basal bands; Southeast

West ONLY 6 eo ee we eh ee pha alanine de es aca Udaya Thurman

Fig. 14. Base of wing, a. Udaya lucaris

Macdonald & Mattingly; b. Mimomyia

(Etorleptiomyia) luzonensis (Ludlow); c.

Mimomyia (Ravenalites) deguzmanae _

(Mattingly); d. Mansonia (Coquillettidia)

crassipes (Van der Wulp).

30.

31.

32.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 9

Without this combination of characters........ Aedes Meigen!®

Small or very small species; alula with fringe

of narrow scales; antenna with first flagello-

mere at least twice as long as fifth, usually

longer; palpus less than one-fifth length of

proboscis; scutellum with narrow scales

only; wing with anterior fork cell less than

twice as long as its stem; Old World tropics 14

OULY since ih eee ne ias a acl PR wa eaten cakte Ficalbia Theobald :

Alula bare or with flat, decumbent scales 14.15

(Fig. 14b, c); Old World tropics only .. eiccudeiees Theobald’ *?

Alula with narrow fringe scales (Fig. 14d). ......-2ec+s86«-+ 32

Hind tarsal claws very small and inconspic-~

uous; all tarsi with well developed La

villi (Fig. 15a) st tpl iis a ‘ pe ele Culex Linnaeus

Fig. 15. a. Culex sp., apex of hind tarsus;

b. Orthopodomyia anopheloides (Giles), mid

tarsus.

Hind tarsal claws not unusually small; 16

pulvilli absent os. 4: a «hg, wierewnd e805 Mansonia Blanchard (part )

I. MALES!”

Proboscis long, strongly attenuated and recurved

with prominent setulae confined to base; poste-

rior edge of wing emarginated just beyond tip

of vein Cu2; palps of the same order of length

as the proboscis (Fig. 1b, c) (Subfamily

Toxorbynechitinae) :¢ 0 wise gece nee oc9 Toxornynchites Theobald?

Proboscis and wing otherwise: WAIN VAViOUG fem ica: wiles exe & eros 2

. Abdominal sterna wholly or largely bare;

scutellum evenly rounded, not trilobed

(Fig. 3b); fore tarsus with a single claw only .......... 3

Without this combination of characters . 66 2. se © ecwier eens 4

. Veins Cul and M, distad of the cross vein, wavy

(Fig. 2); all wing scales dark; palps various;

Moluccas, Melanesia and northern Australia

CRs: eo ee. sce ee a eo cee as Bironella Theobald

These veins wavy only in ‘a few Species with 3

brightly ornamented wing’; palps at least

three-quarters of the length ee the proboscis,

usually longer... slay ieee aes ac arkansas da kuin Anopheles Meigen

Abdomen devoid of scales; fore tarsi with paired

claws; New World only ..... 2s ceeeeecceee biol Cruz

Abdomen densely SCalv i .arsoiw aie y 4 8 4 8 eo 8 wi eee 5

10.

AL,

12.

13.

. Proboscis strongly modified, as in the

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

female (Fig. 5); Old World tropics only . . Malaya Leicester“? 9#5

Proposcis other wises oo ps ee ee ee 6

Scutum with double median stripe of broad,

flat scales; spiracular bristles present;

postspiraculars absent; palps minute, as

in female; Southeast Asia and New Guinea

Os es 4, oi oa ee ee eee eles Topomyia Leicester

2; 3% 5

Without this combination of characters .......e+-e.-e-ceeccee 7

Outstanding scales on distal half of wing emargin-

ated at tips (Fig. 6b); antenna as in female,

non-plumose and with all flagellomeres, in-

cluding the last 2, subequal .......--- Hodgesia Theobald”

Wing scales otherwise; antennae variouS. .......s+se+2eeeee 9

Pleuron with conspicuous vertical stripe of

broad, silvery scales (Fig. 7); 1 fore and

Denied Claw LOOtheG os ss. ae ee ee Zeugnomyia Leicester

Pleuron otherwise; fore and usually also mid

claws both simple. ....... Uvanotaenia Lynch Arribalzaga” 18

Spiracular area with scales or 1 or more

bristles CRO I4) Ses oy See Pe ae UNS Sides 7 11

Spirecular area Dare so 0 ss ae ee ee a ee oe oP : 18

Prealar bristles numerous (Fig. 4); postspirac~

ular bristles absent; bristles present dorsally

on stem vein and usually also ventrally at base

of subcosta (see Fig. 6); palps at most slightly

shorter than proboscis; Nearctic Region and

Old World-only'. 66-5 Ee oe ee Culiseta “Felt*

Without this combination of characters an

GisStrTNliOn 6 ek Se ee he ge SS OE ag 12

Gl Worldonly: . 5 Pha Pe Se ee .... Tripteroides Giles?» ®

New Worle only eee se a eae ae ee Der er Be a eity Ba a Ege 13

Postspiracular bristles present; postnotum

bare; palps longer than proboscis . . Psorophora Robineau-Desvoidy

Postspiracular bristles absent; postnotum

normally with a tuft of setulae (Fig. 3c, d);

palps often only a quarter of the length of

the proposels Of 1686 555. ee Se as we ee 8 be 14

Mattingly: Illustrated Keys to the Genera of Mosquitoes

14. Proboscis shorter than antenna, and with

a conspicuous scale tuft at tip (Fig. 16a),

or with an abrupt flexure beyond half

way (Fig. 16b); palps minute as in female;

scutal scales with gold and purple metal-

lie TETeCUCIN bar ee a i OP eG gee Limatus Theobald

Fig. 16. Male antenna, palp and proboscis.

a. Limatus asulleptus (Theobald); b. Limatus

durhamii Theobald; c. Phoniomyia davisi

Lane & Cerqueira; d. Trichoprosopon

perturbans (Williston).

Proboscis otherwise; palps and scutal orna-

mentation various e e e e@ e e ® e e e e e e e e e e e e e e e e e@ 6

15. Palps about a quarter of the length of the

proboscis or less; antenna at most about

half the length of the proboscis, usually

less; proboscis long and slender, at least

about one- sixthas long again as fore femur

(Fig. 16c); scutum covered with broad,

flat, often metallic scales; scutellum with

silvery scales on part of mid lobe at least;

one or more tarsi nearly always with white

narkines . 65.5 Soe . 2... Phoniomyia Theobald!®

Without this combination of characters. .......eeeeeccee

16. Palps less than a quarter of the length of

the proboscis; scutum with broad scales

with bright, iridescent, metallic reflec-

tion; one or more pairs of legs often with

conspicuous "paddles" (as in Fig. 8a);

prealar bristles absent (see Fig. 4); ante-

rior pronotal lobes very large, almost

touching in mid line 6. ess ass ‘ Sabethes Robineau-Desvoidy20

Without this combination of characters;

legs never with "paddles" ......4.6- Pere Ga ee las Sees «

17. Palps usually more than half the length of

the proboscis; if not then either scutum

with narrow scales or proboscis long and

slender and antenna much more than half

the length of the proboscis with the 2 ter-

minal flagellomeres greatly elongated

(Fi. FOGG). 2 eS, «Ghee ee Ex ee

Palps at most about a quarter of the length

of the proboscis, usually less; scutum

covered with broad, flat scales; antennae

various but never with the 2 terminal

flagellomeres greatly elongated. ..... Wyeomyia Theobald

18. Antenna with the 2 terminal flagellomeres

markedly thickened (Fig. 9b); hind femur

with a large apical scale tuft (Fig. 9c);

Trichoprosopon @icobald

19, 20

palps very short, as in female. ........ Aedeomyia Theobald

Antenna and hind femur otherwise; palps

VATIONS SS Se ee Re Soe ee Se OO eee eee

12 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

19. Antenna with flagellomeres 2-4 each with a

stout dorsal spine; back of head with nu-

merous short hairs: palps with ce ela-

vate (Fig. 10b); New Zealand only . Be ES Opifex Hutton

Antenna, palps and back of head quite

OUST WIDE. os ae ee gi REE ete Se ate a te eee we OE 20

20. Palps at most about one-fifth of the length

of the proboscis; scutum covered with

broad, flat scales with bright, metallic

reflection or postnotum with a group of

setulae (as in Fig. 3c, d) or both; an-

tenna usually with verticillary hairs short

as in female (Fig. 17a); southern and 29

Caetera ARs ONIVS “ok Sf oe eS Heizmannia Ludlow

Fig. 17. a. Heizmannia scintillans Ludlow,

male antenna, palp and proboscis; b. Galindo-

myia leei Stone & Barreto, male fore tarsal

claws.

Without this combination of characters....... Pete? Rubee me ts tae 21

21. Antenna with basal flagellomere (and some-

times also some succeeding flagellomeres)

greatly elongated, at least 3 times as long

as the terminal flagellomere; verticillary

hairs short as in female; proboscis not no-

ticeably swollen apically Fig. 11b); New

WOOP I On ee yy ae Ss a Se PS Deinocerites eke

22, Palps very short as in female; antennal

flagellum with verticillary hairs short

and scanty; apical flagellomere at least

half as long as the basal one; proboscis

distinctly swollen apically (Fig. 11d);

inner claw of fore tarsus with 2-5 teeth

arising from the convex surface (Fig. 17b);

other claws all simple; New World tropics

GRE ee Be ca ee .. Galindomyia Stone & Barreto!9; 23

Without this combination of characters ES So eet er es Bee ae Pe ene 23

23. Scutum covered with bright, metallic,

iridescent scales; anterior pronotal

lobes unusually large (see Fig. 3); palps

at most two-thirds of the length of the

proboscis, often much less; New World

tropics and subtropics only ........e. oii hts Williston

Without this combination of characters. ....... oe eet ihe 2

24. Postspiracular bristle or bristles present ........2.e-e-ee-e 25

Postspiracular area without bristles .......2.e.seeceseee 29

Mattingly: Illustrated Keys to the Genera of Mosquitoes 13

25. Paratergite broad and bare; postnotum

usually with a group of setulae (as in

Fig. 3c, d); back of head, pleura and

posterolateral corners of abdominal

terga with conspicuous patches of broad,

silvery scales; tropical Africa only... . Pictimpireles Theobald

Without this combination of characters. ......-+..-e-e008 26

26. Decumbent scales of vertex broad, flat; post-

spiracular and lower mesepimeral bristles

present; acrostichals and dorsocentrals ab-

sent; proboscis curved and laterally com-

pressed (Fig. 12b); southern Asia, Japan

and Melanesia only 9 Sc Gale eae Te Armigeres Theobald (part)

Without this. combination of characters ..2:+:s s+ + 0. «s-« 6's»

27. Upper surface of wing with all or most scales

very broad, many often asymmetrical (Fig.

13); species confined to New World or if

occurring in Old World then with apical

segment of palp greatly reduced (Fig. 18a)

dig siecle Rie Rp pas ee es Mansonia Blanchard (part) 4

Fig. 18. Male palps and proboscis, a.

Mansonia (Mansonioides) uniformis (Theobald);

b. Ficalbia minima Theobald; c. Mansonia

(Coquillettidia) crassipes (Van der Wulp).

Wing seldom with scales of this type and then

only in Old World species; the latter with

apical and subapical segments of palps either

both well developed or both greatly reduced. .......... 28

28. Palps long, slender, almost hairless, the 2 ter-

minal joints together more than half the length

of the shaft; squama bare or at most with 6

short hairs; alula with broad scales; poste-

rior pronotum without scales; hind tarsomeres

II-IV with conspicuous white basal bands;

Southeast Asia only. ........ er ara Udaya Thurmgp

Without this combination of characters . bitin Titel ae eam Aedes Meigen

29. Alula bare or with flat, decumbent scales 14.15

| (Fig. 14b, c); Old World tropics only . Mimomyia Theobald” “

Alula with narrow fringe scales (Fig. 14d) . Sait: yokes Gis Sie Ree 30

30. Proboscis greatly swollen on distal third

or more (Fig. aa Old World iunaniod 14

only. .......2...-. bs gis Ficalbia ecg

31. Pulvilli present (best seen on hind legs);

hind tarsal claws unusually small (as in

Fig. 15b); palps various, in some cases

very much shorter than proboscis. ......... Culex Linnaeus

Pulvilli absent; hind claws not unusually

small; palps always at least as long or

aAlmost.ag lone GS DTOOOSCIS ..4: so <n we 8 we wee Pr roe ee 32

14 | Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

32. Palps longer than proboscis, the terminal

segment not much shorter than the sub-.

terminal; subterminal segment and tip

of shaft with numerous long hairs (Fig.

PGC) oa oe es ee Pra: Mansonia Blanchard (part)26

Palos Ou erwise sc. 4 ub cae ee a 6 WL ee we Fe Ca 33

33. Palps with the last 2 segments not greatly

reduced (Fig. 19b); southern Asia 97

ONY. ai cece os os ae a ee Go CArmigeres. Theobald < (part)

Fig. 19. Male palps and proboscis, a. Culex

postspiraculosus Lee; b. Armigeres (Leicesteria)

(ey. Barraud; c. Orthopodomyia anopheloides

Giles).

Palps with the apical segment (and some-

times also the subapical) greatly re- 29

duced (91g. 19C) 0 se Sg ee ee Orthopodomyia Theobald

Tl. PUPAE

The pupa of genus Galindomyia is undescribed.

Fig. 20. Generalized mosquito pupa. as Apical

paddle seta, fh Float hair, p Paddle, /7v

Trumpet.

1. Segment X with a conspicuous branched

hair; seta 9 on segment VIII greatly re-

duced; paddles without apical seta (Fig. 5

21a) (Subfamily Toxorhynchitinae).... Toxorhynchites Theobald

Fig. 21. Terminal segments of pupa. a.

Toxorhynchites splendens (Wiedemann), b.

Anopheles balabacensis Baisas, c. Chagasia

bathana (Dyar).

Segment X without setae; seta 9-VIII and

paddle various ..-.--.+.-+-e- oa Ge ee Cea few 2

2, Paddles nearly always with an accessory seta

arising anterior to and in line with the api-

cal seta (Fig. 21b, c); if not then seta 9 on

segments IV-VI in the form of a short,

stout, dark spine arising from the extreme

posterior corner of the segment (as in Fig.

21b) and trumpets short, flared and split

nearly to base (Subfamily Anophelinae)“? ..........6.2. 3

Paddles with accessory seta absent or, if

present, arising level with and laterad of

the apical seta; seta 9 usually and trumpets ,

almost always otherwise (Subfamily Culicinae) ......... 4

Mattingly: Illustrated Keys to the Genera of Mosquitoes 15

3. Seta 2-III-VII a short, stout, dark spine

(Fig. 21c); New World tropics only......... Chagasia Cruz

Seta 2 on these segments otherwise ........ Anopheles Meigen

Bironella Theobald

Trumpets modified for insertion into sub- |

aqueous plant tissues (Fig. 22a, b); 30

float ‘hair Suppressed 2240 e oe oe es Mansonia Blanchard

Fig. 22. Pupal trumpets. a. Mansonia

Mons uniformis (Theobald), b.

ansonia (Coquillettidia) microannulata

(Theobald), c. Hodgesia malayi Leicester,

d. Topomyia spathulirostris Edwards, e.

Topomyia tipuliformis Leicester, f.

Tripteroides lorengaui Peters, g. Tripter-

oides fuscipleura Lee.

Trumpets otherwise or float hair well developed

OF DOU 6 ase a See ke: sol SR a eg, Se eh, oe at a ene 9)

Trumpets with a hinged tragus (Fig. 22c)..... Hodgesia Theobald

Trumpets otherwise ....... Se ie ee ee a a Oe ay ot 6

Paddles small, usually more or less

pointed, without apical seta (Fig. 23)

(Tribe Sabethinig) soe 2 eS Sie gee a ms eg a ae i

Fig. 23. Sabethine pupal paddles. a. Topomyia

decorabilis Leicester, b. Topomyia barbus

Baisas, c. Tripteroides stonei Belkin, d.

Limatus durhamii Theobald, e. Phoniomyia

pallidoventer Theobald, f. Tvichoprosopon

magnum (Theobald).

Paddles ovine pers nearly always with

apical setart cee sk ey 3 eR RR A SHE Be tees 4 14

oA World only: eas a ee PPO eae 6a RNIN 9 ag 8

New: World. onlys>. ese eSNG OP OEE RE OMG AY ptt tag tay glia Aeneas 10

Trumpets subcylindrical with inner and

outer walls widely separated (Fig. 22d, e).......2.-e ese 9

Trumpets subconical or with inner and

outer walls closely apposed or both 9

Pe 2B eye eres: 4 Sag et gies Satchel gan ah eg Tripteroides Giles?

Seta 6-VII relatively well developed,

eae. well cephalad of seta 9-VII

(Pig, ZAG). 6 ie eck oe Pe Re 8 ae ie rn Malaya Leicester

Fig. 24. Segment VII of pupal abdomen. a.

Malaya genurostris Leicester, b. Topomyia

spathulirostris Edwards.

Seta 6-VII usually less well developed and

arising close to and laterad of seta 9-VII 3

CTA an a ek ee OE RS Oe Topomyia Leicester 3

16 Contrib, Amer. Ent. Inst., vol. 7, no. 4, 1971

10. Paddles short and broad, much shorter than

seta 9-VUI, with edges sometimes ser-

rated but tips entirely bare; posterior

border of segment VIII deeply excavated;

trumpets narrow, cylindrical or sub-

conical { Figs, -23d, 20a) ese wie! aot mae niu Limatus Theobald

Fig. 25. Sabethine pupal trumpets. a.

Limatus durhamii Theobald, b. Phoniomyia

edwardsi Lane & Cerqueira, c. Tvichoprosopon

soaresi Lane & Cerqueira, d. Sabethes

soperi Lane & Cerqueira, e. Tvichoprosopon

magnum (Theobald), f. Wyeomyia circumcincta

Dyar & Knab.

Without this combination of characters. ........e-+.-+-scecee 11

11. Trumpets slender, tubular, narrowing

basally, not or only slightly expanded

at apex (Fig. 25b); paddles broad with

apex pointed and usually spiculate (Fig. 34

BOG tei ck em ko, a eh GS Steen Phoniomyia Theobald

Trumpets broader, usually conical or

beaker- shaped with basal portion ex-

panded and inner lining well separated

Be. eG. oO CS aoa ee Se SE bee ai Ad beeieen 2 eee 12

12. Either with long, delicate fringe on both

borders of paddle (Fig. 26a) or with

genital sac sunk in a deep embrasure 35

CP DGD) ae eee ENN Ee BS Wyeomyia Theobald (part)

Fig. 26. Posterior segments of pupal abdomen.

a. Wyeomyia felicia (Dyar & Nunez Tovar), b.

Wyeomyia codiocampa Dyar & Knab, c. Sabethes

purpureus (Theobald), d. Trichoprosopon

soaresi Lane & Cerqueira.

Paddles without such a fringe; posterior

border of segment VII with at most a

Bie low @xeavelion (Pigs COC, Ue SOV kk eke See ee ee ee 13

13. Seta 5-VI longer, usually much longer, than

segment VIL; seta 9-VI variously devel-

oped but always conspicuous (Fig. 26c)

Pena ke oe Sabethes Robineau-Desvoidy36

Wyeomyia Theobald (part)?°

Seta 5-VI less strongly developed, often

shorter than segment VI; seta 9-VI

very small and inconspicuous (Fig. 36

Boy OU rays Sa ee eS Trichoprosopon Theobald,

Wyeomyia Theobald (part)?°

14. Paddles smooth on both borders with apex

convex; apical paddle seta at least two-

thirds as long as paddle; seta 9- VIII long,

single, simple (Fig. 27a); New World

ORES PSone Se Ska eo Se ek ee eo ee Deinocerites Theobald

15.

16.

a1.

18.

19.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 17

Fig. 27. Pupal abdomen. a. Deinocerites

mcdonaldi Belkin & Hogue, b. Aedeomyia

catasticta Knab, c. Opifex fuscus Hutton.

Without this combination of characters .......2se-e«-ce-rceceree 15

Paddle smooth on both borders, deeply cleft

at apex; apical seta at least half as long

as paddle; seta 5-IV-VI with long, frayed

median branch and short lateral branches

(Fic, Bie «ewan ate |

Paddles and seta 5-IV-VI otherwise ......2-s-2eseeceevee

Seta 9-VII very short, single, simple as

on anterior segments (Fig. 27c); New

on anterior segments or distribution

otherwise or both. :.y)-o 16 peieha eee tal i whe wi eilericie ree beie ies 17

Trumpets at least about 10 times as long

as their breadth at half way, usually

more; paddles narrow or very narrow,

not or only very slightly inflated on inner

aspect; fringe, usually of irregular spic-

ules, on both borders; apical seta minute

or absent (Fig. 28); Old World tropics 14. 15. 37

OLY. Gate WI Ra Is aha ee Cale Mimomyia Theobald” "’~*?

Fig. 28. Pupal trumpets and paddles.

Genus Mimomyia. a. M. perplexens

(Edwards), b. M. hybrida (Leicester),

c. M. chamberlaini Ludlow, d. M.

(Etorleptiomyia) luzonensis (Ludlow),

e. M. (Ravenalites) deguzmanae

(Mattingly).

Without this combination of characters . ....-+.-«-e-eseeseee 18

Paddles with long, delicate fringe on both

borders; float hair arising unusually near

mid line, tending to point forward in

mounted specimens; seta 1-II long, stout,

single or bifid; seta 5-III- VII in each case

longer than the following segment (Fig.

29); Southeast Asia only ..... bial Zeugnomyia Leicester

Fig. 29. Pupal abdomen. Zeugnomyia

lawtoni Baisas.

Without this combination of characters... . e+e see ee ees 19

Paddles with inner half deeply excavated

towards base, usually much broader than

outer half; segment IX usually with a pair

of small setulae; paddles fringed or

toothed on both borders, usually exten-

Sively.@o' (Pig. GQ) 4 sk 4 44 Uvranotaenia Lynch Arribalzaga®8

18 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

Fig. 30. Terminal segments of pupa. Genus

Uranotaenia. a. U. mendiolai Baisas, b. U.

ascidiicola De Meijere, c. U. modesia Martini.

Without this combination of characters. ...... a Gilligan, ae tip ar 20

20. Paddles small, with long, delicate fringe

on both borders; apical paddle seta long

and stout; seta 8-C much longer and

stouter than 9-C (Fig. 3la, b); tropical

INEGICAOUTY 5 os 6s ee ee ae ae Evretmapodites Theobald?9

Fig. 31. Terminal segments and cephalo-

thorax of pupa. a. Evetmapodites inornatus

Newstead, b. Evetmapodites dracaenae

Edwards, c. Aedes Stegomyia) ruwenzori

Haddow & Van Someren.

Without this combination of charactersanddistribution,....... 21

21. Paddles with long, delicate fringe on both

borders; seta 6-VI very strongly devel-

oped, longer and much stouter than 5-VI

(nearly always spinose or multibranched,

often subplumose); seta 9-VI a minute,

colourless setula (Fig. 32a, b); southern 4

Asia, China, Japan and Melanesia only... Armigeres Theobald 0

Fig. 32. Terminal segments of pupa. a.

Armigeres subalbatus (Coquillett), b.

Armigeres malayi (Theobald), c. Heizmannia

achaetae (Leicester).

Paddles without such a fringe or segment

VI otherwise; distribution various. ........45e-e+s2ee0e-68 : 22

22. Paddles with long, delicate fringe on both

borders; mid rib of paddle very poorly

developed, barely visible, if at all,

even towards base; Southeast Asia only

aoe ee oe eae. ee eS ae as es Udaya Thurman

Fig. 33. Terminal segments of pupa. a.

Udaya argyrurus (Edwards), b. Aedes

Stegomyia) meronephada (Dyar & Shannon),

c. Aedes Stegomyia) annandalei (Theobald),

d. Heizmannia scintillans Ludlow.

Paddles various, if with long fringe then

mid rib always strongly developed, con-

Serclots (ae oe eS ee es aces 6 ee pepe phage 23

23. Paddles with long, delicate fringe on both

borders, oval, usually more or less

pointed, never indented at tip; seta 9-VI

minute, colourless (Fig. 33d) or if not

then either seta 5-II long, stout, dark

Mattingly: Illustrated Keys to the Genera of Mosquitoes

(Fig. 34a) or seta 5 on all segments very

short (Fig. 34b); southern Asia only. .... Heizmannia Ludlow

Fig. 34. Pupal abdomen. Genus Heizmannia.

a. H. complex (Theobald), b. H. aureochaeta

(Leicester).

Without this combination of characters ......+«+s-e+2esceec2ecee

Fig. 35. Terminal segments of pupa. a.

Aedes (Lorrainea) fumidus (Edwards), b.

Ae. (Diceromyia) franciscoi Mattingly, c.

Ae. (Diceromyia) periskelatus (Giles).

Fig. 36. Pupal abdomen. Aedes subgenus

Stegomyia, a. Ae. albopictus (Skuse), b.

Ae. desmotes (Giles).

24. Trumpets at least 7 times as long as their

breadth at half way, the pinna occupying

at least half the length, meatus entirely

tracheoid or almost so; seta 9- VIII very

feebly developed; paddle edge strongly

serrated on outer half, inner half smooth

(Fig. 37); Old World tropics only. ....... Ficalbia Theobald

Fig. 37. Pupal trumpet and terminal segments.

Genus Ficalbia. a. F. circumtestacea (Theobald),

b. F. malfeyti (Newstead).

Without this combination ‘of characters 2°. 60 6 SOU ai oe eee a

25. Paddles more or less rectangular with

thickened basal portion of outer edge

sometimes spiculate but whole border

otherwise smooth, hyaline; accessory

paddle seta absent; apical paddle seta

very short; seta 9- VI-VII long, stout,

plumose, on VII about half the length

of the paddle or more (Fig. 38) .... Orthopodomyia Theobald

Fig. 38. Pupal cephalothorax and terminal

segments. Genus Orthopodomyia. a. O.

flavicosta Barraud, b. O. wilsoni Macdonald.

Without this combination of characters . ......«-.+seseces

26. Trumpets with tubular portion occupying most

of the length and with rudimentary basal

tracheation at most; seta 9-VIII arising

from the posterolateral corner of the seg-

ment, not, or only very slightly, displaced

anteriorly; either with posterior corner of

abdominal segment IV toothed or with prom-

inent ventral lobes on posterior border of

segment VIL, partly covering the bases of

the paddles, or with accessory paddle seta

present (Fig. 39); New World only. Psorophora Robineau- Desvoidy

20 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

Fig. 39. Pupa. Genus Psorophora. a. Cephalo-

thorax and abdomen, Ps. ciliata (Fabricius), b.

Segment IV of abdomen, Ps. ferox (Humboldt)

c. Segments I-Il, Ps. infinis (Dyar & Knab) 43

Without this combination of characters ........see-sesece at

27. Seta 8-C arising anterior or at most slightly

posterior to base of trumpet, very much

anterior to 9-C; trumpet nearly always

with rudimentary basal tracheation at

most; seta 9- VIII rarely arising cephalad

of the posterior border of the segment

a rhs la ocd 6 > Rok es eee a Seek’ A eat ay ape 28

Fig. 40. Pupal cephalothorax and abdomen.

a. Haemagogus spegazzinii Brethes, b.

Haemagogus capricornii Lutz, c. Aedes

aegypti (Linnaeus).

Seta 8-C arising level with or posterior to

the base of the trumpet, more nearly

level with 9-C; trumpets frequently with

extensive subbasal tracheation; position

of seta 9- Vill various. (Fig, 4)bjewis aosjios Hie es ee 29

Fig. 41. Pupal cephalothorax and abdomen. a.

Culex pipiens Linnaeus, b. C. antillummagnorum

Dyar, c. Culiseta longiareolata (Macquart).

28. Setae 8-C and 9-C poorly developed; setae

\)-II and 5-III very feebly developed, not

or barely reaching onto the following seg-

ment; either with seta 5-VII as long as or

longer than the following segment or with

mid rib of paddle deeply pigmented and

seta 5-IV-VI shorter than the following

segment; seta 9-III-VI minute; seta 9-VII

with 4 or more branches, about half the

length of the paddle or more (Fig. 40a, b); “a

New World tropics and subtropics only .. Haemdgogus Williston

Without this combination of characters or 45

distribution. otherwise: .tessieoeGo awe Suess. Aedes Meigen

29. Trumpets with well developed subbasal

tracheation or seta 9- VII arising well

cephalad of the posterior border of the

segment, usually both (Fig. 4la, hb,

BRO) oe etka ie at RM APO a tieexe fe Culex Linnaeus*®

Fig. 42. Pupal cephalothorax. a. Culex

davisi Kumm, b. Culex bamborum Rozeboom

& Komp, c. Culex pseudomelanoconia

Theobald, d. Aedes auvantius (Theobald),

e. Aedes longirostris (Leicester), f. Aedes

gilli Barraud.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 21

Trumpets with rudimentary basal trachea-

tion at most; seta 9- VII always arising

from the posterior border of the seg- 4.46

ment (Fig. Ateys es) saa eee eee SR 3 ae Culiseta Felt”?

IV. FOURTH STAGE LARVAE

The larva of genus Galindomyia is undescribed.

Respiratory siphon absent; seta 1 usually

palmate on most abdominal segments

(Fig. 43) (Subfamily Anophelinae) .........-. a alas aka 2

Fig. 43. Terminal segments of anopheline

larvae. a. Chagasia bathana (Dyar), b.

Anopheles balabacensis Baisas.

Siphon present; seta 1 never palmate ......+ 2. eee eeees 4

Anterior flap of spiracular apparatus produced

into along, spinelike process, ventral valves

with fringe offine hairs; palmate hairs character-

istically shaped (Fig. 43a); New World tropics only Chagasia Cruz

Spiracular apparatus and palmate hairs other-

wise e e e e e e e e e e e e e e e e e e e e e @ e e ® e e e e e e e 3

. Inner clypeal setae close together; seta 1

of mesothorax palmate (Fig. 44); northern

Australia, Melanesia and Moluccas only. ... Bironella Theobald

Fig. 44. Larval head and thorax. Bironella

hollandi Taylor. icl Inner clypeal setae.

Without this combination of characters and

distripution i's 2s Ciera a Gate Anopheles Meigen

Mouthbrushes with about 10 flattened, non-

pectinate blades; antenna with setae 2-A

and 3-A arising basad of seta 1-A; comb

and pecten absent (Fig. 45) (Subfamily 5

Toxor hynchitinge ic: Ws ate. .clss SHU Be Toxorhynchites Theobald

Fig. 45. Larval head and terminal segments.

Toxorhynchites splendens (Wiedemann).

Mouthbrushes with numerous hairs; antenna

with setae 2-A and 3-A distad of 1-A;

comb almost always present; pecten

present or absent (Subfamily Culicinae) ...... ++ eeees ee

. Ventral brush with at most 2 pairs of setae,

OT a eee ne re ee ei cso, 6

22 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

Fig. 46. Terminal segments of New World

sabethine larvae. a. Sabethes purpureus

(Theobald), b. Phoniomyia fuscipes

(Edwards). vbr Ventral brush, Ics

Lower caudal seta.

Ventral brush with 3 pairs of setae or more

or speetes contined to Old World... 00. 6 sie ee oe es 10

6. Setae of ventral brush as long, or almost

as long, as lower caudal setae; siphon

relatively slender, at least about 3.5

times as long as saddle; comb teeth in

a single row or with at most 3 or 4 de-

taened (fie.. 468) . 6 wa ee ek es Sabethes Rr

Wyeomyia Theobald (part)4#

setae of ventral brush much shorter than

lower caudal setae or siphon short and

stout or comb teeth in at least 2 com-

plete rows or with any combination of

PHO OR re ae ye ee a ee 7

7. Siphon long, slender, strongly tapering,

about 5 times as long as its breadth at

base or more, with numerous long, un-

branched setae dorsally and ventrally;

comb teeth in 2 or more rows (Fig. 46b). . Phoniomyia Theobald

Wyeomyia Theobald (part)49

Siphon otherwise or comb teeth in a

single, regular row (Sometimes aris-

ing from a sclerotized plate) or both. ....... ox er Ge: 8

8. Siphon short, stout, at most about 3 times

as long as its breadth at base with

several branched setae dorsally and

ventrally; head setae 4-C, 5-C and 6-C

single; maxillary "horns" not developed;

comb with about 4-7 teeth in a single

row, not arising from a sclerotized i

plate (Fico 47). ecw OU eee — Limatus Theobald

Fig. 47. Larval head and terminal segments.

Limatus durhamii Theobald. mx Maxilla.

Without this combination of characters;

maxillary "horns" present in some spe-

Clee re es ee a Kk we area 9

Fig. 48. Larval head and terminal segments.

Trichoprosopon frontosum (Theobald). mx

Maxilla.

9. Mandible greatly enlarged (Fig. 49a) or

maxilla with a large "horn" (Fig. 48);

siphon with a dense midventral row of

setae extending for almost the whole

length (Fig. 48) or setae of ventral brush

at least 3 times as long as saddle .... Trichoprosopon Theobald

Mattingly: Illustrated Keys to the Genera of Mosquitoes

Fig. 49. Larval head and terminal segments.

a. Tricnoprosopon digitatum (Rondani), b.

Wyeomyia confusa (Lutz). md Mandible,

mx Maxilla.

Mandibles never thus; maxilla seldom with

conspicuous "horn", if so then ventral

brush or siphon otherwise. ..... Wyeomyia Theobald (part

10. Ventral brush with a single pair of setae

(1 or 2 small, supplementary hairs

present in occasional individuals); an-

tenna short, without articulated apical

segment; siphon with 2 or (usually)

more subdorsal setae as well as vari-

ous ventral or subventral setae (Fig. 50). .........

Fig. 50. Larval head, prothorax and terminal

segments. a. Malaya genurostris Leicester,

b. Topomyia gracilis Leicester.

Without this combination of characters?+ ...........

11. Prothoracic setae 5 and 6 large fan- shaped

tufts arising from a common tubercle;

comb usually a patch of teeth in 2 or

more rows (Fig. 50); seta 6 of meso-

thorax and seta 7 of metathorax never

stout spines; tropical Africa, southern

Asia and Melanesia only: . we ee eee eh we te ee

Fig. 51. Larval head and terminal segments.

a. Mimomyia (Ravenalites) deguzmanae

(Mattingly), b. Culex (Acallyntrum) belkini

Stone & Penn.

Prothoracic setae otherwise; seta 6 of meso-

thorax and/or seta 7 of metathorax often

a stout spine; comb teeth in a single row,

‘sometimes arising from a sclerotized

plate, occasionally absent or reduced to

a single tooth (Fig. 52), or distribution

Gther wise... 6 SF ete ae eet Tripteroides Giles

Fig. 52. Head, thorax and terminal seg-

ments of larva. Genus Tripteroides. a.

T. powelli (Ludlow), b. T. stonez Belkin.

12. Either abdominal segments IV-VI (at least)

with 1 or more pairs of stellate setae

with numerous short, stiff branches

(Fig. 50b, 53a) or maxillae with con-

spicuous "horns" (Fig. 53b) or siphon

at least 6 times as long as saddle (Fig.

53c); southern and eastern Asia and

Melanesia only -. ... 2s eee ee Gar Topomyia Leicester

52,53

13.

14.

15.

16.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

03. Genus Topomyia. a. Segment V

= larval abdomen. T. tenuis Edwards, b.

Larval head., T. decorabilis Leicester,

c. Terminal segments of larva. T.

spathulirostris Edwards. mx Maxilla.

Setae of this kind never present; maxillae

never with "horns"; siphon at most about

4 times as long as saddle (Fig. 50a);

Old World tropics from Africa to eastern

Asia and Melanesia: 2) 60s wicegew wie ais Malaya “Leicester

Siphon modified for piercing plant tissues,

with sclerotized saw-toothed process

Se PE OSE) os a os CE Ee. ee, BS Mansonia Blanchard

Fig. 54. Terminal segments of ee a.

Mansonia uniformis (Theobald), b

Mimomyia hybrida (Leicester).

Siphon not so modified or, if so, without,

any saw-toothed process (Fig. 54b)°*..........020000. 14

Antenna broad, flattened; some thoracic

setae enormously long, others stellate;

tip of siphon with paired hooks and

branched: setae (Fie. 55): gos ee 8 Aedeomyia Theobald

Fig. 05. Larval head, thorax and terminal

segments. Aedeomyia catasticta Knab.

Antenna, thoracic setae and siphon otherwise. .........s.e-. 15

Siphon with a single pair of subventral

setae; metathoracic setae 9-12 very

short, unbranched (Fig. 56); New

Zealand OR a ee PR a Ce ee A tS Opifex Hutton

Fig. 56. Larval thorax and terminal seg-

ments. Opifex fuscus Hutton.

Without this combination of characters

or distribution otherwise ...... Sd tn a al ei Sa ay : 16

Siphon with a single pair of subventral

setae arising at not more than one-fifth

of the distance from base to apex; comb

a single row of at most 20 teeth; siphon

less than twice as long as saddle (Fig.

57); Old World tropics only. .... ake A PENTA.) 4 BET, 17

Fig. 57. Larval head and terminal seg-

ments. a. Hodgesia malayi Leicester, b.

Ficalbia minima (Theobald).

Without this combination of characters

or distribution otherwise 9%........ ee UU Gt bh ie tug b gs SUS a ke 18

18.

19.

20.

21,

Mattingly: Illustrated Keys to the Genera of Mosquitoes 25

Head seta 5 arising almost directly behind

6 which is single; head seta 4 nearly

as long as 5; pecten with at least 3

teeth, usually more (Fig. 57a) ....-.-..--s Hodgesia Theobald

Head setae otherwise; pecten with at most

2 teeth Fics S7b)ia v4 ow wba 2.5 al Ficalbia Theobald°®

Distal portion of antenna freely articulated

(Fig. 58); siphon with a single pair of

subventral setae; pecten with at most

4 teeth on either side, often fewer;

ventral brush with 2-4 pairs of setae,

1 or 2 supernumerary setae occasion-

ally present in addition; Old World 14.15.54

tropics ONY 646 ea: x le Re So Mimomyia Theobald’? ~~?

Fig. 58. Larval head. Genus Mimomyia.

a. M. (Mimomyia) chamberlaini Ludlow,

b. M. (M.) plumosa (Theobald), c. M.

(Ravenalites) deguzmanae (Mattingly).

Antenna never thus; siphon often otherwise;

ventral brush with at least 3 pairs of

setae, often with 5 or more pairs ......64-+522e ee eee 19

Pecten absent; siphon very short and broad

with a single pair of subventral setae

arising beyond half way; antennal seta

minute (Fig. 59a); southern Asia, Japan

and Melanesia only .......-. + ere 2% Armigeres Theobald?

Fig. 59. Larval head and terminal segments.

a. Armigeres subalbatus (Coquillett), b.

Orthopodomyia wilsoni Macdonald.

Without this combination of characters

and distribution 2. aoncmaies ese mie iaoerinndils wie mak was 20

Pecten absent; antennal seta arising on

basal half, with 4 or more branches;

head setae 5 and 6 long and branched;

siphon at least about 2.5 times as

long as its breadth at base, often much

longer, with a single pair of subventral

setae; ventral brush with 6 pairs of 57

setae or more (Fig. 59b). . «0 9s « Orthopodomyia Theobald

Without this combination of characters ....... ee aa ee 21

Head with a pair of conspicuous lateral

pouches; siphon with a pair of large

subventral setae and 2 pairs of smaller

setae distal to this, 1 subdorsal, the

other subventral; saddle poorly devel-

oped; a small accessory sclerotized

plate usually present basad of the ven-

tral brush (Fig. 60a); New World only. ... Deinocerites Theobald

22.

23.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

Fig. 60. Larval head and terminal segments.

a. Deinocerites cancer Theobald, b. Culiseta

longiareolata (Macquart), c. Culiseta littleri

(Taylor).

Without this combination of characters ......e«.e eee es

Siphon with a single pair of subventral setae

arising at not less than a quarter of the

distance from base to apex, usually more;

(a pair of minute subdorsal setae also 58

usually present near tip) (Figs. 62-65)°°. ....... ee.

Siphon with subventral setae more numerous

or (in 1 or 2 neotropical species) entirely

absent or if with a single pair of such

setae then these arising at about one-

fifth of the distance from base to apex

or lesé (Pigs. GOB 0a) ea ee EY Se

Maxillary suture absent or incomplete,

not reaching posterior tentorial pit

(Fig. 61a); head seta 5 or 6 or both

often flattened, barbed, spinelike;

comb often arising from a large scle-

rotized plate (Fig. 62a). ... Uvanotaenia Lynch Arribalzaga

Fig. 61. Larval head in ventral view. a.

Uranotaenia sapphirina (Osten-Sacken), b.

Aedes aegypti (Linnaeus). m Mentum, ms

Maxillary suture, pip Posterior tentorial pit.

Fig. 62. Larval head and terminal segments.

a. Uvanotaenia sapphirina, b. Aedes Stego-

myia) annandalei(Theobald).

Maxillary suture well developed, extending

from the level of the mentum to the poste-

rior tentorial pit (Fig. 61b); head setae

) and 6 sometimes single and barbed but

never spinelike; comb plate, if present,

Smaller (Fie: G2b) soe eRe ee a, ee

24. Antennal seta and head setae 4-7 small,

delicate, inconspicuous; comb teeth

never in a regular row; siphon at most

about 3.5 times as long as its breadth

at base, usually much shorter; pecten

with at most 7 teeth, usually fewer en

times absent); ventral brush with 4 (rarely 5)

pairs of stout, strongly plumose setae,

some of them usually single (Fig. 63a);

Ethiopian Region only. ........-. Evetmapodites Theobald

Fig. 63. Larval head and terminal segments.

a. Evetmapodites chrysogaster Graham, b.

Aedes aegypti (Linnaeus), c. Segment V of

larval abdomen, E. chrysogaster.

07,99

Mattingly: Illustrated Keys to the Genera of Mosquitoes 27

Without this combination of characters

and Gistrilieior cei nt celia ei eo cage be niece lego peal ah act 25

25. Comb with at most 10 teeth, ina single row;

head seta 5 single, 4 and 6 shorter than

5 with 2 or more delicate branches, 7

single and much longer than any of these

(Fig. 64); Southeast Asia only ......-2.-eee-e Udaya Thurman

Fig. 64. Larval head and terminal segments.

Genus Udaya. a. U. argyrurus (Edwards), b.

U, lucaris Macdonald & Mattingly.

Comb various; head setae otherwise; dis-

tribution various! 40..." as gO Sakae “Baw ie era ee Pasi teetGl Meat 26

26. Comb teeth in a single row; saddle incom-

plete with strongly developed spines

along the distal edge; head seta 7 slender,

delicate, single or bifid, 5 and 6 long,

single, somewhat stouter than 7 but not

conspicuously thickened (Fig. 65a); 61

Southeast Asia Only oF aa eS Zeugnomyia Leicester

Fig. 65. Larval head and terminal segments.

a. Zeugnomyia aguilavi Baisas & Feliciano,

b. Heizmannia scintillans Ludlow.

Without this combination of characters

aid Cisivibution <<. Gs As ee SS ORE 6 ee Ske AAS Sele 27

27. Head seta 4 large and conspicuous, 6

markedly anterior to 5 and 7, 7 with

at least 5 branches, usually more;

stellate setae absent; thoracic integu-

ment devoid of spicules; comb teeth

never fused at base or arising from

a sclerotized plate; pecten teeth with

secondary denticles, if any, confined

to base; saddle incomplete (Fig. 65b);

southern and eastern Asia only. ...... Heizmannia Ludlow®”

Without this combination of characters

and distribution e @ o>. 6.8 ee) &. & . & @ e es e & e ® e e e e e 6 e e e e 28

Fig. 66. a. Larval head. Aedes

(Christophersiomyia) gombakensis Mattingly,

b. Larval head and thorax. Aedes (Finlaya)

poecilus (Theobald).

28. Comb teeth in a single, regular row; anal

segment completely ringed by saddle,

the latter pierced in the mid line by the

proximal setae of the ventral brush which

form a midventral row extending almost

to the base of the anal segment (Fig. 67a,

b): New World only 4 os ess < Psorophora Robineau-Desvoidy®

28 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

Fig. 67. Larval head and terminal segments.

a. Psorophora howardii Coquillett, b. Psoro-

phora signipennis (Coquillett), c. Aedes

diantaeus Howard, Dyar & Knab, d. Aedes

atlanticus Dyar & Knab.

Without this combination of characters

and CIStripution Se we Se ee Re gk eg ee te geNe 29

29. Antenna short, smooth or almost so;

antennal seta very small, single or

bifid, rarely trifid; head setae 5 and

6 slender, single or bifid, 6 markedly

anterior to 7; saddle incomplete; ven-

tral brush arising from a sclerotized

boss (Fig. 68a); New World tropics 64

and SubtPopres Only Woe Gs a ge Haemagogus Williston

Aedes Meigen (part)

Fig. 68. Larval head and terminal segments.

a. Haemagogus capricornii Lutz, b. Aedes

serratus (Theobald).

Without this combination of characters 64

and -distripution sore 4s Ao SPE Ay Aedes Meigen (part)

30. Siphon with a single pair of subventral

setae arising near base (with or with- ,

out a midventral row of setae beyond 55.58

this) €Pies.60bs 69a) s Sie. 'eah ee ee ae ae Culiseta Felt**?

Fig. 69. Terminal segments of larva. a.

Culiseta melanura (Coquillett), b. Culex

pipiens Linnaeus, c. Culex modestus Ficalbi.

Siphon with subventral setae usually well

removed from mid line (Fig. 69b), oc-

casionally forming a midventral row

(Fig. 69c) but never with a separate |

Pair arising ear DASE 24). ee Se a we ee Culex Linnaeus

10.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 29

NOTES ON THE KEYS

Outside the genus Bironella this condition is found only in certain

members of the Anopheles leucosphyrus DOdnitz complex.

Genera Malaya, Hodgesia, Uranotaenia, Zeugnomyia and, in part,

Topomyia are characterized by the fact that vein 1A turns down abruptly

to reach the wing margin before or at most very slightly beyond the base

of the fork of vein Cu (Fig. 6a). This is a very distinctive character

shared only by Aedes subgenus Cancraedes and the males of a few Culex.

In some Limatus spp. there is a tendency for the tip of vein 1A to turn

down abruptly but this seems always to take place well beyond the base of

the fork of vein Cu. In Zeugnomyia the tip of vein 1A turns down abruptly

but sometimes reaches the wing margin slightly beyond the base of the

fork of vein Cu. Doubtful specimens are easily recognized by the vertical

silvery stripe extending from the prealar area down to the mid coxa (Fig.

7).

Topomyia and some species of the closely related genus Malaya (for-

merly Harpagomyia) are very Similarly ornamented, having a conspicuous

median longitudinal white or silvery stripe on the scutum (occasionally

brownish in Topomyia). They can, however, always be distinguished with

ease by the proboscis. This is strongly swollen towards the tip in some

Topomyia but never shows the extensive modifications, associated with

feeding on the regurgitations of ants, which are found in Malaya.

Formerly called Theobaldia by workers in the Old World.

Tripteroides, Malaya and Topomyia, together with the New World gen-

era in couplets 14-17, form the tribe Sabethini of the subfamily Culicinae.

The other subfamilies of Culicidae are the Anophelinae (Anopheles,

Bironella, Chagasia) and the Toxorhynchitinae with the single genus

Toxorhynchites (Megarhinus of earlier authors).

It has been proposed to place the New Zealand species T. argyropus

(Walker) in a monotypic genus Maorigoeldia but this is retained here as a

subgenus of Tripteroides.

One species of Phoniomyia has the tarsi entirely dark in a proportion

of individuals. The very short antenna, of the order of a third of the

length of the proboscis, distinguishes it from other New World sabethines

with comparable proboscis. Some Tvichoprosopon or a few Wyeomyia

Spp. with unusually long proboscis might be confused but these can be re-

cognized by the entirely dark scutellum or tarsi or both. A few Tvicho-

prosopon spp. with white tarsal markings have a peacock blue, rather than

silvery, scale patch on the mid lobe of the scutellum.

A few Trichoprosopon spp. have scutal scales with dull, bluish lustre

but they are never brightly iridescent. A few Wyeomyia spp. with bright

metallic scutal scaling might run down here but these are distinguished by

the presence of prealar bristles.

Two species lacking postnotal bristles have been placed in a separate

genus, Mattinglyia. I prefer to treat this as a subgenus of Heizmannia

and am describing an annectant species elsewhere.

Known only from a single species with undescribed early stages and

uncertain affinities.

it.

12.

13.

14.

15.

16.

Ly.

18.

ro.

20.

21.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

Armigeres differs markedly from Mansonia in the broad scaled vertex

and from Udaya in the heavily scaled pleuron. Aedes is distinguished,

except from Armigeres s. str., by the presence of postspiracular bris-

tles. Aedes subgenus Alanstonea resembles Armigeres s. str. closely in

general facies but differs in having no lower mesepimeral bristle. All

other Aedes lack the curved proboscis. Another character distinguishing

many Aedes is the presence of acrostichal or dorsocentral bristles.

Some authors recognize Mansonia and Coquillettidia as distinct genera,

each with two subgenera. I prefer to include all four subgenera in the

genus Mansonia. Apart from a single species, all the subgenera, except

Coquillettidia, run to the present couplet. Coquillettidia and one species

of subgenus Rhynchotaenia run to couplet 32.

Culex postspiraculosus Lee, from Australia, would also run down here.

It is the only known Culex in which postspiracular bristles occur other-

wise than as an occasional aberration. It is recognizable as a Culex by

the well developed pulvilli. (See Fig. 15a). One species of Aedes, for-

merly placed in a separate genus, Ayuvakitia, lacks postspiracular bris-

tles but runs down correctly, via couplet 25, to the present couplet since

it has toothed fore and mid tarsal claws and palps less than a quarter of

the length of the proboscis. (See also note 27)

The genus Ficalbia is currently held to include four subgenera. In my

view, however, the nominotypical subgenus differs too widely from the

others for this treatment to be acceptable. I prefer, therefore, to treat

Ficalbia s. str. as a separate genus, referring the other three subgenera

to genus Mimomyia. Diagnostic characters are given in the keys.

One species extends beyond the tropics in the extreme eastern part of

its range, as far north as Okinawa. Two others extend some distance

south of the tropics in Queensland.

Subgenus Coquillettidia and one species of subgenus Rhynchotaenia.

lacking postspiracular bristles. (See note 12). One species of Armigeres,

with simple claws, would also run down here but this can be immediately

recognized by the flat scaled vertex, palps more than half as long as pro-

boscis and the curved proboscis (as in Fig. 12).

Based on external characters only.

Males of Aedes subgenus Cancraedes, from southern Asia, would also

run down here. They differ from Uvanotaenia in having postspiracular

bristles and 2 or more lower mesepimerals. (One or none in Uvanotaenia).

A few species of Wyeomyia, with unusually long proboscis, might run

down here but all of these have either the scutellum or the tarsi entirely

dark scaled or both. One species of Phoniomyia has the tarsi entirely

dark in a porportion of individuals but this is recognizable from any

Wyeomyia with a comparable proboscis by the very short antenna (not

more than about one third of the length of the proboscis).

One or two species of Wyeomyia with bright metallic scutal scaling

might run down here but these can be recognized by the presence of pre-

alar bristles.

The very long male palps will separate most Trvichoprosopon spp. from

other New World sabethines. Some of the few Trichoprosopon spp. with

22.

23.

24.

25.

26.

27,

28.

29.

30.

31.

32.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 31

short male palps have narrow scutal scales which are completely diagnos-

tic. The others have the two terminal antennal flagellomeres greatly e-

longated, together about 10 times as long as the antepenultimate flagel-

lomere or more (Fig. 16d). No Trichoprosopon spp. have bright, metal-

lic scutal scaling though a few have scutal scales with dull bluish reflec-

tion. :

See note 9. Males of subgenus Mattinglyia differ from those of the

nominotypical subgenus in having quite strongly plumose antennae and in

lacking postnotal setae. :

Some Haemagogus have short male palps and reduced flagellar setae

but their antenna in no way resembles that of Galindomyia. It is subplu-

mose with all flagellomeres, except the last,very short.

All except subgenus Coquillettidia and one species of subgenus Rhyn-

chotaenia. (See notes 16 and 27).

The Australian Culex postspiraculosus would run down here. (See note

13). It can be recognized by the long, upturned, tapering palps (Fig. 19a).

Subgenus Coquillettidia only. (See note 24).

Subgenus Leicesteria only. Individuals of Mansonia (Rhynchotaenia)

arvribalzagai Theobald lacking postspiracular bristles would also run down

here but this species and subgenus are found only in the New World trop-

ics. Aedes (Kompia) purpureipes Aitken, which is almost unique among

Aedes in lacking postspiracular bristles, wouldalso run down here but this

Species is found only in the United States and Mexico.

Aedes subgenus Ayurakitia would run down here. (See note 13). It in-

cludes only one species and is known only from Thailand. It differs from

Orthopodomyia in many details of ornamentation, among them the restric-

tion of the pleural scaling to four small silvery spots.

In some Bironella spp. the accessory paddle seta is absent or arises

level with the apical paddle seta. These can at once be recognized as

anopheline by the character of seta 9 and the trumpets. Among non-anoph-

eline genera only Aedeomyia (Fig. 27b) has seta 9 approximately as in

Anopheles. One species of Uranotaenia has a trumpet of anopheline type,

but this is recognizable by the other characters given.

Some Mimomyia have a rather similar trumpet but, apart from the

African M. perplexens, these have the float hair well developed. M.

perplexens has the trumpet modified in quite a different way from any

Mansonia sp. (Fig. 28a). Pupae of subgenus Coquillettidia, and apparently

also Rhynchotaenia, have the tips of the trumpets equipped with backwardly

directed barbs which prevent them being withdrawn. The tips break off

short when the pupa rises to the surface at the time of emergence. They

are consequently seldom seen in cast skins.

Absent only in some Mimomyia with a highly characteristic type of

paddle.

Complete separation of Tripteroides pupae from those of other Old

World sabethines is not at present possible but almost all can be recog-

nized by the combination of characters given in the key.

33.

34.

35.

36.

37.

38.

39.

40.

41.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

All but two of the known Topomyia pupae have seta 6-VII as in Fig. 24b.

Both the others have it as in Malaya but differ from that genus in having

all or most of the paddle surface spiculate (Fig. 23a).

Trichoprosopon magnum (Fig. 25e) has longer trumpets than are usual

in the genus but differs from all known Phoniomyia in having them strongly

expanded at the tip and from known Limatus spp. in the large paddles and

relatively shallow excavation of the posterior border of segment VII (Fig.

23f). Wyeomyia circumcincta would key here with Phoniomyia but has the

trumpets much longer than any known in that genus (Fig. 25f).

Pupae of Wyeomyia cannot be entirely separated from those of Sabethes

or T'vichoprosopon on currently available material and descriptions.

Some Sabethes spp. have seta 9-VI more strongly developed, others

less strongly developed, than in the figure but it is always more conspic-

uous than in any known Trvichoprosopon. All those Sabethes spp. with

feebly developed seta 9- VI which are known to me have seta 5-VI more

strongly developed than in any known Trichoprosopon except perhaps T7.

magnum with its distinctive trumpet (Fig. 25e and see note 34).

Subgenera Etorleptiomyia and Ravenalites of genus Mimomyia have

highly distinctive pupae unlikely to be confused with any others (Fig. 28d,

e). The resemblance of some pupae of subgenus Mimomyia to those of

Mansonia has already been mentioned (note 30). A few Uvanotaenia re-

semble Mimomyia superficially in the shape of the paddle and the long

trumpets but can be recognized by the excavation of the basal part of the

inner half of the paddle and the presence of paired setulae on abdominal

segment IX (Fig. 30a). The African M. splendens Theobald is unique in

having a fringe of long, hairlike spicules on both borders of the paddle

but it should run down correctly.

The characteristic paddle shape is usually conspicuous, least so in one

or two aberrant species resembling the Sabethini but differing from these

in the presence of an apical paddle seta (Fig. 30b). A few Aedes (Lor-

vainea) and Armigeres s. str. have a similar type of paddle but with a

delicate fringe on both borders which is very rare in Uvanotaenia (Figs.

32a, b, 35a). The few Uvanotaenia with such a fringe can be recognized

from both the above genera by the reduction of seta 9-VIII and the paddle

seta (Fig. 30c). One or two Haemagogus spp. have a somewhat similar

paddle but are easily recognized by the combination of characters given

in couplet 27. Paired setulae are present on abdominal segment IX in

aa Uranotaenia (Fig. 30a) but absent in a few atypical ones (Fig. 30b,

Cc).

The only species which might be confused are a few members of the

Aedes (Stegomyia) africanus group (Fig. 31c) but these have the paddle

fringe and apical seta less strongly developed and differ in various other

characters among them the cephalothoracic setae as indicated in the key.

Armigeres malayi (Fig. 32b) has seta 6-VI less strongly developed

than in other species but still much better developed than 5-VI. One spe-

cies of Heizmannia (Fig. 32c) would run here but this differs from Arm.

malayi in the much more strongly developed seta 9-VIII and from other

Armigeres in having seta 6-VI much less strongly developed.

Aedes (Lorrainea) spp. have a similar paddle fringe but are recogniz-

able by the characteristic paddle shape (Fig. 35a and see note 38). Ae.

42.

43.

44.

45.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 33

(Diceromyia) spp. differ in the indented tip of the paddle (Fig. 35b) except

for Ae. periskelatus which has the apical paddle seta longer and stouter

than in Heizmannia (Fig. 35c) as do some Stegomyia (Fig. 36a). A few

other Stegomyia present difficulties but can be recognized by having seta

5 on most segments intermediate in type (Fig. 36b), less reduced than in

fi ‘ealaubaaon (Fig. 34b), more so than in other Heizmannia (Figs. 33d,

a).

Some Aedes, particularly Ae. (Finlaya) spp., have a similar paddle

but differ either in the much longer apical paddle seta or in the reduction

of seta 9-VII-VIII or both. The relative position of setae 8-C and 9-C also

furnishes an absolute distinction from Aedes. (See Fig. 40c).

Subgenera Psorophora and Janthinosoma are distinguished from other

genera by the lobed posterior border of segment VIII and/or the toothed

posterolateral corner of segment IV. Most species of subgenus Gvab-

hamia are distinguished, except from Culex and one species of Culiseta

by the presence of an accessory paddle seta. Culex are readily distin-

guishable by the tracheated trumpet or anterior displacement of seta 9-

VU, usually both. The only New World Culiseta with an accessory paddle

seta, C. melanura, is distinguished by having the trumpet cleft nearly to

base, i.e. with very short tubular portion. Aedes atropalpus is said some-

times to have an accessory paddle seta but this species can be recognized

by the short, broad paddle, almost as broad as, or broader than, long

with apex flattened and more or less indented. In Gvabhamia the paddle

is distinctly longer than broad or has a rounded or pointed apex, usually

both. One species of Gvabhamia from the Caribbean area lacks the acces-

sory paddle seta and would run to Aedes in couplet 28. It can be recog-

nized by having seta 2-II well inside 3-II and seta 5-II almost directly ante-

rior to 3-II (Fig. 39c).

The only Haemagogus sp. known to me which might not run down here

is H. chalcospilans. It is recognizable from any known Aedes or Psoro-

phora by the combination of very short setae 5-II and 5-III, setae 5-IV and

o-V only a little longer than the following segments, seta 9-III-VI minute,

seta 9-VII with several branches, 9-VIII about half the length of the paddle,

paddle with apex pointed and short, branched apical paddle seta. A number

of Aedes otherwise resembling Haemagogus have seta 5 on one or more

abdominal segments as long as the two following segments together. In

Haemagogus seta 5 is never more than a little longer than the following

segment. Aedes aegypti is at once distinguished by the well developed

seta 9-III-VI (Fig. 40c).

The relative position of setae 8-C and 9-C appears to be almost com-

pletely diagnostic as between Aedes and Culex. In the former, however,

seta 8-C occasionally arises somewhat posterior to the trumpet (Fig. 40c)

while in the latter it arises further forward in some species than in others.

In comparing pupal skins, therefore, the specimens must be carefully ori-

ented. A few small New World Culex apparently have these setae as in

Aedes but these have trumpets with subbasal tracheation of a kind unknown

in the New World aedine genera (Fig. 41b). In the great majority of Culex

seta 9- VII arises well cephalad of the posterior border of the segment

(Fig. 41a). The few species known to me in which this is not so have

slender trumpets with extensive tracheation of a kind almost unknown in

Aedes (Fig. 42a-c). They are all small or very small species. The few

Aedes with comparable tracheation are all Old World species and are

either large or very large species or have seta 8-C much further forward

than in any Old World Culex (Fig. 42d, e). In contrast to this seta 9-III

46.

47.

48.

49.

00.

52.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

rarely arises cephalad of the posterior border of the segment in Aedes.

All the species in which it is known to do so have trumpets with poorly

developed basal tracheation and seta 8-C arising well forward (Fig. 42f).

(See also note 43).

The relative position of setae 8-C and 9-C appears to be absolutely di-

agnostic as between Aedes and Culiseta (Figs. 40c, 41c). In the latter

seta 8-C always arises far back. Additional partial characters are the ab-

normally long seta 1-IV-VI in most Culiseta (but not the tropical African

species) and the presence of an accessory paddle seta in about half the

known species. These characters should not, however, be needed for di-

agnosis.

Two species of-Sabethes have a pair of small, supernumerary setae in

addition to the single pair of well developed setae by which the ventral

brush is represented in all other known sabethines.

Most Wyeomyia spp. have the setae of the ventral brush short (often

stellate). In some they are longer, though still much shorter than the

lower caudal seta. All but about 3 of the remaining species can be recog-

nized by the arrangement of the comb teeth as indicated in the key. Some

Sabethes spp. have a pair of dorsal, chitinous hooks on abdominal seg-

ment VI. These are unknown in any other New World genus.

A few Trichoprosopon spp. have the siphon moderately long, slender,

tapering with several short subdorsal setae. In these, however, the ven-

tral setae form a long row of branched tufts quite unlike anything found in

Phoniomyia. ‘Two or three Wyeomyia spp. would run down here and are

best separated at the species level.

None of the few species of Wyeomyia with the siphon as in Fig. 48 has

a conspicuous maxillary horn. Two species which might run down to the

first half of the present couplet are W. confusa (Fig. 49b) and W. occulta.

The former, however, has an enormously elongated maxilla, quite unlike

that of any other species in either genus, while the latter differs from any

known Trichoprosopon lacking a mid-ventral row of setae on the siphon in

having a comb of numerous scales in a patch three rows deep. An addi-

tional character serving to distinguish most, though not quite all, species

of Trichoprosopon with the siphon otherwise than in Fig. 48 is the very

small number of comb teeth (5 or less; not less than 10 in Wyeomyia spp.

with maxillary horn). |

Two African Culex spp. lack the ventral brush entirely. In Mimomyia

subgenus Ravenalites it comprises two pairs of setae only. Some mem-

bers of this subgenus have a superficial resemblance to Tripteroides but

they can be recognized at once by the articulated terminal joint of the an-

tenna and the presence of only one pair of subventral setae on the siphon

(Fig. 51a). In Culex (Acallyntrum) spp. from Melanesia the ventral brush

is represented by a single pair of setae. Larvae of these species differ

from those of the sabethine genera in lacking dorsal or subdorsal setae on

the siphon (Fig. 51b). In addition the larvae of the sabethine genera al-

most always have the antenna and antennal seta much reduced while in

many the thorax and abdomen are covered with large stellate setae (Fig.

52a), a condition not met with in Acallyntrum .

Some Topomyia have the comb reduced to a single row or almost so.

These, however, can at once be separated from Tripteroides, as can all

other Topomyia and Malaya, by the large fan-shaped setae 5 and 6 on the

O3.

o4.

OD.

06.

Ot.

08.

o9.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 35

prothorax wae 50). A few Topomyia Shp: have well developed stellate

setae on the thorax and/or abdomen but these are never so strongly devel-

oped as in many Tvipteroides (Figs. 50b, 52a). The modified mesothco-

racic seta 6 and/or metathoracic seta 7 (Fig. 52a), when present as they

are in the majority of species, are completely diagnostic of 'Tripteroides.

T. (Maorigoeldia) argyropus is unique in the genus in having a comb with

numerous scales in a patch but this species is confined to New Zealand.

Two of the three known species of Topomyia with strongly developed

maxillary horns lack stellate setae on the abdomen. They are, however,

readily distinguished from Malaya in which, so far as is known, maxillary

horns never occur. Horns of this kind are found in various Australasian

Tripteroides but not in either of the known Australasian Topomyia (one un-

described). One species of Topomyia (T. spathulirostris) lacks maxillary

horns and has the stellate setae poorly developed. It can, however, be

on PA gaeera by the siphon which is much longer than in any Malaya

Fig. 53c).

A few African, Southeast Asian and Melanesian Mimomyia have a

piercing siphon (Fig. 54b) but this is much less elaborately modified than

in Mansonia and there should be no danger of confusion.

Most Culiseta have a pair of subventral setae arising at or near the

base of the siphon and are without other siphonal setae, but only three of

them are found in the tropics. Two of these are recognizable by the fact

that the comb teeth are arranged in a patch (Fig. 60b) and the third by the

presence of a midventral row of setae on the siphon.

An additional character is the presence in most. Hodgesia larvae of

several long, very delicate setae on the anal segment anterior to the bar-

red area of the ventral brush (Fig. 57a). These are easily broken off,

however, and when this happens their bases are virtually impossible to

detect. One or two short, delicate setae anterior to the ventral brush are

also sometimes seen in Ficalbia.

The only culicine genera consistently lacking a pecten are Armigeres

and Orthopodomyia, Among the other genera remaining to be keyed this

condition is found in whole or in part in two species of Culex, two of

Uvanotaenia and half dozen Evetmapodites. The latter are confined to the

Ethiopian Region where Armigeres is absent and Orthopodomyia very rare.

They differ from Orthopodomyia in many characters including the reduced

antennal seta, head setae and ventral brush (Fig. 63a). Of the two Culex

spp. C. moucheti lacks the ventral brush entirely while C. dispectus has

4 pairs of subventral tufts on the siphon. The two Uranotaenia spp. both

occur outside the known geographical range of Armigeres and Orthopodo-

myia, One (U. browni from the Seychelles) has the siphon densely covered

with long spicules. The other (U. colocasiae from Fiji) has the ventral

brush much as in Eretmapodites.

One species of Culiseta, known only from southern Australia, would

run down here instead of to the second half of the couplet. It is easily re-

cognized from any other species with which it might be confused by the

oe aa complete saddle and long head setae 5 and 6 and antennae

Fig. 60c).

New World Uvranotaenia larvae are recognizable also by the more or

less thickened, barbed, spinelike head setae 5 and 6, very large comb

plate and apically fringed pecten scales. Many of the Old World species

60.

61.

62.

63.

64.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

also exhibit some or all of these characters but some do not and for these

the only absolutely diagnostic character is the absence or extreme reduc-

tion of the maxillary suture (Fig. 6la).

The only species likely to be confused are some Aedes subgenus Stego-

mytia. These are easily recognized by the long, slender, non-plumose se-

tae of the ventral brush (Fig. 63b). Most of them also differ in having the

comb teeth in a single row and in the more numerous pecten teeth.

character which is completely diagnostic for Eretmapodites, when present,

is the occurrence of rugose, sclerotized bosses at the bases of the princi-

pal lateral abdominal setae (Fig. 63c).

Among the remaining Southeast Asian genera with a single pair of sub-

ventral setae on the siphon Heizmannia is distinguished by the much more

strongly developed head seta 7 (and usually also 5) and less strongly spic-

ulate saddle edge (Fig. 65b). Many Aedes differ in having head seta 7 (at

least) more strongly developed or the comb teeth in more than one row.

Those which do not differ in either of these characters, e.g. most Stego-

myia are distinguished by having the saddle edge finely spiculate at most

while some also differ in having the comb teeth arising from a sclerotized

plate. One Southeast Asian Finlaya (Aedes harperi from the Philippines)

seems from the published description to be distinguishable only at the spe-

cies level.

Larvae of this genus closely resemble those of some Aedes. They often

have a characteristic head seta 6 with two unequal branches (Fig. 65b).

This, when present, is absolutely diagnostic except from Aedes subgenus

Christophersiomyia, easily recognized by the much smaller head seta 4

(Fig. 66a) and one or two Aedes (Finlaya) spp. differing in other charac-

ters given in the key. Ae. (Christophersiomyia) spp., like many other

Aedes, are also distinguished by the fact that head seta 6 arises only very

slightly, if at all, anterior to 7. Aedes spp. resembling Heizmannia in

the well developed head seta 4 and the position of head seta 6, e.g. many

Ae. Stegomyia) spp., can be recognized by the feebly developed head seta

7 (Fig. 62b). Other characters serving to separate a few doubtful species

are the presence of stellate setae on thorax and abdomen (Fig. 66b) or the

possession of a spiculated thoracic and abdominal integument.

Larvae of subgenus Psorophora s. str. have the mouthbrushes modified

for predation (Fig. 67a) and are unlikely to be mistaken for anything else.

Those of the other subgenera show some resemblances to Aedes. How-

ever, most New World Aedes have the comb teeth in two or more rows

while those which do not either have the saddle incomplete (Fig. 67c) or

the ventral brush confined to that part of the anal Segment which is not

ringed by the saddle (Fig. 67d). None of them have the comb arising from

a sclerotized plate as in many Psorophora (Fig. 67b). Some Haemagogus

have the comb arising from a small sclerotized plate but these all have the

saddle incomplete and lack the anterior extension of the ventral brush.

They are also distinguished by the much reduced antenna (Fig. 68a).

Most New World Aedes are distinguishable from Haemagogus either by

the longer or more strongly spiculate antenna, position and branching of

the head setae, complete saddle or absence of lateral sclerotizations from

the barred area at the base of the ventral brush (Fig. 68b) or by a combi-

nation of these. There remain a number of Aedes, particularly in the sub-

genera Finlaya and Howardina, which are best separated at the species

level.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 37

BIBLIOGRAPHY

As a source of references to the taxonomic literature the Synoptic

Catalog (Stone et al., 1959) and supplements (Stone, 1961-70) are indispens-

able. They can be kept up to date with the aid of one of the monthly bibliogra-

phies. The Bulletin Analytique d'Entomologie Medicale et Veterinaire, pub-

lished by the Office de la Recherche Scientifique et Technique Outre-mer at

Bondy, Seine-et-Oise, is the best. As a general introduction to the fauna of

the various regions, and a source of further references, the following are re-

commended.

Palaearctic Region 3

Gutsevitch et al. (1970), Guy (1959), Kramar (1958), La Casse & Yamaguti

(1950), Mattingly & Knight (1956), Mihalyi (1963), Mohrig (1969), Natvig (1948),

Rioux (1958), Senevet & Andarelli (1959), University of Maryland (1963).

Ethiopian Region

Edwards (1941), Gillies & de Meillon (1968), Grjebine (1966), Haddow et al.

(1951), Hamon (1963), Hopkins (1952), Leeson (1958), Mattingly (1952, 1953),

Mattingly & Brown (1955), Mattingly & Lips (1953), Muspratt (1955, 1956),

Ovazza et al. (1956), Van Someren et al. (1955).

Oriental Region

Barraud mee | Bonne-Wepster (1954), Borel (1930), Bram (1967a),

Christophers (1933), Delfinado (1966), Lien (1968), Macdonald (1957, 1960),

Mattingly (1965), Reid (1968), Reuben (1969), Thurman (1959), Tyson (1970),

Wharton (1962).

Australian Region (including Micronesia)

Belkin Heed 1968), Bohart (1956), Dobrotworsky (1965), Huang (1968a, b),

Marks (1954), O'Gower (1958), Ramalingam & Belkin (1965), Sirivanakarn

(1968), Steffan (1966, 1968), Van den Assem & Bonne-Wepster (1964).

Nearctic Region

Barr (1958), Beckel (1954), Carpenter (1968), Carpenter & La Casse (1955),

Darsie (1951), Gjullin et al. Beye King et al. (1960), Ross & Horsfall (1965),

Smith (1969), Steward & McWade (1961), Vockeroth (1954), Zavortink (1970).

Neotropical Region

Belkin et al. (1970), Belkin & Hogue (1959), Berlin (1969a, b), Bram (1967b),

Correa & Ramalho (1956), Cova-Garcia (1961), Cova-Garcia et al. (1966),

Foote (1954), Forrattini (1962, 1965a, b), Galindo et al. (1954), Garcia &

Ronderos (1962), Guedes et al. (1965), Lane (1953), Ronderos & Bachmann

(1963b), Schick (1970).

General

Certain of the above overlap two or more adjacent regions, notably Mattingly

& Knight, 1956 (Palaearctic, Oriental and Ethiopian), University of Maryland,

1963 (Palaearctic and Oriental), Bonne-Wepster, 1954 (Oriental and Austral-

ian), Belkin & Hogue, 1959; Bram, 1967b; Foote, 1954; Galindo et al., 1954;

Zavortink, 1970 (Neotropical and Nearctic). Recent taxonomic works with

World coverage are Maslov (1967) and Zavortink (1968). Foote & Cook (1959)

is useful as a general survey of vector species.

38 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

REFERENCES

BARR, A.R.

1958. The mosquitoes of Minnesota (Diptera: Culicidae: Culicinae).

Tech. Bull. Univ. Minn. agric. Exp. Stn, 228. 154 pp.

BARRAUD, P. J.

1934. The Fauna of British India. Diptera, 5 (Culicidae: Megarhinini

and Culicini). London: Taylor & Francis. 463 pp.

BATES, M.

1949. The Natural History of Mosquitoes. New York: Macmillan.

(Republished by:-Harper Torchbooks, 1965). 378 pp.

BECKEL, W. E.

1954. The identification of adult female Aedes mosquitoes (Diptera,

Culicidae) of the black-legged group taken in the field at

Churchill, Manitoba. Can. J. Zool. 32:324-330.

BELKIN, J.N.

1962. The Mosquitoes of the South Pacific (Diptera, Culicidae).

Berkeley & Los Angeles: Univ. Calif. Press. 2 vols. 1020 pp.

1968. Mosquito studies (Diptera, Culicidae) VI. The Culicidae of

New Zealand. Contr. Am. ent. Inst. 3(1):1-182.

BELKIN, J.N., HEINEMANN, S.J. & W.A. PAGE

1970. Mosquito studies (Diptera, Culicidae) XXI. The Culicidae of

Jamaica. Contr. Am. ent. Inst. 6(1):1-458.

BELKIN, J.N. & C.L. HOGUE

1959. A review of the crabhole mosquitoes of the genus Deinocerites

(Diptera, Culicidae). Univ. Calif. Publ. Ent. 14:411-458.

BERGE, T.O., SHOPE, R.E. & T.H. WORK

1970. Catalogue of arthropod-borne viruses of the World. Am. J.

trop, Med. Hyg. 19:1082-1160.

BERLIN, O.G.W. |

1969a. Mosquito studies (Diptera, Culicidae) XM. A revision of the

Neotropical subgenus Howardina of Aedes. Contr. Am. ent.

Inst. 4(2):1-190. 7

1969b. Mosquito studies (Diptera, Culicidae) XVII. The subgenus

Micraedes of Culex. Contr. Am. ent. Inst. 5(1):21-63.

BOHART, R.M.

1956. Insects of Micronesia. 12(1). Diptera: Culicidae. Honolulu:

Bishop Museum. 85 pp.

BONNE-WEPSTER, J.

1954. Synopsis of a hundred common non-anopheline mosquitoes of

the Greater and Lesser Sundas, the Moluccas and New Guinea.

Publs R. trop. Inst. Amsterdam, 111. 147 pp.

BOREL, E.

1930. Les moustiques de la Cochinchine et duSud-Annam. Monogr.

Colln Soc. Path. exot. 3. 423 pp.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 39

BRAM, R.A.

1967a. Contributions to the mosquito fauna of Southeast Asia II. The

genus Culex in Thailand (Diptera: Culicidae). Contr. Am. ent.

Inst. 2(1):1-296.

1967b. Classification of Culex subgenus Culex in the New World

(Diptera: Culicidae). Proc. U.S. natn. Mus. 120:1-120.

BUSVINE, J.R.

1966. Insects and Hygiene. 2nd Edn. London: Methuen. 467 pp.

CARPENTER, S.J.

1968. Review of recent literature on mosquitoes of North America.

Calif. Vector Views 15:71-98.

CARPENTER, S.J. & W.J. LA CASSE

1955. Mosquitoes of North America. Berkeley & Los Angeles: Univ.

Calif. Press. 360 pp.

CHRISTOPHERS, S.R.

1933. The Fauna of British India. Diptera, 4 (Culicidae: Anophelini).

London: Taylor & Francis. 371 pp.

1960a. Aedes aegypti (L.), the Yellow Fever Mosquito. Cambridge

Univ. Press. 739 pp.

1960b. The generic name Aedes. Trans. R. Soc. trop. Med. Hyg.

54:407-408.

CLEMENTS, A.N.

1963. The Physiology of Mosquitoes. London: Pergamon Press. 393 pp.

CORREA, R.R. & G.R. RAMALHO

1956. Revisao de Phoniomyia Theobald, 1903 (Diptera, Culicidae,

Sabethini). Folia clin. biol. 25:1-176.

COVA-GARCIA, P.

1961. Notas sobre los Anofelinos de Venezuela y su Identificacion.

2nd Edn. Caracas: Editora Grafas. 213 pp.

COVA-GARCIA, P., SUTIL, E. & J.A. RAUSSEO

1966. Mosquitos (Culicinos) de Venezuela. 2vols. Caracas: Mini-

sterio de Sanidad. 823 pp.

DARSIE, R. F.

1951. Pupae of the culicine mosquitoes of the Northeastern United

States (Diptera, Culicidae, Culicini), Mem. Univ. Cornell

agric. Exp. Stn 304. 67 pp.

DELFINADO, M.

1966. The culicine mosquitoes of the Philippines, Tribe Culicini

(Diptera, Culicidae), Mem. Am. ent. Inst. 7. 252 pp.

de MEILLON, B. et al.

1967. Papers on Culex pipiens fatigans. Bull. Wid Hlth Org. 36(1):

1-100 & 163-180.

40 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

DETINOVA, T.S.

1968. Age structure of insect populations of medical importance.

A. Rev. Ent. 13:427-450.

DOBROTWORSKY, N.V.

1965. The Mosquitoes of Victoria. Melbourne Univ. Press. 237 pp.

EDESON, J. F.B. & T. WILSON

1964. The epidemiology of filariasis due to Wuchereria bancrofti and

Brugia malayi. <A. Rev. Ent. 9:245-268.

EDWARDS, F.W.

1941, Mosquitoes of the Ethiopian Region III. Culicine adults and

pupae. London: Br. Mus. (Nat. Hist.). 499 pp.

FOOTE, R.H.

1954. The larvae and pupae of the mosquitoes belonging to the Culex

subgenera Melanoconion and Mochlostyrvax. Tech. Bull. U.S.

Dep. Agric. 1091. 126 pp.

FOOTE, R.H. & D.R. COOK

1959. Mosquitoes of Medical Importance. Agric. Handb. Forest

Serv. U.S. 152. 158 pp.

FORATTINI, O. P.

1962. Entomologia Medica. I. Parte Geral. Diptera, Anophelini.

Univ. de Sao Paulo. 662 pp.

1965a. Entomologia Medica. II. Culicini: Culex, Aedes e Psorophora.

Univ. de Sao Paulo. 506 pp.

1965b. Entomologia Medica. HI. Culicini: Haemagogus, Mansonia,

Culiseta, Sabethini. Toxorhynchitini. Arboviroses. Filariose

bancroftiana. Genetica. Univ. de Sao Paulo. 416 pp.

GALINDO, P., TRAPIDO, H. & S.J. CARPENTER

1950. Observations on diurnal forest mosquitoes in relation to sylvan

yellow fever in Panama. Am. J. trop. Med. 30:533-574.

GALINDO, P., BLANTON, F.S. & E.L. PEYTON

1954. A revision of the Uvanotaenia of Panama with’notes on other

American species of the genus. Ann. ent. Soc. Am. 47:107-

wee

GARCIA, M. & R.A. RONDEROS :

962. Mosquitos de la Rep ublica Argentina I. Tribu Anophelini

(Diptera-Culicidae-Culicinae), Anu. Comis. Invest. Cient.

Prov. B. Aires 3:105-164.

GARNHAM, P.C.C.

1966. Malaria Parasites and other Haemosporidia. Oxford: Black-

well. 1114 pp.

GILLETT, J.D. |

1962. Contributions to the oviposition-cycle by the individual mosqui-

toes in a population. J. Insect Physiol. 8:665-681.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 41

GILLIES, M.T. & B. de MEILLON

1968. The Anophelinae of Africa South of the Sahara. PublsS. Afr.

Inst. med. Res. 54. 343 pp.

GJULLIN, C.M., SAILER, R.I., STONE, A. & B.V. TRAVIS

1961. The mosquitoes of Alaska. U.S. Dep. Agric. Hdbk 182. 98 pp.

GRJEBINE, A.

1966. Biologie et Taxonomie des Anophelinae de Madagascar et des

Isles Voisines. Doctoral thesis. Universite de Dijon. 487 pp.

GUEDES, A.S., SOUZA, M.A., MACIAL, C.S. &8.H. XAVIER

1965. Catalogo illustrado dos mosquitos da colecao do Instituto

Nacional de Endemias Rurais I. Genero Psorophora Robineau-

Desvoidy, 1827. Revta bras. Malar. Doenc. trop. 17:3-24.

GUTSEVICH, A.V., MONCHADSKI, A.S. & A.A. SHTAKEL'BERG

1970. Fauna of USSR. Vol. III. No. 4. Mosquitos. Family Culicidae.

Leningrad: Zoological Institute. 384 pp. In Russian.

GUY, X.

1959. Les Anopheles du Maroc. Mem. Soc. Sci. nat. phys. Maroc,

Zool, n.s.. ‘21-236.

HADDOW, A.J.

1955. Studies of the biting habits of African mosquitoes. An appraisal

of methods employed, with special reference to the twenty- four-

hour catch. Bull. ent. Res. 45:199- 242.

HADDOW, A.J., VAN SOMEREN, E.C.C., LUMSDEN, W.H.R., HARPER,

J.O. & J.D. GILLETT

1951. The mosquitoes of Bwamba County, Uganda, VIII. Records of

“inact behaviour and habitat. Bull. ent. Res. 42:207-

HAMON, J.

; 1963. Les moustiques anthrophiles de la region de Bobo- Dioulasso

(Republique de Haute-Volta). Annis Soc. ent. Fr. 132:84-144.

HOPKINS, G.H.E.

1952. Mosquitoes of the Ethiopian RegionI. Larval Bionomics of

Mosquitoes and Taxonomy of Culicine Larvae. 2nd Edn.

London: Br. Mus. (Nat. Hist.). 355 pp.

HORSFALL, W.R.

1955. Mosquitoes: their Bionomics and Relation to Disease. London:

Constable. 723 pp.

HUANG, Y.-M.

1968a. A new subgenus of Aedes (Diptera, Culicidae) and illustrated

key to the subgenera of Aedes of the Papuan Subregion. J.

med. Ent. 5:169-188.

1968b. Aedes (Verrallina) of the Papuan Subregion (Diptera: Culicidae).

Pacif. Insects Monogr. 17:1-73.

JACHOWSKI,

1954.

KING, W.V.;

1960.

KRAMAR, J.

1958.

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

L. A.

Filariasis in American Samoa V. Bionomics of the principal

vector, Aedes polynesiensis Marks. Am. J. Hyg. 60:186- 203.

BRADLEY, G.H., SMITH, C.N. & W.C. McDUFFIE

A handbook of the mosquitoes of the southeastern United States.

U.S. Dep. Agric. Handbk 173. 188 pp.

Fauna of Czechoslovakia. 13. Biting Mosquitoes - Culicinae.

Csechoslovenske Akad. Ved. 286 pp. In Czech.

LA CASSE, W.J. & S. YAMAGUTI

1950.

LAIRD, M.

1956.

LANE, J.

1953.

LEESON, H.S

1958.

LIEN, J.C.

1968.

Mosquito Fauna of Japan and Korea. 3rd Edn. Kyoto: Office

of Surgeon, 8th Army H.Q. 487 pp.

Studies of mosquitoes and fresh water ecology in the South

Pacilic. . Bull. R. Soc. N.Z. 6. 213 pp.

Neotropical Culicidae. Univ. de Sao Paulo. 2vols. 1112 pp.

An annotated catalogue of the culicine mosquitoes of the Federa-

tion of Rhodesia and Nyasaland and neighbouring countries, to-

gether with locality records from Southern Rhodesia. Trans.

R. ent. Soc. Lond. 110:21-51.

New species of mosquitoes from Taiwan (Diptera: Culicidae)

Part V. Three new subspecies of Aedes and seven new species

of Culex. Trop. Med., Nagasaki 10:217- 262.

MACDONALD, W. W.

1957.

1960.

Malaysian Parasites XVI. An interim review of the non-anoph-

eline mosquitoes of Malaya. Stud. Inst. med. Res. Malaya

28:1-34.

Malaysian Parasites XXXVIII. On the systematics and ecology

of Armigeres subgenus Leicesteria (Diptera, Culicidae). Stud.

Inst. med. Res. Malaya 29:110-153.

MACDONALD, W.W. & R. TRAUB

1960.

MARKS, E.N.

1954.

Malaysian Parasites XXXVII. An introduction to the ecology of

the mosquitoes of the lowland dipterocarp forest of Selangor,

Malaya. Stud. Inst. med. Res. Malaya 29:79-109.

A review of the Aedes scutellaris subgroup with a study of vari-

ation in Aedes pseudoscutellaris (Theobald) (Diptera: Culicidae).

Bull. Br. Mus. nat. Hist. Ent. 3:349-414.

MARSHALL, J. F.

1938.

The British Mosquitoes. London: Br. Mus. (Nat. Hist.). 341

pp. oe by Johnson Reprint Company, New York in

1966.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 43

MASLOV, A.V. |

1967. Bloodsucking mosquitoes of the sub-tribe Culisetina (Diptera,

Culicidae) of the world fauna. Opredeli Faune SSSR. 93. 182

pp. In Russian.

MATTINGLY, P. F.

1952. The subgenus Stegomyia (Diptera: Culicidae) in the Ethiopian

Region. I. A preliminary study of the distribution of species

occurring in the West African Sub-region with notes on taxonomy

and bionomics. Bull. Br. Mus. nat. Hist. Ent. 2:235-304.

1953. The subgenus Stegomyia (Diptera: Culicidae) in the Ethiopian

Region. II. Distribution of species confined to the East and

South African Sub-region. Bull. Br. Mus. nat. Hist. Ent. 3:

1-65,

1958. A revision of Pavaédes Edwards and Cancraédes Edwards

(Diptera: Culicidae). Proc. R. ent. Soc. Lond., B, 27:76-83.

1962a. Towards a zoogeography of the mosquitoes. Publs Syst. Ass.

4:17-36.

1962b. Mosquito behaviour in relation to disease eradication program-

mes. A. Rev. Ent. 7:419-436.

1965. The Culicine Mosquitoes of the Indomalayan Area VI. Genus

Aedes Meigen subgenus Stegomyia Theobald (Groups A, B and

D). London: Br. Mus. (Nat. Hist.). 67 pp.

1969. The Biology of Mosquito-borne Disease. London: Allen &

Unwin. 184 pp.

AGT “ie a aspects of mosquito evolution. Riv. Parassit. (in

press).

197 . Cyclical behaviour in mosquitoes. Trans. 13 internat. Congr.

Ent., Moscow (in press).

MATTINGLY, P.F. & E.S. BROWN

1955. The mosquitos (Diptera: Culicidae) of the Seychelles. Bull.

ent. Res. 46:69-110.

MATTINGLY, P. F. & K.L. KNIGHT

1956. The 2 lama of Arabia. I. Bull. Br. Mus. nat. Hist. Ent.

4:91-141.

MATTINGLY, P. F. & M. LIPS

1953. Notes on the Culicini of the Katanga (Diptera, Culicidae). Revue

Zool. Bot. afr. 47:311-343 and 48:49-72.

MIHALYI, F.

1963. Biting Mosquitos of Hungary. Budapest: Akademiai Kiado.

229 pp. In Hungarian.

MOHRIG, W.

1969. Die Culiciden Deutschlands. Parasit. SchrReihe 18:1- 260.

44 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

MUIRHEAD- THOMSON, R.C.

1951. Mosquito Behaviour in Relation to Malaria Transmission and

Control in the Tropics. London: Arnold. 219 pp.

1968. Ecology of Insect Vector Populations. London & New York:

Academic Press. 174 pp.

MUSPRATT, J.

1955. Research on South African Culicini (Diptera, Culicidae) II. A

check list of the species with notes on taxonomy, bionomics and

identification. J. ent. Soc. sth. Afr. 18:149- 207.

1956. The Stegomyia mosquitoes of South Africa and some neighbour-

ing territories. Mem. ent. Soc. sth. Afr. No. 4. 138 pp.

NATVIG, L.R.

1948. Contributions to the knowledge of the Danish and Fennoscandian

mosquitoes. Norsk. Ent. Tidsskr. Suppl. I. 567 pp.

O'GOWER, A.K.

1958. The mosquitoes of North Western Australia. Serv. Publs Dep.

Hlth Aust. Sch. publ. Hlth trop. Med. No. 7. 46 pp.

OVAZZA, M., HAMON, J. & P. NERI

1956. Contribution a l'etude des dipteres vulnerants de l'Empire

d'Ethiopie. I. Culicidae. Bull. Soc. Path. exot. 49:151-182.

PAMPANA, E.J.

1969. A Textbook of Malaria Eradication. 2nd Edn. London: Oxford

Univ. Press. 593 pp.

RAMALINGAM, S. & J.N. BELKIN

1965. Mosquito studies (Diptera, Culicidae) Il. Two new Aedes from

Tonga and Samoa. IV. The mosquitoes of the Robinson- Peabody

Museum of Salem expedition to the Southwest Pacific, 1956.

Contr. Am. ent. Inst. 1(4):1-34.

REEVES, W.C.

1965. Ecology of mosquitoes in relation to arboviruses. A. Rev.

Ent. 10:25-46. :

REID, J.A. :

1968. Anopheline mosquitoes of Malaya and Borneo. Stud. Inst. med.

Res. Malaya 31. 520 pp.

REUBEN, R.

1969. A redescription of Culex vishnui Theo., with notes on C.

pseudovishnui Colless and C, tritaeniorhynchus Giles from

southern India. Bull. ent. Res. 58:643-652.

RIOUX, J.A. :

1958. Les Culicides du "Midi" Mediterraneen. Encyclopedie Entomo-

logique. 35. Paris: Lechevalier. 303 pp.

RONDEROS, R.A. & A.O. BACHMANN

1963a. A proposito del complejo Mansonia (Diptera, Culicidae). Revta

Soc. ent. argent. 25:43-51.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 45

RONDEROS, R.A. & A.O. BACHMANN

1963b. Mansoniini neotropicales I (Diptera- Culicidae). Revta Soc.

ent. argent. 26:57-65.

ROSS, H.H. & W.R. HORS FALL

1965. A synopsis of the mosquitoes of Illinois. Illinois nat. Hist.

Surv. biol. Notes 52. 50 pp.

RUSSELL, P. F., WEST, L.S., MANWELL, R.D. & G. MACDONALD

1963. Practical Malariology. 2nd Edn. London: Oxford Univ. Press.

750 pp. |

SCHICK, R.X.

1970. Mosquito studies XX. The terrens group of Aedes (Finlaya),

Contr. Am. ent. Inst. 5(3):1-158.

SENEVET, G. & L. ANDARELLI

1959. Les moustiques de l'Afrique du Nord et du bassin mediterra-

neen. Encyclopodie Entomologique. 37. Paris: Lechevalier.

383 pp. |

SIRIVANAKARN, S.

1968. The Culex subgenus Lophoceraomyia in New Guinea and Bis-

marck Archipelago (Diptera: Culicidae). Pacif. Insects

Monogr. 17:75-186.

SMART, J.

1965. A Handbook for the Identification of Insects of Medical Impor-

tance. 4th Edn. London: Br. Mus. (Nat. Hist.). 303 pp.

SMITH, M.E.

1969. The Aedes mosquitoes of New England (Diptera, Culicidae) II.

Larvae. Can. Ent. 101:41-51.

STEFFAN, W.A.

1966. A checklist and review of the mosquitoes of the Papuan Subre-

gion. J. med. Ent. 3:179-237. —

1968. Avmigeres of the Papuan Subregion (Diptera: Culicidae). J.

med. Ent. 5:135-159.

STEWARD, C.C. & JI.W. McWADE

1961. The mosquitoes of Ontario (Diptera: Culicidae) with keys to the

tari and notes on distribution. Proc. ent. Soc. Ont. 91:

-188.

STONE, A.

1961. A synoptic catalog of the mosquitoes of the world, Supplement

I (Diptera: Culicidae). Proc. ent. Soc. Wash. 63:29-52.

1963. A synoptic catalog of the mosquitoes of the world, Supplement

II tiptera, Culicidae). Proc. ent. Soc. Wash. 65:117-140. |

1967. A synoptic catalog of the mosquitoes of the world, Supplement

Ill (Diptera, Culicidae). Proc. ent. Soc. Wash. 69:197- 224.

1970. A synoptic catalog of the mosquitoes of the world, Supplement

IV (Diptera, Culicidae). Proc. ent. Soc. Wash. 72:137-171.

46 Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

STONE, A. & P. BARRETO

1969. A new genus and species of mosquito from Colombia, Galindo-

myia leei (Diptera, Culicidae, Culicini). J. med. Ent. 6:143-

146.

STONE, A., KNIGHT, K.L. & H. STARCKE

1959. A Synoptic Catalog of the Mosquitoes of the World. Thomas

Say Foundation. Vol. 6. Washington, D.C.: Entomological

Society of America. 358 pp.

TAYLOR, B. & M.D.R. JONES

1969. The circadian rhythm of flight activity in the mosquito Aedes

aegypti (L. ); the phase-~ setting effect of light-on and light- off.

J. exp. Biol. 51:59-70.

TAYLOR, R.M.

1967. Catalogue of Arthropod-borne Viruses of the World. Public

Health Service Publication No. 1760. Washington, D.C.: U.S.

Govt Printing Office. 898 pp.

THURMAN, E.B.

1959. A contribution to a revision of the Culicidae of northern Thailand.

Bull. Md agric. Exp Stn. A-100. 180 pp.

TRAPIDO, H. & P. GALINDO

1957. Mosquitoes associated with sylvan yellow fever near Almirante,

Panama. Am. J. trop. Med. Hyg. 6:114-144.

TYSON, W.H.

1970. Contributions to the mosquito fauna of Southeast Asia VII. Ge-

nus Aedeomyia Theobald in Southeast Asia. VIII. Genus Aedes,

subgenus Mucidus Theobald in Southeast Asia. Contr. Am. ent.

Inst. 6(2):1-80.

UNIVERSITY OF MARYLAND

1963. Index Catalogue to Russian, Central and Eastern European, and

Chinese Literature in Medical Entomology. I. Diptera.

College Park, Md: Dept. of Zoology. 243 pp.

VAN DEN ASSEM, J. & J. BONNE-WEPSTER

1964. New Guinea Culicidae, a synopsis-of vectors, pests and com-

mon species. Zool. Bijdr. 6. 136 pp.

VAN SOMEREN, E.C.C., TEESDALE, C. & M. FURLONG

1955. The mosquitos of the Kenya coast; records of occurrence, be-

haviour and habitat. Bull. ent. Res. 46:463-493.

VOCKEROTH, J.R.

1954. Notes on the identities and distributions of Aedes species of

northern Canada with a key to the females (Diptera: Culicidae).

Can. Ent. 86:241- 255.

WARD, R.A. & J.E. SCANLON (Eds)

1970. Conference on anopheline biology and malaria eradication.

Misc. Publs ent. Soc. Am. 7:1-196.

Mattingly: Illustrated Keys to the Genera of Mosquitoes 47

WHARTON, R.H.

1962. The biology of Mansonia. mosquitoes in relation to the trans-

mission of filariasis in Malaya. Bull. Inst. med. Res. Malaya

11. 114 pp.

WORLD HEALTH ORGANIZATION

1970. Insecticide resistance and vector control. Tech. Rep. Ser.

Wld Hlth Org. 443. 279 pp.

WRIGHT, J.W. & R. PAL (Eds)

1967. Genetics of Insect Vectors of Disease. Amsterdam, London &

New York: Elsevier. 794 pp.

ZAVORTINK, T.J.

1968. Mosquito studies (Diptera, Culicidae) VIII. A prodrome of the

genus (Orthopodomyia. Contr. Am. ent. Inst. 3(2):1-221.

1970. The treehole Anopheles of the New World. Contr. Am. ent.

Inst. 5(2):1-35.

ACKNOWLEDGEMENTS

I am indebted to Dr. Botha de Meillon and the staff of the Southeast

Asia Mosquito Project, and to Dr. Alan Stone, for much kindness and hospi-

tality in Washington and for checking the keys and making many souc criti-

cisms and suggestions. All mistakes are my own.

SS Toxorhynchites splendens

o b i ~postnotal

setulae

ape re

Chagasia Anopheles Wyeomyila Trichoprosopon

fajardo/ balabacensis aporonoma digitatum

Hodgesia malay! 6b

Trichoprosopon way

allidiventer -

e Sc

Zeugnomyia

be M. Druckenbrod gracilis

ff Sabethes belisario/

i Q mid leg

hind femur

SS : ies

Nee —— 2

iii

Ni Ni i= UN

si ms

ric

Aedeomyia catasticta

Opifex fuscus

Deinocerites cancer

<=>

Ss" t.

= oN IZ PLA)

ee, Y A

Mansonia (Mansonioides) uniformis

14a

Mimomyia (Etorleptiomyia)

Udaya Jucaris /uzonensis

Mansonia (Coquillettidia)

crassipes

Mimomyia (Ravenalites)

deguzmanae

mid leg

fore tarsal claw ventral

/ 3

M. Druckenbroo ~

hind tarsal claw ventral 1ba

Culex sp. Orthopodomyia anopheloides

16d

— 5 = “ae

Ne By A AY,

; Uy

WW PAN

— we

Limatus durhamii

Trichoprosopon perturbans

Limatus

asulleptus

aE HI see

> en ie eo Hl SK

Gira ae 4

SN EE ER

CARY

PI. 6

17b

M. Druckenbrod

Galindomyia /eei

fore tarsal claw

Heizmannia scintillans

19c

EAS

| RSS

1 / Wi =

| ‘i

ae Akt

= > —€—

Ficalbia minima

SS.

—

Culex postspiracu/osus

TTR :

WK \

WAGs

y ISS

ONY

(NVA

NAAN:

; ..

‘. SS

| =

Orthopodomyia

anopheloides

Y SS ROK BR

of te => aS =e

Va IK SS Ss = RANTS

AA AES . Y

SSA

Ry AN NY

SSN

wes

SSS

i<SSS Ss

VBA. 3NWSWWN“{’

(Ce fj

Mansonia (Mansonioides) uniformis

crassipes

oes Zee)

Foy

dentatus

Armigeres (Leicesteria)

Mansonia (Coquillettidia)

Generalized mosquito pupa

balabacensis

‘

Topomyia

spathulirostris

Mansonia

Hodges/a

malayi

Mansonia (Mansonioides) (Coquillettidia)

uniformis microannulata

Topomyia

tipuliformis

- Tripteroides florengaui Tripteroides fuscipleura

Jopomyia

barbus

Bri

’ Tripteroides

stone/

HESS

Limatus

Aurhamii

Trichoprosopon

magnum

. “ Phoniomyia

pallidoventer

24a |

Malaya genurostris \ Topomyia spathulirostris\}

7 'M.Druckenbrod

}4 Oba | Phoniomyia

edwards/

Limatus

durhamii

Trichoprosopon

soares/

mete Sabethes —-

oe ' SOPeCr Be vi cae

o oe _Wyeomyia circumcincta

\\ Yy

\ :

\ V

Wyeomyia 4 ere SS

felicia ii case a eae

/ , ; \ \ . \ » 4 i: \ X / “ \ SS |

if HS \\ \SS

1 cans NS

MAAN i ee 2 Wy YN \

y / i \ ; ovil Ke AWA \

ST ante

AN MA Wi} 26d

Sabethes f' Trichoprosopon

purpureus// 77 ff | soares/

{1M Druckenbrod

S_=— 2

"8c

Opifex fuscus

chamberlain

Mimomyla

M. Druckenbrod

S Ke!

“ew

& co

% N

cy SeS

> Q

S =

RS)

AA}

aS)

(Ss)

S @&§

s ae

feb) S56

Q ==

Mimomyia (Etorleptiomyia) Minnis

/uzonensis (Ravanalites: Meneenee.

Uranotaenia ascidiicola

Uranotaenia modesta

Eretmapodites

dracaenae Aedes ( Stegomyia)

| ruwenzor!

Eretmapadites inornatus

Ahh Ag

> Se ee ss

Zenon ty

SS. :

ae =

Data ft N tra Of

Armigeres suba/batus Armigeres malayi Heizmannia_ achaetae

Udaya argyrurus

edes (Stegomyia ;

meronephada

Uf Ff “BRN 2

. . < A y *

Upp Al RTT

Dy” RRR

Heizmannia complex Heizmannia aureochaeta Aedes (Lorrainea) fumidus

Aedes (Diceromyia)

francisco!

: ! iii” Aedes

(Diceromyia) (Stegomyia) (Stegomyia) desmotes

periske/atus albopictus

Ficalbia circumtestacea (a mauehenerad Ficalbia malfeyti

—

Se eee ae * N

& 5

3S a

ae) >

S g

A e

> )

: :

aS 5)

9 <

(Se ~

(e) N

< S

,e)

Le)

ie)

Psorophora

infin

init

Haemagogus spegazz

Haemagogus capricornii Aedes aegypti

Culex antillummagnorum

M Druckenbrod Naan: Saas

ulex pipiens

Culex davisi

M. Oruckenbrod

Culiseta l/ongiareolata Culex bamborum

PI. 21

;

Aedes gill

‘

oy \y

Aedes aurantius

M.Dre

cA t

Saef ny

ese

U .

’

pce,

tris

ITOs

long

Culex

pseudomelanoconia

43b

—=an- ae LC

Toxorhynchites

splendens

q Se

f\ a

(| Q

pues

\ \ Se

ANN

NAN

Vie

Seay

Cty

é > 4

, BS

Sack cdl

ES h 7

Chagasia bathana

Bironella hollandi

46a

Sabethes

purpureus

eZ,

Limatus

durhamii

46b

Phoniomyia

fuscipes

a 0

La EES)

Trichoprosopon

after Lane, 1953

vill

, Hoy ty Wyeomyila

confusa

Trichoprosopon digitatum

NV!

— ta.

rai

Sa Ss

eon

—-__.

Topomyia gracilis

Malaya genurostris

(Acallyntrum)

— : >

—— ee

ie!

belkini

Mimomyila

(Ravenalites)

deguzmanae

<2)

1@)

“we

Tripteroides powell

53a 4

Topomyia tenuis

O3b & Me iy

el pe es ec Ret /

awa

Topomyia decorablis

Topomyia_ spathul/irostris

Vill

Airy 1

04b

Mimomyia hybrida

Mansonia uniformis

} : <2

: LO ae

L\ KA 4 ; eo FFE

On: gp my OOD

~ | $Olere Sf ire

: : eS 2 ae K

<i se AB'ge WN oe x

= se bt = ~~ SS

ike Pre hS % _

Y— WS Wee. S }

i We49.65.7— 3

¢ Sey ~ tered ot os

: aa ms A - OSS Sicanee pee .

: 2 e

Tees . > ; ee J

ak CSR SS \ gle 7

sy DSSS Se : a

Sis — SS OED yy

=> SF

cA \ x SS <> eee = ‘

A SSS S SS:

3 arg Oat RO SRS \ >

- oy SSNS SS a

CSS See BA \ \_ 55

z a ECBO SOY NOS NN SN 5

—= SS PrEee Se Si)

nes ete Ss OE Se ware Y

2 S CL Ea, SSS

SNe Ss = “a ae 7

7 eo CHT S27

Z LILES =F

SDS

SSS

Aedeomyia

catastica

Vill

———

‘

Wf),

Hy ie

CQ ~~ re

‘ae Qe. mee \ Wee aS i

SSS SSSA

nee

4 Se

FEN

Vill

SIS

AAS Siete ar

S R LAT a TAN Ne, an, epee PY ade Nae INE NN NNN EIN cc 8 Sig IE 8 Hp hia ae SR a, ae eft. aE TSR

CR EKK :

SSS we

Se OR A 622 7 ay

SSS Ss (ELLE DE MEG!

ULE a Ae gle

ZF PEL pd

CLS apy

LL LEE Mire

OEE Free |

Opifex fuscus

te N

M. (Ravenalites )

deguzmanae

Ficalbia minima

M. (Mimomyia)

plumosa

—=aS.- Fe CO

58a

M. (Mimomyia)

chamber/aini

: Z,

SS ——

Hodgesia malayi

— 2

“ae : s eee arg BN er

ian a ae er ne oe wis

Orthopodomyia

wilsoni

i a

60b

Culiseta /ongiareo/ata

Dejinocerites cancer

Oris

at4,e

YS i,

Armigeres suba/batus

sapphirina

Uranotaenia

sapphirina

Aedes (Stegomyia) annandalei

littleri

Vill

Culiseta

Aedes

aegypti

Eretmapodites

chrysogaster

64a

Udaya argyrurus Udaya /ucaris

Pl. 32

Zeugnomyia

agui/ari

Heizmannia

scinti/lans

N MAK,

Aedes

(Christophersiomyia)

gombakensis

Psorophora

howardi

a esis

Aedes (Finlaya)

poeci//us

PI. 33

/ i

6/c

Aedes dianteus

Psorophora_ signipennis

—

68a

Haemogogus capricornii

Aedes atlanticus

68b

Aedes serratus

eu Ze Zs

Y -

\" ce S

—=

an ¢W a

= YJ. r

= 3h Z

. 6

~St

cociige FO 2)

[S- ge

S—_—S

= =

==> —S=

vO == SS

Vill 4

4A LAY ny ARK H gee ee

Cit aii oe

Culex pipiens 69b

69c

Culex modestus

Contrib. Amer. Ent. Inst., vol. 7, no. 4, 1971

INDEX

Genera are in upper and subgenera in lower case.

note numbers which start on page 29.

In brackets are the

Figure numbers are underlined and a

number followed by a "k" refers to the page on which the genus keys out.

Acallyntrum (51)

AEDES 1, 2, 9k, 13k, 20k, 28k,(2, 11,

13,18, 27, 41, 42, 43, 44, 45,

46, 61, 62, 63, 64)

aegypti 40c, 61b, 63b

africanus (39) —

albopictus 36a

annandalei 33c, 62b

atlanticus

atropalpus (43)

aurantius 42d

desmotes 36b

diantaeus 6%c

franciscoi 35b

fumidus 35a

gilli A2b

gombakensis

harperi (61)

longirostris 42e

meronephada 33b

periskelatus 35c, (41)

poecilus 66b

polynesiensis 2

purpureipes (27)

ruwenzori 31c

Serratus 68b

AEDES 1

AEDEOMYIA 2, 7k, 11k, 17k, 24k, (29)

catasticta 9a, b, ¢, 2b, 55

AEDEOMYIA 1

AEDOMYIA 1

Alanstonea (11)

ANOPHELES 2, 3, 4k, 9k, 15k, 21k,(5,

29)

66a

albimanus 2

albitarsis 2

annularis 2

aquasalis 2

balabacensis 3b, 21b, 43b

barbirostris

bellator 2

eruzil 2

culicifacies 2

darlingi 2

fluviatilis 2

funestus 2

gambiae 2

hyrcanus 2

labranchiae 2

leucosphyrus (1)

maculatus 2

minimus 2

moucheti 2

nili 2

nuneztovari 2

pharoensis 2

pseudopunctipennis 2

pulcherrimus 2

punctulatus 2

sacharovi 2

sergentii 2

stephensi 2

Sundaicus 2

Superpictus 2

umbrosus

ARMIGERES _ 8k, 13k, 14k, 18k, 25k,(11,

16, 38, 57)

19b

32b, (40)

12a, b, 32a, 59a

dentatus

malayi

subalbatus —

Ayurakitia :

BIRONELLA 4k, 9k, 15k, 21k,(1, 5, 29)

hollandi 2,44

Cancraedes (18)

CHAGASIA 4k, 9k, 15k, 21k,(5)

bathana 2ic,43a

fajardoi 3a

Christophersiomyia (62)

Coquillettidia 1, (12,16, 24, 26, 30)

CULEX 2, 9k, 13k, 20k, 28k,(2, 13, 43,

45, 51,57) 15a

antillummagnorum™ 41b

bamborum 42b

belkini 51b

davisi 42a

dispectus (57)

modestus 69c

moucheti (5%)

pipiens 41a, Las

pipiens fatigans —

postspiraculosus cs ak 25)

pseudomelanoconia

CULISETA 1, 2, 6k, 10k, | “D1, 28k, (43, 46,

55, 58)

littleri 60c

longiareolata 41c,60b

melanura 69a (43)

DEINOCERITES 2, 7k, 12k, 16k, 25k

cancer lla,b, 60a

mcdonaldi 27a

Diceromyia (41)

ERETMAPODITES 2, 8k, 13k, 18k, 26k,

(57, 60)

chrysogaster 63a

Mattingly: Illustrated Keys to the Genera of Mosquitoes

dracaenae 31b

inornatus 3ia

Etorleptiomyia 1, (37

) 3

FICALBIA 1, 9k, 13k, 19k, 25k,(14,56) flavicosta 384

circumtestacea 37d

ORTHOPODOMYIA (08 ae 19k, 25k,

anopheloides 15b,19c

wilsoni 38b,59b

-malfeyti 37b Paraédes 1

minima I[1le,18b,57b PHONIOMYIA 6k, 11k, 16k, 22k,(7, 19,

Finlaya (42,64) ~ ee ;

GALINDOMYIA 1, 7k, 12k, 14, 21,(23) davisi 16c

leei 1l1c,d,17b edwardsi 25b

Grabhamia fuscipes 46b

(43)

HAEMAGOGUS 2, 7k, 12k, 20k, 28k,

(23, 38, 44, 63, 64)

capricornii 40b, 68a

chalcospilans (44) ~

Spegazzinii 40a

HARPAGOMYIA 1, (3)

HEIZMANNIA 7k, 12k, 19k, 27k,(9,

40, 41, 61, 62)

achaetae 32c

aureochaeta 34b, (41)

complex 34a

scintillans 17a, 33d, 65b

HODGESIA 5k, 10k, 15k, 25k,(2, 56)

malayi 6a,b, 22c,57a

Howardina (64) —

Janthinosoma (43)

Kerteszia 2

Kompia (27)

Leicesteria (27)

LIMATUS 2, 6k, 11k, 16k, 22k, (2, 34)

assuleptus 16a

durhami 16b, 23d, 25a, 47

MALAYA 1, 4k, 10k, 15k, 24k,(2, 3,5,

33, 52,53)

genurostris 5, 24a,50a

MANSONIA 1, 2,3, 8k, 9k, 13k, 14k,

15k, 24k, (11, 12, 30, 37,

crassipes 14d,18c

microannulata 22b

uniformis 13, 18a, 22a,54a

Maorigoeldia (6,52) —

Mattinglyia (9, 22

MEGARHINUS _ 1,(5)

MIMOMYIA 1, 9k, 13k, 17k, 25k,(14,

30, 31,37, 51, 54)

chamberlaini 28c,58a

deguzmanae 14c, 28e,51a,58c

hybrida 28b,54b

luzonensis 14h, 28d

perplexens 28a, (30)

plumosa 58

splendens (37)

OPIFEX ‘7k, 12k, 17k, 24k

fuscus 10a, b, 27c,56

pallidoventer 23e

PSOROPHORA 2, 6k, a 19k, 27k,(43,

; 63

ciliata 39a

ferox 39b-

howardi 67a

infinis 39c

Signipennis 67b

Ravenalites I,(37,51)

Rhynchotaenia (16, 24, 30)

SABETHES 2, 6k, 16k, 22k, (35, 36,

, 48

belisarioi 8a

purpureus 26c, 46a

Soperi 25d

Stegomyia (41,60, 61, 62)

Taeniorhynchus 1

THEOBALDIA | 1,(4)

TOPOMYIA _ 5k, 10k, 15k, 23k, (2, 3, 5, 33,

92,53

barbus 23b

decorabilis 23a, 53b

gracilis 50b

spathulirostris 22d, 24b, 53c, (53)

tenuis 9d3a

tipuliformis 22e

TOXORHYNCHITES 1, 2, 4k, 9k, 14k,

21k, (5)

splendens la,b,c, ala, 45

TRICHOPROSOPON 2, 6k, 11k, 16k, 22k,

ores ata

digitatum 3d,49a

frontosum 48

magnum 23f, 25e,(34, 36)

pallidiventer 8c

perturbans 16d —

soaresi 25c, 26d

TRIPTEROIDES 5k, 10k, 15k, 23k,(5, 6,

32,51, 52, 53)

(6, 52)

pee

argyropus

fuscipleura

lorengaui 22f

powelli 52a

stonei 23c,52b

UDAYA 8k, 13k, 18k, 27k,(11)

ee Contrib. Amer; Ent. Inst. , vol’7; no. 4, 1971

argyrurus 33a, 64a

lucaris 14a, 64b

URANOTAENIA 5k, 10k, 17k, 26k,(2,

18, 29, 37, 38, 57, 59)

acidiicola 30b

browni (57) —

colocasiae (57)

mendiolai 30a

modesta 30c _

sapphirina 6la,62a

WYEOMYIA 2, 6k, 11k, 16k, 22k, 23k,

(7, 8,19, 20, 35, 48, 49, 50)

aporonoma 3c

circumcincta 25f, (34)

codiocampa 26b

confusa 49b, (50)

felicia 26a

moerbista 8b

occulta, (50)

ZEUGNOMYIA _ 5k, 10k, 17k, 27k,(2)

aguilari 65a

gracilis

lawtoni 29

4, At i

anf - pei!

‘ 4 A :

tage

int lad

wre

tae

‘s

et a3 -

a) Ape ly

7’

* Sa

nit

ede ty

* ~ Whi ya

as i oe

)

en ee

Ketone

te “au

hea) ‘

LP a

‘a

pene Wot pvt hg coe

ge)

Deere

4 SD PL ae Mea he ee “hia Pye!

“Dod. of " “a i , j

Sunt Pind ann : ann)

ae ) i f Py a 5 wt [ ‘ Arahas

ire

Pieroni sera

Poe

A f. 4

ar 4)

=i i c \ aan

. ! e

(Continued from inside front cover)

Parts of Volume 4, with prices

Mosquito studies (Diptera, Culicidae), 1968.

X. Peters, T. Michael. Dixinae originally described from North America. 7 pages.

XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes

originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay.

29 pages. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XII. Berlin, O. G. W. A revision of the Neotropical subgenus Howardina of Aedes.

Carlson, Robert W. and Fred B. Knight, 1969.

Biology, taxonomy, and evolution of four sympatric Agrilus beetles (Coleoptera:

Buprestidae). 104 pages, 47 figures. Price: $3.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XII]. Barr, Ralph A. and Sylvia Barr. Pupae of the genus Psorophora in America

north of Mexico. 19 pages, 11 figures.

XIV. Berlin, O. G. W. Redescription of Toxorhynchites (T.) gigantulus from the

Philippines. 7 pages, 2 figures.

XV. Zavortink, Thomas J. A new species of treehole breeding Anopheles from

the southwestern United States. 12 pages, 4 figures.

Price: $1.25, postpaid.

Matthews, Robert W., 1970. A revision of the genus Spathius in America north of Mexico

(Hymenoptera, Braconidae). 86 pages, 31 figures. Price: $3.50, postpaid.

postpaid.

Parts of Volume 5, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XVI. Zavortink, Thomas J. A new species of treehole breeding Aedes (Ochlerotatus)

from southern California. 6 pages, 2 figures.

XVII. Adames, Abdiel J. and Charles L. Hogue. Two new species of Deinocerites from

Costa Rica. 6 pages, 6 figures.

XVIII. Berlin, O. G. W. The subgenus Micraedes of Culex. 30 pages, 12 figures, 1 map.

Price: $2.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XIX. Zavortink, Thomas J. The treehole Anopheles of the New World. 35 pages, 12

figures. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XX. Schick, Robert X. The Terrens Group of Aedes (Finlaya). 158 pages, 60 figures.

Price: $5.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

V. Reinert, John F. Genus Aedes, subgenus Diceromyia Theobald in southeast Asia.

43 pages, 15 figures. Price: $1.50, postpaid.

Howden, Anne T. The Tanymecini of the West Indies (Coleoptera: Curculionidae). 74 pages,

76 figures. Price: $2.75, postpaid.

Kethley, John B., 1970. A revision of the family Syringophilidae (Prostigmata: Acarina).

76 pages, 38 figures. Price: $2.75, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

VI. Mattingly, P. F. The genus Heizmannia Ludlow in southeast Asia. 104 pages,

51 figures. Price: $4.00, postpaid.

Parts of Volume 6, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXI. Belkin, John N., Sandra J. Heinemann, and William A. Page. The Culicidae of

Jamaica. 458 pages, 126figures. Price: $15.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

Vil. Tyson, W. H. Genus Aedeomyia Theobald in southeast Asia. 27 pages, 6 figures.

Vill. Tyson, W. H. Genus Aedes, subgenus Mucidus Theobald in southeast Asia. 54 pages,

21 figures. Price: $3.50, postpaid.

Parts of Volume 7, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXII. Zavortink, Thomas J. A new subgenus and species of Aedes from Arizona.

11 pages, 2 figures.

XXII. Schick, Robert X. Additions and corrections to the revision of the Aedes

terrens group. 27 pages, 8 figures. Price: $1.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1971.

XXIV. Adames, Abdiel J. A revision of the crabhole mosquitoes of the genus

Deinocerites. 154 pages, 58 figures. Price: $6.00, postpaid.

(Continued from inside back cover)

No. 3. Contributions to the mosquito fauna of southeast Asia, 1971.

IX. Zavortink, Thomas J. The genus Orthopodomyia Theobald in Southeast Asia.

37 pages, 12 figures.

X. Dobrotworsky, N. V. The genus Culiseta Felt in Southeast Asia. 24 pages, 8 figures.

XI. Sirivanakarn, Sunthorn. A proposed reclassification of Neoculex Dyar based

principally on the male terminalia. 24 pages, 3 figures. Price: $3.50, postpaid.

No. 4. Contributions to the mosquito fauna of southeast Asia, 1971.

XII. Mattingly, Peter F. Illustrated keys to the genera of mosquitoes (Diptera, Culicidae).

84 pages, 34 plates. Price: $3.50, postpaid.

No. 5. Mosquito studies (Diptera, Culicidae), 1971.

XXV. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes

originally described from Brazil. 64 pages. Price: $2.50, postpaid.

Contributions

of the

American Entomological Institute

Volume 7 Number 5, 1971

MOSQUITO STUDIES (Diptera, Culicidae)

XXV. Mosquitoes originally described from Brazil.

By John N. Belkin, Robert X. Schick, and

Sandra J. Heinemann.

SEP 16 19/71

CiBRARIE?

CONTRIBUTIONS

of the

AMERICAN ENTOMOLOGICAL INSTITUTE

The Contributions are for larger papers on insects. Each paper is a separate number, with separate

pagination and index. Separate numbers aggregating about 500 pages constitute a volume. Issues appear

irregularly, as suitable manuscripts are available.

Copies are sold separately or in subscriptions to complete volumes. Complete volumes are $12.00.

The price of separate numbers varies. Subscribers are billed for each volume with its beginning number,

and receive the parts as issued. Orders for separate numbers that total less than $8.00 must be accom-

panied by payment. Address orders or correspondence to the American Entomological Institute, 5950

Warren Road, Ann Arbor, Michigan 48105, U.S.A.

Parts of Volume 1, with prices

No. 1. Dasch, Clement E., 1964. The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae).

75 pages, 69 figures. Price: $2.25, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1965.

I. Belkin, John N., etal. A project for a systematic study of the mosquitoes of Middle America.

II. Belkin, John N., et al. Methods for the collection, rearing and preservation of mosquitoes.

78 pages, 4 figures. Price: $2.25, postpaid.

No. 2a. Same as no. 2, but in Spanish. Price: $2.25, postpaid.

No. 3. Matthews, Robert W., 1965. The biology of Heriades carinata Cresson. (Hymenoptera,

Megachilidae). 33 pages, 23 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1965.

Ill. Ramalingam, Shivaji, and John N. Belkin. Two new Aedes from Tonga and Samoa.

IV. Belkin, John N. The mosquitoes of the Robinson-Peabody Museum of Salem expedition to

the southwest Pacific, 1956.

34 pages, 3 figures. Price: $1.00, postpaid.

No. 5. Mosquito studies (Diptera, Culicidae), 1965.

V. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from Middle America. 95 pages. Price: $2.75 postpaid.

No. 6. Mosquito studies (Diptera, Culicidae), 1966.

VI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from North America. 39 pages. Price: $1.00, postpaid.

No. 7. DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae (Homopotera,

Cicadellidae). A synopsis of the genus Ponana.

86 pages, 257 figures. Price: $2.50, postpaid.

No. 8. Contributions to the mosquito fauna of southeast Asia, 1967.

I. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald in Thailand.

55 pages, 20 figures. Price: $1.75, postpaid.

Parts of Volume 2, with prices

No. 1. Contributions to the mosquito fauna of southeast Asia, 1967.

II. Bram, Ralph A. The genus Culex in Thailand (Diptera: Culicidae).

296 pages, 104 figures. Price: $9.00, postpaid.

No. 2. Schuh, Toby, 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region.

35 pages, 30 figures. Price: $1.25, postpaid.

No. 3. Snider, Richard J., 1967. The chaetotaxy of North American Lepidocyrtus s. str. (Collembola:

Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid.

No. 4. Contributions to the mosquito fauna of southeast Asia, 1968.

Ill. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald of southeast

Asia. 76 pages, 26 figures. Price: $2.25, postpaid.

No. 5. Contributions to the mosquito fauna of southeast Asia, 1968.

IV. Knight, Kenneth L. Species of the subgroup Chrysolineatus of group D, genus Aedes,

subgenus Finlaya Theobald. 45 pages, 12 figures. Price: $1.50, postpaid.

Parts of Volume 3, with prices

No. 1. Mosquito studies (Diptera, Culicidae), 1968.

Vil. Belkin, John N. The Culicidae of New Zealand.

182 pages, 30 figures. Price: $5.50, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1968.

VIll. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia.

221 pages, 41 figures. Price: $6.75, postpaid.

No. 3. Fleming, Richard C., 1968. Head musculature of sphinx moths (Lepidoptera: Sphingidae).

32 pages, 37 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1968.

IX. Belkin, John N. The type specimens of New World mosquitoes in European museums.

69 pages. Price: $2.00, postpaid.

(Continued on back cover)

MOSQUITO STUDIES (Diptera, Culicidae)

XXV. MOSQUITOES ORIGINALLY DESCRIBED

FROM BRAZIL

John N. Belkin, Robert X. Schick and Sandra J. Heinemann’

This is the fifth in a series of papers providing information on the source and loca-

tion of the original type material of mosquitoes described from the Americas (Bel-

kin, Schick and Heinemann, 1965, 1966, 1968; Peters, 1968). A sixth paper, with

corrections and additions up to the end of 1970 for the entire series, is in prepara-

tion. In addition to the species originally described from Brazil, the present paper

includes all the species originally described from South America without indication

of the country of origin; the type localities of all of these are here restricted to lo-

calities in Brazil. We are greatly indebted to the following individuals for assistance

to one of us (JNB) in locating type material in various institutions and for other

courtesies: T.H.G. Aitken, R.R. Correa, L.M. Deane, J.P. Duret, O.P. Forattini,

H. de Souza Lopes, B. Lutz, S. de Oliveira, E.X. Rabello, L.E. Rozeboom, A. Stone

and L.P. Travassos. We also thank Claire M. Price for the painstaking preparation of

the copy for lithoprinting.

For explanation of the arrangement and method of presentation in the sections

on Nominal Taxa and Localities, the first paper of the series should be consulted

(Belkin, Schick and Heinemann, 1965). The nominal taxa described up to the end

of 1969 are included. For the subfamily (family, auct.) Culicinae the listing follows

the taxonomic order and interpretation of the world catalog as modified by the

supplements (Stone, Knight & Starcke, 1959; Stone, 1961, 1963, 1967, 1970);

a few additional changes in synonymy are indicated. The subfamilies (families,

auct.) Chaoborinae and Dixinae follow, with the taxonomic interpretation adopted

by Stone (1966a, 1966b). For the localities, we have used as standard the gazetteer

of Brazil prepared by U.S. Board on Geographic Names (1963), except for the eli-

mination of all diacritic marks, accents and so on.

Lectotype designations are made here for the first time for several species with-

out previously designated holotype or lectotype; for the procedure followed see

Belkin (1968:2). We have also designated restricted type localities for the majority

of species without originally specified locality. In case the locality given was Rio

de Janeiro, Para or Bahia we have interpreted it to mean the city of Rio de Janeiro

‘Contribution from project ‘Mosquitoes of Middle America”’ supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478.

: Department of Zoology, University of California, Los Angeles, California 90024.

2 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

(Guanabara), Belem (Para) and Salvador (Bahia) respectively. For species described

from Brazil or South America without specified locality, we have been guided in

designating a locality by the source of other species described by the author in ques-

tion as well as by the known distribution of the species as currently interpreted. In

complex cases, an explanation is given in the section on Authors.

The section on Authors provides an index to the nominal species described by

every author and the location of the type material of these species. Discussions of

special problems relevant to the determination of the type material and type locali-

ties are also given..

In the section on Depositories is an alphabetical list of institutions abbreviated

as in the world catalog (Stone, Knight and Starcke, 1959); a few institutions have

been added. This section provides an index to the type material contained in each

institution and, in a few instances, discussions of special problems.

The section on References Cited provides full bibliographic entries for all litera-

ture citations given in the text. The citations of periodicals conform to the rules of

entry in ‘‘Anglo-American Cataloging Rules; North American Text” (Amer. Libr.

Ass., 1967) and the abbreviations follow the “‘Revised and Enlarged Word-A bbrevia-

tion List for American National Standard for Periodical Title Abbreviations” (Nat.

Clearinghouse Period. Title Word Abbreviations, 1966).

In the Index to Scientific Names, the number in parentheses following a specific

name refers to the number assigned to that name in the section on Nominal Taxa;

all other references are to page numbers.

NOMINAL TAXA

I. Chagasia fajardi (Lutz, 1904). Type: Female(s), Sao Paulo (Sao Paulo) (LU,

not in IOC). Bionomics: [Larvae in mats of aquatic vegetation along margins of

swift streams]. 7 |

*2. Chagasia neivae Cruz, 1906 [= fajardi|. Type: Female(s), Juiz de Fora (Minas

Gerais), C. Chagas (LU). Bionomics: [As for J. fajardi].

3. Chagasia maculata Peryassu, 1921 [= fajardi|. Type: Adult(s), in forest near

park in Cambuquira (Minas Gerais) (LU). Bionomics: [As for 1. fajardi]. |

4. Chagasia stigmopteryx Martini, 1932 [= fajardi|. Type: Holotype female, Bu-

tantan, Sao Paulo (Sao Paulo), 28 Mar 1920, R. Fischer (DEI). Bionomics: [As for

1. fajardi|. :

5. Chagasia rozeboomi Causey, Deane & Deane, 1944. Type: Egg(s), Ceara; Type

LOCALITY here restricted to vicinity of Sao Benedito (NE). Bionomics: [Larvae

probably in vegetation along margins of streams].

6. Anopheles (Stethomyia) lineatus (Lutz, 1905) [= nimbus]. Type: Holotype

male, probably near Sao Paulo (Sao Paulo) (possibly IOC, tubes 433,434,1066).

Bionomics: [Larvae probably in mats of algae along margins of shaded swift

streams |.

7. Anopheles (S.) lewisi Shannon, 1931; thomasi Shannon, 1933, new name.

Type: Holotype male, Rio Cururipe, near Salvador (Bahia) (USNM; see Stone and

Knight, 1956b:278). Bionomics: [Larvae in forest springs and streams].

8. Anopheles (A.) anchietai Correa & Ramalho, 1968. Type: Holotype male

(15856) with genitalia slide (3114) and slide of larval and pupal skins (3115),

Cangaiba, Sao Paulo (Sao Paulo) (FH). Bionomics: [Larvae probably in permanent

or semipermanent shaded ground waters].

Belkin et al: Topotypic Brazilian Mosquitoes 3

9. Anopheles (A.) bustamentei Galvao, 1955. Type: Holotype female (10909,

4112), Ribeirao Pequeno, municipio de Laguna (Santa Catarina), 13 June 1950,

J. Bento (FH). Bionomics: [Larvae probably in permanent or semipermanent ground

waters]. Females on animal bait. sae: :

| 10. Anopheles (A.) geometricus Correa, 1944 [= var. of eiseni]. Type: Males

larvae, pupae, eggs, Guaruja, Ilha de Santo Amaro (Sao Paulo) (NE). Bionomics:

[Larvae probably in shaded waters with vegetation and organic matter].

11, Anopheles (A.) evandroi Lima, 1937. Type: Holotype female (3040) with

slide of wing (3285), Sao Bento, Baixada Fluminense (Rio de Janeiro), Feb 1935,

Evandro Chagas (IOC). Bionomics: [Larvae probably in permanent or semiperma-

nent shaded ground waters].

12. Anopheles (A.) fluminensis Root, 1927. Type: Holotype male, Itaperuna

(Rio de Janeiro), 17 June 1925, F.M. Root (USNM). Bionomics: Larvae along

edges of a small brook.

13. Anopheles (A.) intermedius (Peryassu, 1908). Type: Females, Rio de Janeiro

(Guanabara) and Xerem (Rio de Janeiro), July (possibly in IOC, slide (3295) of

wing, 3045). Bionomics: [Larvae in shaded ground waters with dense vegetation;

forest ponds or pools].

*14. Anopheles (A.) maculipes (Theobald, 1903). Type: Holotype female, Sao

Paulo (Sao Paulo), A. Lutz (BM). Bionomics: [Larvae probably in permanent or

semipermanent shaded ground waters]. |

15. Anopheles (A.) mattogrossensis Lutz & Neiva, 1911. Type: Holotype female

(3530), Lagoa de Manicore (Mandicore) [?Mandiore] (Mato Grosso), Aug 1908,

J.C. Diogo (IOC). Bionomics: [Larvae in marshes, drainage ditches and small rain-

water pools in open country; lagoons in forests].

16. Anopheles (A.) amazonicus Christophers, 1923 [= mattogrossensis|. Type:

Holotype female, River Amazon, June 1915, A.A. Clark; TYPE LOCALITY here

restricted to Manaus (Amazonas) (BM). Bionomics: [As for 15. mattogrossensis] .

*17. Anopheles (A.) mediopunctatus (Theobald, 1903). Type: Holotype male,

Sao Paulo (Sao Paulo), A. Lutz (BM). Bionomics: [Larvae in sunlit, shallow fresh-

water pools with abundant vegetation; in forest ponds and pools].

18. Anopheles (A.) rockefelleri (Peryassu, 1923) [= mediopunctatus|. Type: Fe-

male(s), Brazil without locality specified, Mar and Apr; TYPE LOCALITY here re-

stricted to vicinity of Rio de Janeiro (Guanabara) (LU). Bionomics: [As for 17.

mediopunctatus | .

*19. Anopheles (A.) limai Fonseca & Ramos, 1939; costalimai Coutinho, 1944,

new name [= mediopunctatus]. Type: Holotype male (SPM33) with slides of wing

(3) and male genitalia (2), Sao Vicente (Sao Paulo), Aug 1939 (FH). Bionomics:

[As for 17. mediopunctatus | .

20. Anopheles (A.) costai Fonseca & Ramos, 1939 [= mediopunctatus|. Type:

Holotype male (SPM21) with slide of genitalia (1), Sao Vicente (Sao Paulo), June

1934 (FH). Bionomics: [As for 1 7. mediopunctatus].

21. Anopheles (A.) minor Lima, 1929. Type: Syntypes 1 male and | female

(tube 281 not 291) with slides of wings (673,890), Estrella [Imbarie] (Rio de

sine 14 Jan 1929 (IOC). Bionomics: [Larvae in ground waters with slow cur-

rent}. ,

*22. Anopheles (A.) lutzi (Cruz, 1907); peryassui Dyar & Knab, 1908, new name.

Type: Adult(s), shores of Rio Bicudo (Minas Gerais), June, C. Chagas (?10C, 1

slide with 2 wings). Bionomics: [Large bodies of sunlit clear water with abundant

emergent vegetation and some algae; partially shaded streams].

& Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

23. Anopheles (A.) alagoanii Peryassu, 1925 [= peryassui]. Type: Males and fe-

males, Mutange and Bom Parto, suburbs of Maceio, on shores of Lagoa Manguaba

(Alagoas), June-Sept (LU). Bionomics: Larvae in partially shaded ditches and

swamps. Adults fly rapidly and silently. |

24. Anopheles (A.) celidopus Dyar & Shannon, 1925 [= peryassui]. Type: Holo-

type female, Carmo, Rio Branco (Roraima), 1 Sept 1924, J. Bequaert (USNM, —

27747). Bionomics: [As for 22. lutzi]. |

25. Anopheles (A.) pseudomaculipes (Peryassu, 1908). Type: Adult(s), Xerem

nomics: [Larvae probably in permanent or semipermanent ground water].

26. Anopheles (A.) rachoui Galvao, 1952. Type: LECTOTYPE by present desig-

nation, female (1002) with 1 wing and 1 leg missing, Acude Sao Bento, municipio

Santo Amaro (Bahia), 19 Mar 1947, C. Azumbuja (FH, 10080). Bionomics: Larvae

in swamp. | 3 | :

27. Anopheles (A.) shannoni Davis, 1931. Type: Holotype female, Belem (Para),

Apr 1930, D.J. Crawford and N.C. Davis (USNM). Bionomics: [Larvae in forest

ponds and pools]. Adults on horse bait and alighting to feed in woods.

*28. Anopheles (A.) tibiamaculatus (Neiva, 1906). Type: Female(s), Oliveira

(Minas Gerais), May, C. Chagas (LU; not in IOC). Bionomics: [Larvae in rainpool

without vegetation (Davis, 1944), in road ruts and pools on margins of streams in

forest with cold water and without vegetation].

29. Anopheles (Nyssorhynchus) allopha (Peryassu, 1921) [= albitarsis]. Type:

Syntypes males and females, coastal lowlands in Rio de Janeiro (Guanabara) and

state of Rio de Janeiro (Museu Nac Rio de Janeiro). Bionomics: Larvae in swamps

and bromeliads (?).

30. Anopheles (N.) limai Galvao & Lane, 1937 [= albitarsis]. Type: Eggs, adult(s),

Pinheiros and Butantan, Sao Paulo (Sao Paulo) (NE; only slides of stomachs in

FMSP). Bionomics: Larvae in depressions; grassy sunlit rain pools; generally with

clear water but sometimes muddy; by Rio Pinheiros. |

31. Anopheles (N.) marajoara Galvao & Damasceno, 1942 [= albitarsis]. Type:

Males, females, larvae, Ilha do Marajo (Para); TYPE LOCALITY here restricted

to vicinity of Cachoeira do Arari (FMSP 958). Bionomics: [As for 30. limai].

32. Anopheles (N.) imperfectus Correa & Ramos, 1943 [= albitarsis]. Type:

Holotype female (22), Vera Cruz (Sao Paulo), G.R. Ramalho (FH). Bionomics:

[As for 30. limai]. Taken biting horse.

33. Anopheles (N.) domesticus Galvao & Damasceno, 1944 [= ssp. of albitarsis].

Type: Males, females, eggs, Cachoeira [?do Arari], Ilha do Marajo (Para) (LU).

Bionomics: [Larvae in borrow pits, brick pits, ditches, swamps, ponds, springs,

streams].

34, Anopheles (N.) antunesi Galvao & Amaral, 1940. Type: Holotype female

(370) with slide (235) of larval skin, Emilio Ribas, Campos do Jordao (Sao Paulo),

elev. ca 1570 m (FMSP). Bionomics: Larvae in shaded clear water with Sparse vege-

tation, in rockholes along river (Rio Capivari), in small drainage pools of fountain

Seale Simao) and in shaded pools of small streams. Adults taken at night biting

orse.

35. Anopheles (N.) emilianus Komp, 1941 [= aquasalis]. Type: Holotype male,

northern outskirts of Belem (Para), 14 Apr 1941, W.H.W. Komp (USNM). Bio-

nomics: Larvae in grassy pool. Holotype bred from larva.

36. Anopheles (N.) guarujaensis Ramos, 1942 [= aquasalis|. Type: Adults, larvae,

eggs, Guaruja (Sao Paulo), Mar 1939 (LU). Bionomics: [Larvae in brackish or fresh

water in tidal areas]. Adults in house.

Belkin et al: Topotypic Brazilian Mosquitoes 5

*37. Anopheles (N.) argyritarsis Robineau-Desvoidy, 1827. Type: Female(s), un-

specified locality in Brazil; TYPE LOCALITY here restricted to vicinity of Rio

de Janeiro (Guanabara) (NE). Bionomics: [Larvae in partly shaded or sunlit ground

pools, usually small].

38. Anopheles (N.) sawyeri Causey, Deane, Deane & Sampaio, 1943 [= ssp. of

argyritarsis]. Type: Holotype female, plateau of Serra da Ibiapaba, near Sao Bene-

dito (Ceara) (USNM). Bionomics: [Larvae in mountain forest pools (Deane, Causey

and Deane, 1946:45)]. unre tli bred from egg laid by female captured on horse

bait.

39. Anopheles (N.) braziliensis (Chagas, 1907). Type: Syntypes 6 females, mar-

gins of Rio das Velhas (Minas Gerais), 11 July 1907; TYPE LOCALITY here re-

stricted to vicinity of Lassance, see Root, 1926:704 (IOC). Bionomics: [Larvae

in sunlit clear water with abundant emergent vegetation and algae]. .

40. Anopheles (N.) pessoai Galvao & Lane, 1937 [= braziliensis|. Type: Holotype

female (82,25), left bank of Rio Pinheiros, Pinheiros, Sao Paulo (Sao Paulo), G.R.

Ramalho (FMSP). Bionomics: Larvae in small, shallow, sunlit pools with abundant

grass and algae. Holotype bred from larva.

41. Anopheles (N.) darlingi Root, 1926. Type: Lectotype male, Caxiribu, near

Porto das Caixas (Rio de Janeiro), 12 Mar 1925 (USNM; designation by Stone,

1943:30). Bionomics: Larvae in patches of Ceratophyllum in small side bays and

along edges of small river and canal with rather rapid current.

42. Anopheles (N.) paulistensis Galvao, Lane & Correa, 1937 [= darlingi|. Type:

Adults, eggs, Novo Oriente, 3 km from Lussanvira, near Pereira Barreto (Sao Paulo),

Apr-May, 1937 (LU). Bionomics: [As for 41. darlingi].

43. Anopheles (N.) strodei Root, 1926 [= evansae]. Type: Lectotype male (64),

a few miles from Agua Limpa Station, near Juiz de Fora (Minas Gerais), 27 Mar

1925 (USNM; designation by Stone and Knight, 1956b:280). Bionomics: Larvae

in small marshy expansions of mountain streams, side pools of a river and in small

_ pools, marshes and seepage areas with abundant vegetation.

44. Anopheles (N.) ramosi Unti, 1940 [= var. of evansae]. Type: Larvae, Lorena

(Sao Paulo), Mar 1940 (NE). Bionomics: Larvae in shade in extensive swamps with

vegetation, principally ““Aguape”’ and “‘Vassoura do brejo”’.

45. Anopheles (N.) arthuri Unti, 1941 [= var. of evansae]. Type: Eggs, valley of

Rio Paraiba [do Sul] (Sao Paulo), elev. ca 500 m (NE). Bionomics: [As in 43.

strodei].

46. Anopheles (N.) artigasi Unti, 1941 [= evansae var. arthuri|. Type: Eggs,

valley of Rio Paraiba [do Sul] (Sao Paulo), elev. ca 500 m (NE). Bionomics: [As for

43. strodei).

47. Anopheles (N.) albertoi Unti, 1941 [= var. of evansae}]. are Eggs, valley

of Rio Paraiba [do Sul] (Sao Paulo), elev. ca 500 m (NE). Bionomics: [As for

43. strodei|.

48. Anopheles (N.) galvaoi Causey, Deane & pee, 1943. Type: LECTOTYPE

female (R426) by present designation, marked capanemai in register, Rio Branco

(Acre), O.R. Causey (FMSP, 694). Bionomics: [Larvae in small pools with algae»

and grass and in swamps]. Adults reared from eggs.

49. Anopheles (N.) lanei Galvao & Amaral, 1938. Type: Holotype female (86-1,

220), Emilio Ribas, near Campos do Jordao (Sao Paulo), elev. ca 1570 m, Feb-Mar,

1938 (FMSP). Bionomics: Larvae in clear permanent waters. Adults strongly zoo-

philic.

50. Anopheles (N.) lutzii Cruz, 1901. Type: Syntypes 3 females (1965) in tube

6 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

993 and slide (2111) of wing, Lagoa Rodrigo de Freitas, Rio de Janeiro (Guanabara)

(IOC). Bionomics: [Larvae in swamps].

*51. Anopheles (N.) niger (Theobald, 1907) [= lutzii]. Type: Lectotype female,

Cantareira (Sao Paulo), 9 Nov 1904, A. Lutz (BM; designation by Belkin, 1968:10).

Bionomics: [As for 50. lutzii].

52. Anopheles (N.) guarani Shannon, 1928 [= lutzii]. Type: Holotype female,

Iguacu (Parana), 5 Oct 1927, R.C. and E.M. Shannon (USNM). Bionomics: [As for

50. lutzii).

53. Anopheles (N. ) nigritarsis (Chagas, 1907). Type: Female(s), [Oliveira (Minas

Gerais)] (from Peryassu, 1908:60) (LU). Bionomics: [Larvae probably in swamps

or forest pools].

54. Anopheles (N.) noroestensis Galvao & Lane, 1937. Type: LECTOTYPE by

present designation, male genitalia slide (2177;10293), adult apparently lost, (pos-

sible syntype FMSP, 343), Novo Oriente [Pereira Barreto], near Lussanvira (Sao

Paulo) (FH). Bionomics: [Larvae in sunlit or partially shaded pools].

55, Anopheles (N.) ayrozai Unti, 1940 [= noroestensis]. Type: Female(s), lar-

va(e), eggs, Guaratingueta (Sao Paulo), Nov 1939-Aug 1940 (NE; not in FH or

SPM). Bionomics: Larvae in sunlit, clear water, shallow pools; wells and ditches

with slightly colored or muddy water.

56. Anopheles (N.) goeldii Rozeboom & Gabaldon, 1941 [= nuneztovari]. Type:

Holotype male, Boa Vista, Rio Tapajos (Para), C.H.T. Townsend (USNM). Biono-

mics: [Larvae in muddy pools and small lagoons, sunlit or partially shaded].

57. Anopheles (N.) dunhami Causey, 1945 [= nuneztovari|. Type: Holotype male

with genitalia slide, Tefe (Amazonas) (USNM). Bionomics: [As for 56. goeldii].

58. Anopheles (N.) oswaldoi (Peryassu, 1922). Type: Syntypes males and females,

Vale do Rio Doce (Espirito Santo) and Baixada Fluminense (Rio de Janeiro), Mar

and Apr (Museu Nac Rio de Janeiro). Bionomics: [Larvae in shaded fresh water in

swamps, pools or stagnant streams] .

59, Anopheles (N.) metcalfi Galvao & Lane, 1937 [= oswaldoi]. Type: LECTO-

TYPE by present designation, male (96) with genitalia on slide (710210-1), Porto

das Caixas (Rio de Janeiro), 29 May 1925, F.M. Root; 1 of several specimens identi-

fied as tarsimaculatus by Root (1926:711), on which Galvao and Lane based their

metcalfi (USNM). Bionomics: Larvae in thick vegetation or flotage of quiet pools,

ponds, marshes, lagoons, side-bays or edges of rivers.

60. Anopheles (N.) konderi Galvao & Damasceno, 1942 [= oswaldoi]. Type:

Male(s), larva(e), pupa(e), Coari (Amazonas) (LU, no material in FMSP, 651; ap-

parently lost). Bionomics: [As for 58. oswaldoi].

_ 61, Anopheles (N.) parvus (Chagas, 1907). Type: Male(s) and female(s), [Oliveira

(Minas Gerais) (Galvao, 1941:537)] (?10C,1966; female (tube 994) and slide (2112)

of wing, both without data). Bionomics: [Larvae in small, shaded forest pools].

62. Anopheles (N.) rondoni (Neiva & Pinto, 1922). Type: LECTOTYPE by pre-

sent designation, large wing mounted on slide (403) together with wing of type of

cuyabensis, marked Matto Grosso, C. Pinto, remainder of adult apparently lost;

TYPE LOCALITY restricted here to Ladario (Mato Grosso) on Rio Paraguai (IOC).

Bionomics: [Larvae in small areas of clear water in ditches and marshes among

aquatic vegetation |.

63. Anopheles (N.) triannulatus (Neiva & Pinto, 1922). Type: Holotype female,

Fazenda Sao Joao, right bank of Rio Cuiaba (Mato Grosso) (LU; not in IOC). Bio-

nomics: [Larvae in vegetation in freshwater pools, lakes and river margins].

Belkin et al: Topotypic Brazilian Mosquitoes 7

64. Anopheles (N.) cuyabensis (Neiva & Pinto, 1923) [= triannulatus|. Type:

LECTOTYPE by present designation, small wing mounted on slide (403) together

with wing of type of rondoni, marked Matto Grosso, C. Pinto, remainder of adult

apparently lost; type locality, Fazenda Sao Joao, right bank of Rio Cuiaba (Mato

Grosso) (IOC). Bionomics: [As for 63. triannulatus].

65. Anopheles (N.) chagasi Galvao, 1941 [= triannulatus]. Type: Females and

eggs, Chaves (Para), Itacoatiara and Manaiis (Amazonas); TYPE LOCALITY here

restricted to Manaus (LU). Bionomics: [As for 63. triannulatus] .

66. Anopheles (Kerteszia) lutzii Theobald, 1901; cruzii Dyar & Knab, 1908,

adolphoi Neiva, 1908, new names. Type: Lectotype female, Rio de Janeiro (Guana-

bara), 4 July 1899, A. Lutz (BM, designation by Belkin, 1968:10). Bionomics:

[Larvae in bromeliads] .

67. Anopheles (K.) montemor Correa, 1950 [= cruzii]. Type: Holotype male

(102) with slides of genitalia (15,G8D1) and larval and pupal skins (15,G8D1),

Caraguatatuba (Sao Paulo), G.R. Ramalho and J. Germano (FH). Bionomics: [Lar-

vae in bromeliads].

68. Anopheles (K.) laneanus Correa & Cerqueira, 1944 [= ssp. of cruzii]. Type:

Holotype male (383.H.13) with genitalia slide (640), Campos do Jordao (Sao Paulo),

elev. 1600 m, J. Lane and F. Lane (FH;2226 in tube 2227). Bionomics: [Larvae

in bromeliads].

*69. Anopheles (Lophopodomyia) gilesi (Peryassu, 1908). Type: Holotype male,

Rio das Velhas (Minas Gerais), June; TYPE LOCALITY here restricted to vicinity

of Lassance (LU). Bionomics: [Larvae in shaded, cool, clear water in mountain

forest streams].

70. Anopheles (L.) pseudotibiamaculatus Galvao & Barretto, 1941. Type: Holo-

type female (380,438), Casa Grande, municipio Mogi das Cruzes (Sao Paulo), 5

Sept 1940, M. Pereira Barretto (FMSP). Bionomics: [Larvae in shaded streams and

pools].

71. Toxorhynchites (Ankylorhynchus) catharinensis (Lima, Guitton & Ferreira,

1962). Type: Holotype female (5553) with slide (4869) of associated pupal skin,

Brusque (Santa Catarina), R. Rachou (IOC). Bionomics: [Larvae in bromeliads] .

72. Toxorhynchites (A.) purpureus (Theobald, 1901). Type: Lectotype female,

Amazon, 1861, H.W. Bates; TYPE LOCALITY here restricted to Manaus (Ama-

zonas) (BM; designation by Belkin, 1968:34). Bionomics: [Larvae in bromeliads] .

73. Toxorhynchites (A.) trichopygus (Wiedemann, 1828). Type: Syntypes 3

males, Brazil, locality not specified, Freireiss; TYPE LOCALITY here restricted to

Salvador (Bahia) (SNG; see Belkin, 1968:34). Bionomics: [Larvae in bromeliads].

74. Toxorhynchites (A.) neglectus (Lutz, 1904) [= trichopygus]. Type: Holotype

female, near Sao Paulo (Sao Paulo) (NE). Bionomics: Bred from larva in bromeliad.

75. Toxorhynchites (Lynchiella) bambusicola (Lutz & Neiva, 1913). Type: Syn-

types males and females, Petropolis (Rio de Janeiro), elev. 800-900 m, J.G. Foet-

terle et al (IOC). Bionomics: Larvae in bamboo internodes (taquaracu [Guadia

sp.]|).

76. Toxorhynchites (L.) horei (Gordon & Evans, 1922) [identity uncertain].

Type: Lectotype male (463) with genitalia on 2 slides, Macapa, near Manaus (Ama-

zonas), 21 Dec 1921, R.M. Gordon (BM; designation by Belkin, 1968:33). Bio-

nomics: Larvae in axils of “bananeira brava’’ [Heliconia sp.].

77. Toxorhynchites (L.) mariae (Bourroul, 1904). Type: Female(s), larva(e), Ilha

de Itaparica (Bahia) (NE). Bionomics: Larvae in bromeliads.

78. Toxorhynchites (L.) pusillus (Lima, 1931). Type: Syntypes 1 male (541),

8 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

1 female (526) with slides of associated larval skins (1102,1121) and pupal skins

(1116,1135), Alto da Boa Vista, Tijuca, Rio de Janeiro (Guanabara), Apr 1930,

C.A. Campos Seabra (IOC). Bionomics: Larvae in bamboo internodes.

79. Toxorhynchites (L.) solstitialis (Lutz, 1904). Type: LECTOTYPE by present

designation, female marked as type by Theobald (data in Belkin, 1968:34), Sao

Paulo (Sao Paulo), 7 Oct 1903 (BM). Bionomics: Larvae in bromeliads, principally

“Aechmea tinctoria’’.

80. Toxorhynchites (L.) chrysocephalus (Theobald, 1907) [= solstitialis]. Type:

Holotype male, Sao Paulo (Sao Paulo), 17 Aug 1903, A. Lutz (BM). Bionomics:

[As for 79. solstitialis | .

81. Toxorhynchites (L.) ferox (Wiedemann, 1828); wiedemanni Dyar & Knab,

1906, new name. Type: Syntypes several males, Brazil, without specified locality;

TYPE LOCALITY here restricted to Salvador (Bahia) (SNG and NMW; see Belkin,

1968:33). Bionomics: [Larvae probably in treeholes or bamboo].

82. Toxorhynchites (L.) posticatus (Lutz & Neiva, 1913) [?= wiedemanni].

Type: Syntypes 2 females (549,550) and possibly 1 male (540), Petropolis (Rio

de Janeiro) (IOC). Bionomics: Larvae in bromeliads.

83. Toxorhynchites (L.) violaceus (Wiedemann, 1820). Type: Lectotype male,

Bahia [Salvador (Bahia)] (NMW; designation by. Belkin, 1968:34). Bionomics: Hoare

vae in bromeliads].

84. Toxorhynchites (L.) ambiguus (Dyar & Knab, 1906) [identity uncertain].

Type: Holotype male, Brazil, locality not specified, coll. Winthem; TYPE LOCA-

LITY here restricted to Salvador (Bahia). (LU; originally in Winthem Collection

in Hamburg, possibly now in NMW; see Belkin, 1968:32). Bionomics: [Larvae pro-

bably in treeholes, bamboo or bromeliads].

85. Toxorhynchites (L.) fluminensis (Peryassu, 1908) [identity uncertain]. Type:

Described from unspecified number of males, females and larvae, Rio de Janeiro

(Guanabara) (possibly IOC, male (tube 535), with genitalia slide (1128), Rua Conde

de Bonfim, Rio de Janeiro, 16 Apr 1907, C. Chagas leg.). Bionomics: Larvae in

bromeliads.

86. Trichoprosopon (T.) compressum Lutz, 1905. Type: Syntypes males and fe-

males, states of Sao Paulo and Rio de Janeiro; following material in box 13 possi-

bly part of type series: 1 female (2653), Sao Paulo; 1 male without genitalia (2651),

1 female (2652), Petropolis; 1 female (2654), 1 adult (2655), without data; 2 male

genitalia slides (1313,1314), “da coll. do Dr. Lutz’? OC). Bionomics: Larvae in

bamboo.

87. Trichoprosopon (T.) digitatum (Rondani, 1848). Type: Female(s), Brazil lo-

cality not specified; TYPE LOCALITY here restricted to vicinity of Rio de Janeiro

(Guanabara) (LU; see Belkin, 1968:35). Bionomics: [Larvae in cacao pods, coconut

husks and fallen palm spathes].

88. Trichoprosopon (T.) splendens Lutz, 1904 [= digitatum]. Type: Adult(s),

Manaus (Amazonas) (NE). Bionomics: [As for 8&7. digitatum].

89. Trichoprosopon (T.) townsendi Stone, 1944 [= var. of digitatum|. Type:

Holotype male (56740), Boa Vista, Rio Tapajos (Para), 20-22 July, C.H.T. Town-

send (USNM). Bionomics: [Probably as for 87. digitatum] .

90. Trichoprosopon (T.) obscurum Lane & Cerqueira, 1942. Type: Holotype

male, Mangaratiba (Rio de Janeiro), Apr-Oct 1938 (IOC). Bionomics: [Larvae pro-

bably j in bamboo].

*91. Trichoprosopon (T.) pallidiventer (Lutz, 1904). Type: LECTOTYPE by pre-

sent designation, male dissected on slide, marked Holoconops pallidiventer Lutz by

Belkin et al: Topotypic Brazilian Mosquitoes 9

Theobald, Sao Paulo (Sao Paulo), A. Lutz (BM; see Belkin, 1968:36-37). Bionomics:

[Larvae in bamboo internodes].

92. Trichoprosopon (T.) soaresi Lane & Cerqueira, 1942. Type: Holotype male,

Sao Joao de Petropolis (Espirito Santo) (IOC). Bionomics: [Larvae in bamboo inter-

nodes].

*93. Trichoprosopon (Limamyia) brevipes (Lima, 1931). Type: LECTOTYPE by

present designation, male (tube 462) with genitalia on slide (1029) and larval and

pupal skins on slides, apparently mixed with those of female, Laranjeiras, Rio de

Janeiro (Guanabara), Jan 1930, J.F. Ladeiras (IOC). Bionomics: Larvae in bamboo

internode. 7 | :

#94. Trichoprosopon (Shannoniana) fluviatile (Theobald, 1903). Type: Holotype

female, marked by Theobald Heleconops longipalpis Lutz, Sao Paulo (Sao Paulo),

A. Lutz (BM; see Belkin, 1968:35-36). Bionomics: [Larvae in bored internodes of

bamboo or ‘“‘taquarucu” (Chusquea gaudichaudii)].

95. Trichoprosopon (S.) longipalpis (Lutz, 1905) [= fluviatile]. Type: Syntypes

male(s) and female(s), [Sao Paulo (Sao Paulo)] (IOC, possibly male (2635) with

genitalia slide, 1315). Bionomics: [As for 94. fluviatile].

96. Trichoprosopon (Isostomyia) luederwaldti (Lane, 1936). Type: Holotype fe-

male (439), Fazenda Jose Euphrasio, Avare (Sao Paulo), 21 Mar 1936, J. Lane and

F. de Andrade (FH,426 not 428). Bionomics: [Larvae probably in leaf axils of

Araceae or leaf axils and flower bracts of Musaceae]. | :

97. Trichoprosopon (Ctenogoeldia) walcotti Lane & Cerqueira, 1942. Type: Holo

type female, Caravelas (Bahia), Jan 1931, N.C. Davis (IOC). Bionomics: [Larvae

probably in leaf axils or flower bracts of Marantaceae or Musaceae].

98. Trichoprosopon (Runchomyia) castroi Lane & Cerqueira, 1942. Type: Holo-

type male, Teresopolis (Rio de Janeiro), Mar-May, 1938 (IOC). Bionomics: [Larvae

probably in bromeliads or leaf axils and flower bracts of other water holding

plants] |

99. Trichoprosopon (R.) cerqueirai Stone, 1944. Type: Holotype male (57190),

Rio de Janeiro (Guanabara), Feb 1940, L. Whitman (USNM). Bionomics: [Larvae

in bromeliads]. |

100. Trichoprosopon (R.) edwardsianum Lane & Cerqueira, 1942. Type: Holo-

type female, Belterra, Santarem (Para), Sept 1938 (IOC). Bionomics: [Larvae in

crown of palms, Mauritia sp.].

101. Trichoprosopon (R.) humboldti Lane & Cerqueira, 1942. Type: Holotype

male, Paineiras, Rio de Janeiro (Guanabara), Aug 1939 (IOC). Bionomics: [Larvae

probably in bromeliads or leaf axils and flower bracts of other water holding

plants].

102. Trichoprosopon (R.) trichopus (Dyar, 1919) [= longipes|. Type: Holotype

female (21996), Tefe (Amazonas), June 1906, Ducke (USNM). Bionomics: [Larvae

in leaf axils of Montrichardia arborescens (Araceae), Kumm and Novis, 1938:512].

*103. Trichoprosopon (R.) lunatum (Theobald, 1901). Type: Lectotype female,

~Maua, Rio de Janeiro (Guanabara), 22 July 1899, C. Moreira (BM; designation by

Belkin, 1968:36). Bionomics: [Larvae in leaf axils of Musaceae and Araceae].

104. Trichoprosopon (R.) reversum Lane & Cerqueira, 1942. Type: Holotype

male, Mage (Rio de Janeiro), July 1940 (IOC). Bionomics: Larva in bromeliad.

105. Trichoprosopon (R.) simile Lane & Cerqueira, 1942. Type: Holotype male,

Campos do Jordao (Sao Paulo), Dec 1935 or Jan 1936, F. Lane (IOC). Bionomics:

_ [Larvae probably in bromeliads or leaf axils and flower bracts of other water holding

plants].

10 _ Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

106. Trichoprosopon (R.) theobaldi Lane & Cerqueira, 1942. Type: Holotype

male, Mage (Rio de Janeiro), June 1940 (IOC). Bionomics: [Larvae in bromeliads.

in Trinidad (Lane, 1945:133)].

107. Wyeomyia (W. ) arthrostigma (LZ y LOW). ao¥ Des LECTOTYPE by present

designation, only remaining specimen, female without locality label but with hand-

written label, “‘Proveniente de larva de taqueras, (larva os predador e desenhada)”’

[Sao Paulo (Sao Paulo), Peryassu, 1908:75] (IOC). Bionomics: Larvae in bamboo.

108. Wyeomyia (W.) downsi Lane, 1945. Type: Holotype female (608), Mirassol

(Sao Paulo), Jan 1936, Andrade and Antenor (FH,4043). Bionomics: [Larvae pro-

bably in leaf axils or flower bracts of plants or in bamboo].

females, Boa Esperanca and Pocinho (Mato Grosso), 19 Aug-6 Sept (LU; not in FH).

Bionomics: [Larvae in cut bamboo]. Females attracted to humans in forest in day-

time. |

110. Wyeomyia (W.) limai Lane & Cerqueira, 1942. Type: Holotype male, Lon-

drina (Parana), Sept 1936, Jan or Feb 1937 (IOC). Bionomics: [Larvae probably

in bamboo].

I11. Wyeomyia (W.) lutzi (Lima, 1930). Type: Syntypes, several males and fe-

males, with larval and pupal skins and genitalia slides, Alto da Boa Vista, Tijuca,

Rio de Janeiro (Guanabara), 28 Feb, 20 Apr 1930, C.A. Campos Seabra (IOC;

tubes 472,513,514; slides 1058,1066,1084,1096-1101,1103,1104,1118,1119,1171-

1176). Bionomics: Larvae in bamboo internodes.

112. Wyeomyia (W.) medioalbipes Lutz, 1904. Type: LECTOTYPE by present

designation, male with attached genitalia mount, rest of abdomen glued on another

attached mount with red type label, [garden of Hospital de Santa Isabel], Bahia

[Salvador] (Bahia) (BM; see Belkin, 1968: 40-41). Bionomics: Reared from larvae

collected in bromeliads.

113. Wyeomyia (W.) oblita (Lutz, 1904). Type: Syntypes male and female (tube

1544), with genitalia on slides (4599,4600), [Pacaembu], Sao Paulo (Sao Paulo),

11 Oct 1904, [A. Lutz] (IOC); female (BM; see Belkin, 1968:41). Bionomics:

Males reared from larvae in “‘tabuas”’ (Typha), Lutz, 1905:271.

114. Wyeomyia (W.) pintoi (Lima, 1930) [?= oblita]. Type: Holotype female

(618), Mendes (Rio de Janeiro), fazenda of Dr. Olympio da Fonseca (IOC). Bio-

nomics: Larvae in bamboo internodes.

115. Wyeomyia (W.) sabethea Lane & Cerqueira, 1942. Type: Holotype male,

Teresopolis (Rio de Janeiro), Apr 1938 (IOC). Bionomics: [Larvae probably in

bamboo internodes]. :

116. Wyeomyia (W.) serrata (Lutz, 1905). Type: LECTOTYPE by present desig-

nation, male, marked as 1 of 2 “‘cotypes”’ by Costa Lima, with genitalia slides (1167

and 1168) and possibly slide of leg (1169) and wing (1170), R. Frio, Pindamon-

hangaba, Sao Paulo (Sao Paulo), 9-12 Feb 1905, A. Lutz (IOC). Bionomics: [Larvae

probably i in bamboo].

*117. Wyeomyia (Cruzmyia) dyari Lane & Cerqueira, 1942. Type: Lectotype male

(1363) with slides of genitalia (355) and midtarsus (356), Petropolis (Rio de Jan-

eiro), May 1938, R.C. Shannon (FH; designation by Lane and Cerqueira, 1958a).

Bionomics: Larvae in bromeliads.

118. Wyeomyia (C.) kummi Lane & Cerqueira, 1942. Type: Holotype female,

Curralinho (Para), Jan, Feb 1936, H.W. Kumm (IOC). Bionomics: [Larvae probably

in bromeliads].

119. Wyeomyia (C.) mattinglyi Lane, 1953. Type: Female(s), state of Bahia;

TYPE LOCALITY here restricted to vicinity of Salvador (LU; not in IOC or FH).

Bionomics: [Larvae probably in bromeliads] .

*120. Wyeomyia (Menolepis) leucostigma Lutz, 1904. Type: Adult(s), Sao Pauls

(Sao Paulo) (NE). Bionomics: Larvae in “‘tabuas’”’ (Typha).

Belkin et al: Topotypic Brazilian Mosquitoes 11

*121. Wyeomyia (Antunesmyia) rooti Lane & Cerqueira, 1942; alani Lane & Cer-

.queira, 1957, new name. Type: Holatype male, Rio de Janeiro (Guanabara), Oct

1940 (IOC). Biohbmiee: [Larvae probably in bamboo].

122. Wyeomyia (Dendromyia) airosai Lane & Cerqueira, 1942. Type: Holotype

male, Santa Teresa (Espirito Santo), May 1940 (IOC). Bionomics: [Larvae probably

in Bronicliads|

123. Wyeomyia (D.) bourrouli (Lutz, 1905). Type: Male(s) and female(s), Esta-

cao de Itaici (Sao Paulo) (NE). Bionomics: Larvae in bromeliads.

124. Wyeomyia (D.) cesari Del Ponte & Cerqueira, 1938. Type: Holotype female

(2209), Cuiaba (Mato Grosso), Mar 1935, G. Cesar (IOC). Bionomics: [Larvae pro-

bably in leaf axils].

#125. Wyeomyia (D.) confusa (Lutz, 1905). Type: Syntype females, woods near

Sao Paulo (Sao Paulo) (NE). Bionomics: [Larvae probably in leaf axils or flower

bracts of Araceae, Marantaceae and Musaceae, and possibly bromeliads].

126. Wyeomyia (D.) finlayi Lane & Cerqueira, 1942. Type: Holotype male, Xe-

rem (Rio de Janeiro), June 1940 (IOC). Bionomics: [Larvae probably in brome-

liads].

127. Wyeomyia (D.) howardi Lane & Cerqueira, 1942. Type: Holotype male,

Muriqueira [Gois Calmon] (Bahia), May 1929, R.C. Shannon (IOC). Bionomics:

[Larvae probably in bromeliads].

128. Wyeomyia (D.) kerri Del Ponte & Cerqueira, 1938. Type: Holotype female

(2210), Cuiaba (Mato Grosso), Feb-July 1935, G. Cesar (IOC). Bionomics: larvae

in ‘“‘palma de assaizeiro” (Garapa guyanensis Hubl).

129. Wyeomyia (D.) knabi Lane & Cerqueira, 1942. Type: Holotype male, Ca-

choeira (Rio de Janeiro), May 1938 (IOC). Bionomics: [Larvae probably in leaf

axils].

*130. Wyeomyia (D.) luteoventralis Theobald, 1901. Type: Lectotype female,

[Belem] (Para), H.E. Durham (BM; designation by Belkin, 1968:40). Bionomics:

[Larvae probably in bromeliads].

131. Wyeomyia (D.) melanoides (Root, 1928) [= melanocephala|. Type: Lecto-

type male (90-2) with genitalia slide and associated pupal skin, Mage (Rio de Jan-

eiro), 26 May 1925 (USNM,44165; designation by Stone and Knight, 1957b:124).

Bionomics: [Larvae probably in leaf axils of Araceae, Musaceae or Typhaceae].

132. Wyeomyia (D.) mystes Dyar, 1924. Type: Lectotype male with genitalia

slide (36.VII.28b), Rio de Janeiro (Guanabara), Aug 1922, F.L. Soper (USNM;

designation by Stone and Knight, 1957b:124). Bionomics: [Larvae in leaf axils, of

aroids (Dyar, 1928:72) or bromeliads (Shannon, 1931:6)].

133. Wyeomyia (D.) negrensis Gordon & Evans, 1922. Type: Lectotype male

(10-5/643) with 2 slides of genitalia, Macapa, near Manaus (Amazonas), 20 Dec

‘1921, R.M. Gordon (BM; designation by Belkin, 1968:41). Bionomics: Larvae in

es bananeira brava’”’ (Heliconia sp.).

134. Wyeomyia (D.) personata (Lutz, 1904). Type: LECTOTYPE by present

designation, male with attached genitalia mount, marked “Type selected by J.

Lane”, Cantareira (Sao Paulo) (BM; see Belkin, 1968:41). Bionomics: Larvae in

“taquaras”” (bamboo).

135. Wyeomyia (D.) brucei Del Ponte & Cerqueira, 1938 [= personata]. Type:

Holotype female (2211), Cuiaba (Mato Grosso), Feb-June 1935, °G. Cesar (LU;

in IOC only “allotype” of Lane and Cerqueira (1942:599) present). Bionomics:

[Larvae in bamboo internodes (Lane and Cerqueira, loc. cit.)].

136. Wyeomyia (D.) rooti (Del Ponte, 1939). Type: LECTOTYPE by present

12 ‘Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

designation, female, same specimen as the holotype of delpontei Lane & Cerqueira,

1942, Cuiaba (Mato Grosso), Feb 1935 (IOC; for explanation see Del Ponte in au-

thors section). Bionomics: [Larvae probably in uncut internodes of bamboo].

137. Wyeomyia (D.) delpontei Lane & Cerqueira, 1942 [= rooti]. Type: Holo-

type female, Cuiaba (Mato Grosso), Feb 1935 (IOC). Bionomics: [Larvae probably

in uncut internodes of bamboo].

138. Wyeomyia (D.) shannoni Lane & Cerqueira, 1942. Type: Holotype male,

Mangaratiba (Rio de Janeiro), Apr 1938 (IOC). Bionomics: [Larvae probably in

leaf axils or flower bracts of Musaceae and Araceae, or possibly in bromeliads or

palms].

139. Wyeomyia (D.) subcomplosa (Del Ponte, 1939) [identity uncertain]. Type:

Syntype females, Cuiaba (Mato Grosso), Feb-Mar 1935, G. Cesar; Rio Canaticu,

Curralinho (Para), Jan 1936, H.W. Kumm; Bahia [Salvador (Bahia)], 1930, R.C.

Shannon and N.C. Davis; Sergipe, 1929, R.C. Shannon and N.C. Davis; Para [Belem

(Para)], Apr 1930, R.C. Shannon and N.C. Davis (LU, possibly FH, IOC or USNM;

for explanation see Del Ponte in authors section). Bionomics: [Pupa in water in

“inaja” palm (Maximiliana regia) | .

140. Wyeomyia (D.) tarsata Lane & Cerqueira, 1942. Type: Holotype male, Ana-

polis (Goias), Sept 1936, Veiga and O. Verano (IOC). Bionomics: Larvae in treehole

(“‘jenipapeiro”’, Genipa americana) in association with Aedes fulvithorax.

141. Wyeomyia (D.) undulata Del Ponte & Cerqueira, 1938. Type: Holotype fe-

male (2213), Cuiaba (Mato Grosso) Feb 1935, G. Cesar (IOC). Bionomics: [Larvae

probably in leaf axils or in bamboo]. |

_ 142, Phoniomyia antunesi (Lane & Guimaraes, 1937). Type: Syntypes males, fe-

males and larvae, Campos do Jordao (Sao Paulo), Dec 1935-Jan 1936, F. Lane

(FH; lectotype not designated). Bionomics: [Larvae in bromeliads] .

143. Phoniomyia bonnei Lane & Cerqueira, 1942. Type: Holotype male, Rio

de Janeiro (Guanabara) (IOC not FH; in latter a male (FH,1362, slide 354) from

Iguacu (Rio de Janeiro), labelled holotype of “‘rockefelleri’’ and bonnei is a para-

type). Bionomics: [Larvae in bromeliads] .

144. Phoniomyia davisi Lane & Cerqueira, 1942. Type: Holotype male, Man-

garatiba (Rio de Janeiro), July 1938, R.C. Shannon (IOC not FH; in latter male

(1515) with genitalia intact, from same locality, May 1938, marked holotype is a

paratype). Bionomics: [Larvae in bromeliads].

145. Phoniomyia diabolica Lane & Forattini, 1952. Type: Holotype male (9110)

with attached mounts of genitalia and larval and pupal skins, Serra do Diabo, rio

Cuiaba, municipio Venceslau (Sao Paulo), 25 Apr 1951, O.P. Forattini (FH). Bio-

nomics: Larvae in epiphytic bromeliads.

146. Phoniomyia edwardsi Lane & Cerqueira, 1942. Type: LECTOTYPE by pre-

sent designation, male (95.x) with genitalia on slide (95 x/y. Mercedes) together

with pupal skin, Porto das Caixas (Rio de Janeiro), 29 May 1925, F.M. Root; 1 of

several specimens identified as quasilongirostris by Dyar (1928:54), Lane and Cer-

queira’s (1942:638, footnote) reference to a single specimen of this series as a holo-

type is incorrect; the present specimen is probably the one from which the figure of

the genitalia in Dyar (1928:fig. 37) was drawn; the larva described and figured by

Dyar (1928: fig. 37) is incorrectly associated and belongs to Weomyia (C.) dyari,

see Lane and Cerqueira, 1942:581 (USNM). Bionomics: Larvae in bromeliads.

147. Phoniomyia flabellata Lane & Cerqueira, 1942. Type: Holotype male, Muri-

queira [Gois Calmon] (Bahia), 1929 (IOC). Bionomics: [Larvae probably in brome-

liads].

Belkin et al: Topotypic Brazilian Mosquitoes 13

148. Phoniomyia galvaoi Correa & Ramalho, 1956. Type: Holotype male (804-4)

with slide (2298, not labelled) of genitalia and larval and pupal skins, Parada 24

de Outubro, municipio Guaruja (Sao Paulo) (FH,10886). Bionomics: [Larvae in

bromeliads]. |

149. Phoniomyia incaudata (Root, 1928). Type: Lectotype male (66) with geni-

talia slide, Rio de Janeiro (Guanabara), 12 Apr 1925, F.M. Root (USNM,44163;

designation by Stone and Knight, 1957b:117). Bionomics: Larvae in bromeliads.

*150. Phoniomyia longirostris (Theobald, 1901). Type: Lectotype female, Rio de

Janeiro (Guanabara), 4 July 1899, A. Lutz (BM; designation by Belkin, 1968:24).

Bionomics: [Larvae in bromeliads].

151. Phoniomyia lopesi Correa & Ramalho, 1956. Type: Holotype male (6.14)

with slide (2300, not labelled) of genitalia and larval and pupal skins, Parada 24

de Outubro, municipio Guaruja (Sao Paulo), 14 Jan 1953 (FH,10888). Bionomics:

Larvae in bromeliads.

152. Phoniomyia pallidoventer Theobald, 1907. Type: Holotype male (10) with

genitalia on slide, Rio de Janeiro (Guanabara), F. Fajardo (BM; see Belkin, 1968:

24). Bionomics: [Larvae in bromeliads].

153. Phoniomyia palmata Lane & Cerqueira, 1942. Type: Holotype male, Rio

de Janeiro (Guanabara) (IOC). Bionomics: Larvae in bromeliads.

154. Phoniomyia pilicauda (Root, 1928). Type: Lectotype male (95.2) with geni-

talia slide and associated larval and pupal skins, Porto das Caixas (Rio de Janeiro),

29 May 1925, F.M. Root (USNM,44164; designation by Stone and Knight, 1957b:

117). Bionomics: Larvae in bromeliads.

155. Phoniomyia quasilongirostris Theobald, 1907. Type: Lectotype female (7),

Mana [Maua], Rio de Janeiro (Guanabara), F. Fajardo (BM; designation by Belkin,

1968:24). Bionomics: [Larvae in bromeliads].

156. Phoniomyia neivai Lane & Cerqueira, 1942 [= quasilongirostris]. Type:

Holotype female, Londrina (Parana) (IOC; in FH (1373) female (11566) from same

locality, Nov 1936, also marked holotype). Bionomics: [Larvae in bromeliads].

157. Phoniomyia theobaldi Lane & Cerqueira, 1942. Type: Holotype male, Rio

de Janeiro (Guanabara), May 1937 (IOC; in FH (1434) male (SFA1) with genitalia

slide, same locality, June 1937, also labelled holotype). Bionomics: [Larvae in bro-

meliads].

*158. Phoniomyia tripartita (Bonne-Wepster & Bonne, 1921). Type: Holotype

male, represented by fig. 2 in Dyar (1919), Sao Paulo (Sao Paulo) (NE, see Belkin,

1968:24). Bionomics: [Larvae probably in bromeliads].

*159. Limatus durhamii Theobald, 1901. Type: Lectotype female, Para [Belem]

(Para), H.E. Durham (BM; designation by Belkin, 1968:23). Bionomics: [Larvae

in treeholes, bamboo, cacao pods, fruit rinds, fallen leaves and artificial containers] .

*160. Limatus curvirostris (Laveran, 1902) [= durhamii]. Type: Male(s) and fe-

male(s), neighborhood of Rio de Janeiro (Guanabara), elev. 300-500 m (NE; see

Belkin, 1968:22). Bionomics: [As for 159. durhamii].

161. Limatus paraensis (Theobald, 1903) [= durhamii]. Type: Holotype female,

Para [Belem] (Para), H.E. Durham (BM). Bionomics: [As for 159. durhamii].

162. Limatus flavisetosus Castro, 1935. Type: Holotype, presumably male, Serra

do Cubatao (Sao Paulo) (LU; not in IOC). Bionomics: Larvae in treeholes.

163. Sabethes (S.) albiprivus Theobald, 1903. Type: Lectotype female, Sao Paulo

(Sao Paulo), 28 Nov, A. Lutz (BM; designation by Belkin, 1968:30). Bionomics:

[Larvae probably in treeholes, bamboo or possibly leaf axils].

164. Sabethes (S.) albiprivatus Lutz, 1904 [= albiprivus|. Type: LECTOTYPE

14 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

by present designation, female, Sao Paulo (Sao Paulo), 28 Nov, A. Lutz, same as

lectotype of 163. albiprivus (BM; see Belkin, 1968:30). Bionomics: [As for 163.

albiprivus |.

165. Sabethes (S.) amazonicus Gordon & Evans, 1922. Type: Holotype female,

ca 300 yds in forest, Macapa, near Manaus (Amazonas), 22 Dec 1921, R.M. Gordon

(BM). Bionomics: [Larvae probably in treeholes, bamboo or possibly leaf axils].

166. Sabethes (S.) longfieldae Edwards, 1928 [= amazonicus]. Type: Holotype

female, Melguierra [?], Ribeirao Amolar, headwaters of Rio Paraguai, south of

Diamantino (Mato Grosso), elev. ca 2000 ft, 24 May 1927, C. Longfield (BM).

Bionomics: [As for 165. amazonicus|.

167. Sabethes (S.) batesi Lane & Cerqueira, 1942. Type: Holotype male, Tingua

(Rio de Janeiro), July 1940, J. Lane (IOC). Bionomics: [Larvae probably in tree-

holes, bamboo or possibly leaf axils].

168. Sabethes (S.) belisarioi Neiva, 1908. Type: Syntypes 2 females, Bicudos

(Minas Gerais), Feb, B. Penna (LU; not in IOC). Bionomics: [Larvae in bromeliads

and occasionally in water on cut trees].

169. Sabethes (S.) argyronotum Edwards, 1928 [= belisarioi|. Type: Lectotype

female, Melguierra [?], Ribeirao Amolar, headwaters of Rio Paraguai, south of

Diamantino (Mato Grosso), elev. ca 2000 ft, 26 May 1927, C. Longfield (BM;

designation by Belkin, 1968:30). Bionomics: [As for 168. belisarioi].

170. Sabethes (S.) nitidus Theobald, 1901 [distinct from bipartipes|. Type:

Lectotype male, Para [Belem] (Para), H.E. Durham (BM; designation by Belkin,

1968:31). Bionomics: [Larvae probably in treeholes, bamboo or leaf axils].

*171. Sabethes (S.) locuples Robineau-Desvoidy, 1827 [= cyaneus]. Type: Fe-

male(s), Brazil, locality not specified; TYPE LOCALITY here restricted to vicinity

of Rio de Janeiro (Guanabara) (NE). Bionomics: [Larvae probably in bamboo or

treeholes].

172. Sabethes (S.) remipes (Wiedemann, 1828) [= cyaneus|. Type: Holotype

female, Brazil, without specified locality; TYPE LOCALITY here restricted to Sal-

vador (Bahia) (ZMC; see Belkin, 1968:32). Bionomics: [As for /7/. locuples|. _ |

173. Sabethes (S.) forattinii Cerqueira, 1961. Type: Holotype male (2018) with

genitalia slide (399), Manaus, km 23 on route 17 to Caracarai (Amazonas), 16 Oct

1959, C. Elias (FH,15017, slide 2963). Bionomics: [Larvae probably in bamboo,

treeholes or possibly leaf axils].

174. Sabethes (S.) lanei Cerqueira, 1961. Type: Holotype male (2015) with geni-

talia slide (465), Manaus, km 23 on route 17 to Caracarai (Amazonas), 14 Oct 1959,

C. Elias (FH,15113, slide 2969). Bionomics: [Larvae probably in bamboo, treeholes

or possibly leaf axils].

175. Sabethes (S.) lutzii Theobald, 1903 [nomen dubium]. Type: Holotype possi-

bly represented by slide (1278) of wing, marked ‘‘da coll. de Dr. Lutz/XI-930/

C.L.”; type locality, Manaus (Amazonas) (?IOC). Bionomics: [Larvae probably in

treeholes, bamboo or possibly leaf axils].

176. Sabethes (S.) purpureus (Theobald, 1907). Type: Lectotype female with

right wing on slide, Rio de Janeiro (Guanabara), E.A. Goeldi (BM; designation by

Belkin, 1968:31). Bionomics: [Larvae in treeholes] .

177. Sabethes (S.) purpureus Peryassu, 1908; remipusculus Dyar, 1924, new name

[= purpureus Theobald, 1907]. Type: LECTOTYPE by present designation, only

remaining specimen, broken (135), head and thorax on 1 pin, leg on point on

another pin, wing on slide (1092), Juiz de Fora (Minas Gerais), 28 Oct 1907, A.

Neiva (LOC,505). Bionomics: [As for 1 76. purpureus Theobald, 1907].

Belkin et al: Topotypic Brazilian Mosquitoes 15

178. Sabethes (S.) quasicyaneus Peryassu, 1922. Type: Female(s), Mato do

Utinga, suburb of Belem (Para) (LU). Bionomics: [Larvae probably in treeholes,

bamboo or possibly leaf axils].

179. Sabethes (S.) shannoni Cerqueira, 1961. Type: Holotype male (2062-43)

with slide (2965) of associated larval and pupal skins (456), Igarape do Leao, Man-

aus (Amazonas), 8 Jan 1960, C. Elias and E. Viera (FH,15109). Bionomics: Larvae

in epiphytic bromeliads, “‘bananeira brava” (Heliconia sylvestris) and tree trunk cut

at about 1 m from ground.

180. Sabethes (S.) spixi Cerqueira, 1961. Type: Holotype male (2013) with geni-

talia slide (393,2962), Manaus, km 23 on route 17 to Caracarai (Amazonas), Oct

1959, C. Elias (FH,15106). Bionomics: [Larvae probably in bamboo, treeholes or

possibly leaf axils]. |

181. Sabethes (Sabethoides) confusus (Theobald, 1903) [= chloropterus|. Type:

Lectotype female (female “‘type” of nitidus), Para [Belem] (Para), H.E. Durham

(BM; designation by Mattingly, 1958:105; see Belkin 1968:30). Bionomics: [Larvae

in treeholes].

182. Sabethes (Sabethoides) glaucodaemon (Dyar & Shannon, 1925). Type:

Holotype female, near San Alberto [?], Rio Branco (Roraima), 28 Aug 1924, J.

Bequaert (USNM,27744). Bionomics: [Larvae in treeholes or bamboo; possibly in

leaf axils of Montrichardia as stated by Lane and Cerqueira, 1942:674].

183. Sabethes (Sabethoides) tridentatus Cerqueira, 1961. Type: Holotype male

(1403.3) with associated larval and pupal skins (62) on slide (2968); slide (422) of

male genitalia missing, Igarape da Bolivia, Manaus (Amazonas), May 1956, C. Elias

et al (FH,15112). Bionomics: Larvae in treeholes in forest. |

184. Sabethes (Sabethinus) aurescens (Lutz, 1905). Type: LECTOTYPE by pre-

sent designation, the specimen identified as the holotype of aurescens Theobald,

1907 by Belkin (1968:30); the latter name is here eliminated (see discussion under

Theobald in section on authors below); Cantareira (Sao Paulo), 16 Apr 1905, A.

Lutz (BM). Bionomics: [Larvae in bamboo internodes].

185. Sabethes (Sabethinus) fabricii Lane & Cerqueira, 1942. Type: Holotype

male, Tingua (Rio de Janeiro), Jan 1941 (IOC). Bionomics: [Larvae probably in

bamboo internodes or treeholes, possibly in leaf axils].

186. Sabethes (Sabethinus) lutzianus Lane & Cerqueira, 1942 [?= identicus].

Type: Holotype male, locality not specified (Rio de Janeiro); TYPE LOCALITY

here restricted to Tingua (IOC). Bionomics: [Larvae probably in bamboo inter-

nodes].

187. Sabethes (Sabethinus) intermedius (Lutz, 1904). Type: LECTOTYPE by

present designation, female marked as lectotype by John Lane, [near Sao Paulo]

(Sao Paulo) (BM; see Belkin, 1968:31). Bionomics: [Larvae in bamboo internodes].

188. Sabethes (Sabethinus) albiprivatus (Theobald, 1907); melanonymphe Dyar,

1924, new name. Type: Lectotype male with thorax and 5 legs on pin, abdomen

and genitalia on | slide, head and left wing on another, Cantareira (Sao Paulo),

16 Apr 1905, A. Lutz (BM; designation by Belkin, 1968:29). Bionomics: [Larvae

in bamboo internodes].

189. Sabethes (Sabethinus) soperi Lane & Cerqueira, 1942. Type: Holotype male,

Piraja (Bahia), Mar 1930, R.C. Shannon (IOC). Bionomics: [Larvae probably in

16 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

bamboo internodes or treeholes; possibly in leaf axils].

190. Sabethes (Sabethinus) whitmani Lane & Cerqueira, 1942. Type: Holotype

male, Sao Joao de Petropolis, Santa Teresa (Espirito Santo), May 1940 (IOC). Bio-

nomics: [Larvae probably in bamboo internodes or treeholes; possibly in leaf axils].

191. Coquillettidia (Rhynchotaenia) albicosta (Peryassu, 1908). Type: LECTO-

TYPE by present designation, only remaining specimen, female, Xerem (Rio de

Janeiro), 28 Oct 1907 (IOC). Bionomics: [Larvae probably attached to roots of

grassy vegetation in permanent or semipermanent ground waters].

192. Coquillettidia (R.) albifera (Prado, 1931). Type: Holotype rales marshes

near Rio Pinheiros, Butantan, Sao Paulo (Sao Paulo), 23 Apr 1931, Franca (IB, —

1040; antigo 72). Bionomics: [Larvae probably attached to roots of grassy vege-

tation in permanent or semipermanent ground waters].

193. Coquillettidia (R.) arribalzagae (Theobald, 1903). Type: Lectotype female,

Para [Belem] (Para), H.E. Durham (BM; designation by Belkin, 1968: 11). Bio-

nomics: [Larvae probably attached to roots of grassy vegetation in permanent or

semipermanent ground waters].

194. Coquillettidia (R.) chrysonotum (Peryassu, 1922). Type: Syntypes males

and females, Vale do Rio Doce (Espirito Santo) and Baixada Fluminense (Rio de

Janeiro), Mar-Apr (Museu Nac Rio de Janeiro and IOC, tubes 431 and 432). Bio-

nomics: [Larvae attached to roots of Brass vegetation in permanent or semiper-

manent ground waters].

195. Coquillettidia (R.) hermanoi (ise & Coutinho, 1940). Type: Holotype

female (1010), Acampamento dos Morros Azues (Mato Grosso), 6 Sept 1937, A.

Bueno de Oliveira (FH,1630; tube 1461). Bionomics: [Larvae probably attached

to roots of grassy vegetation in permanent or semipermanent ground waters].

196. Coquillettidia (R.) juxtamansonia (Chagas, 1907). Type: LECTOTYPE by

present designation, female (tube 428), wing on slide (1001), Juiz de Fora (Minas

Gerais), Oct 1906, A. Neiva (IOC). Bionomics: [Larvae attached to roots of grassy

vegetation in permanent or semipermanent ground waters].

197. Coquillettidia (R.) hypocindyna (Dyar, 1918) [= juxtamansonia|. Type:

Holotype female, [Sao Paulo] (Sao Paulo), A. Lutz (USNM,21720). Bionomics:

[Larvae probably attached to roots of grassy vegetation in permanent or semi- —

permanent ground waters].

198. Coquillettidia (R.) lynchi (Shannon, 1931). Type: Holotype male, Para

[Belem] (Para), N.C. Davis (USNM). Bionomics: [Larvae probably attached to roots

of grassy vegetation in permanent or semipermanent ground waters].

199. Coquillettidia (R.) neivai (Lane & Coutinho, 1940) [distinct from nigricans,

see Belkin, Heinemann and Page, 1970:103]. Type: Holotype male (1137) with

genitalia on slide (453), marked “‘Mansonia chrysa” and Taeniorhynchus nigricans

(pinned specimen only), Juquia (Sao Paulo), 16 Jan 1939, J. Lane (FH,1676).

Bionomics: [Larvae probably attached to roots of grassy vegetation in permanent

and semipermanent ground waters].

200. Coquillettidia (R.) shannoni (Lane & Antunes, 1937). Type: Holotype fe-

male, Cuiaba (Mato Grosso), Sept 1934, J. Lane (FH,809). Bionomics: [Larvae pro-

bably attached to roots of grassy vegetation in permanent or semipermanent ground

waters].

201. Mansonia (M.) amazonensis (Theobald, 1901). Type: Holotype female, S.S.

Faraday, between Gurupa and Monte Alegre (Para), 25 Jan 1896, E.E. Austen:

TYPE LOCALITY here restricted to vicinity of Gurupa (BM). Bionomcs: | Larvae

attached to vegetation in permanent or semipermanent ground waters].

Belkin et al: Topotypic Brazilian Mosquitoes 17

202. Mansonia (M.) cerqueirai (Barreto & Coutinho, 1944). Type: Holotype male,

Maracaju (Mato Grosso) (LU). Bionomics: [Larvae probably attached to vegetation

in permanent or semipermanent ground waters].

203. Mansonia (M.) chagasi (Lima, 1935). Type: Holotype male, Bicudos (Minas

Gerais), 11 Feb 1908, C. Chagas (IOC, 1956). Bionomics: [Larvae probably attached

to vegetation in permanent or semipermanent ground waters].

204. Mansonia (M.) indubitans Dyar & Shannon, 1925. Type: Holotype female,

Belem (Para) 19 Sept 1924, J. Bequaert (USNM,27746). Bionomics: [Larvae at-

tached to vegetation (Pistia) in permanent or semipermanent ground waters].

205. Mansonia (M.) pessoai (Barreto & Coutinho, 1944). Type: Holotype male

with genitalia on slide (512), marked paratype, Curitiba (Parana), G. Ramalho

(FH,1822). Bionomics: [Larvae probably attached to vegetation in permanent or

semipermanent ground waters].

206. Mansonia (M.) pseudotitillans (Theobald, 1901). Type: Lectotype female,

S.S. F[araday], Breves (Para), 25 Jan 1896, E.E. Austen (BM; designation by Belkin,

1968:23). Bionomics: [Larvae probably attached to vegetation in permanent or

semipermanent ground waters].

*207. Mansonia (M.) titillans (Walker, 1848). Type: Holotype female, Brazil, with-

out specified locality; TYPE LOCALITY here restricted to vicinity of Belem (Para)

(BM; see Belkin, 1968:23). Bionomics: [Larvae attached to Pistia and floating

grass]. ,

208. Mansonia (M.) wilsoni (Barretto & Coutinho, 1944). Type: Holotype male,

Sao Paulo (Sao Paulo) (LU, not in FMSP). Bionomics: [Larvae probably attached

to vegetation in permanent and semipermanent ground waters].

209. Uranotaenia albitarsis Gordon & Evans, 1922 [= calosomata]|. Type: Lecto-

type male (15/463) with genitalia slide, sawmill near Macapa (near Manaus) (Ama-

zonas), 20 Jan 1922, R.M. Gordon (BM; designation by Belkin, 1968:37). Bio-

nomics: Larvae in old iron “‘bath’’.

210. Uranotaenia davisi Lane, 1943. Type: Holotype male (436) with genitalia

on slide (897), Salvador (Bahia), P.C.A. Antunes (FH,3557). Bionomics: [Larvae

probably in permanent or semipermanent ground waters].

211. Uranotaenia ditaenionota Prado, 1931. Type: Holotype female, woods of

Butantan, Sao Paulo (Sao Paulo), 15 July 1931, D. Yered (IB, 1131; antigo 175).

Bionomics: [Larvae in pools of dirty water and in a small stream (Lane, 1943:160)].

212. Uranotaenia burkii Lane, 1936 [= ditaenionota]. Type: Holotype male

(232A) with genitalia slide (32) and larval skin slide (31), [Coronel] Ponce (Mato

Grosso), 24-26 July 1934 (FH,367). Bionomics: Larvae in a shaded pool with dirty

water and in a small stream pool.

213. Uranotaenia geometrica Theobald, 1901. Type: Lectotype female, Cubatao

(Sao Paulo), A. Lutz (BM; designation by Belkin, 1968:37). Bionomics: [Larvae

in ground and rock pools with algae].

214, Uranotaenia mathesoni Lane, 1943. Type: Holotype male (1112) with geni-

talia on slide (895), Juquia (Sao Paulo), Nov 1938, J. Lane (FH,3554). Bionomics:

[Larvae probably in permanent or semipermanent ground waters].

215. Uranotaenia noctivaga Neiva & Pinto, 1922 [= nataliae]. Type: Holotype

female, Gavea, Rio de Janeiro (Guanabara) (LU; not found in IOC). Bionomics:

[Larvae probably in permanent or semipermanent ground waters].

216. Uranotaenia argenteopennis Peryassu, 1923 [= nataliae|. Type: Syntype

male(s) and female(s), [vicinity of Rio de Janeiro] (Guanabara) and Baixada Flumi-

nense (Rio de Janeiro) (LU). Bionomics: Larvae in association with anophelines

18 : 3 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

[in permanent and semipermanent ground waters].

217. Uranotaenia pallidoventer Theobald, 1903. Type: Holotype female, Para

[Belem] (Para), H.E. Durham (BM). Bionomics: [Larvae in ground pools].

218. Orthopodomyia albicosta (Lutz, 1904). Type: LECTOTYPE by present de-

signation, female (2663), marked S. Paulo; type locality, Serra da Cantareira (Sao

Paulo), Lutz; 1905:69 (IOC; in box 14). Bionomics: Larvae in bamboo.

*219. Orthopodomyia longipalpis (Newstead & Thomas, 1910) [= fascipes]. Type:

Lectotype female, near Manaus (Amazonas), 23 Aug 1906 (BM; designation by Za-

vortink, 1968:75). Bionomics: [Larvae in treeholes].

220. Orthopodomyia townsendi Lima, 1935 [= fascipes|. Type: Holotype fe-

male, Rio Tapajos, Boa Vista (Para), 11 Oct 1932, C.H.T. Townsend (IOC,582).

Bionomics: [Larvae in treeholes].

221. Orthopodomyia sampaioi Lima, 1935. Type: Lectotype male (1861) with

genitalia in capillary tube, Tijuca, Rio de Janeiro (Guanabara), June 1934, P.C.

Sampaio (IOC, tube 941; designation by Zavortink, 1968:82). Bionomics: [Larvae

in treeholes].

222. Psorophora(P.) tibialis (Robineau-Desvoidy, 1827) [= ciliata]. Type: Male(s)

Brazil, locality not specified; TYPE LOCALITY here restricted to vicinity of Rio de

Janeiro (Guanabara) (NE). Bionomics: [Larvae in rain pools].

223. Psorophora (P.) pilipes (Macquart, 1834) [?= ciliata]. Type: Female(s),

locality not specified, Brazil; TYPE LOCALITY here restricted to vicinity of Rio

de Janeiro (Guanabara) (NE; see Belkin, 1968:28). Bionomics: [Probably as for

222. tibialis].

224. Psorophora (P.) scintillans (Walker, 1848) [= cilipes]. Type: Lectotype male

(45.56) with genitalia intact, Para [Belem] (Para) (BM; designation by Belkin, 1968:

29). Bionomics: [Larvae in rain pools].

225. Psorophora (P.) genumaculata Cruz, 1907 [= lineata]. Type: LECTOTYPE

by present designation, male (2582) without head but genitalia intact, Santos (Sao

Paulo), Oct 1906, T. Ribeiro Gomes (IOC; box 9). Bionomics: [Larvae in rain

pools].

226. Psorophora (Janthinosoma) albigenu (Peryassu, 1908) [distinct from vari-

pes|. Type: Holotype female (3523), Chanaan [Canaan] (Sao Paulo), M. Latif

(IOC, tube 1367). Bionomics: [Larvae probably in rain pools in wooded areas].

227. Psorophora (J.) amazonica Cerqueira, 1960. Type: Holotype male (1443.60)

with slide (2960) of larval and pupal skins (342), slide of genitalia (386) missing,

Igarape do Taruma, Manaus (Amazonas), 16 June 1956, C. Elias (FH,15104). Bio-

nomics: Larvae in a grassy pool at edge of forest.

228. Psorophora (J.) discrucians (Walker, 1856). Type: Syntypes male(s) and

female(s), described from South America; TYPE LOCALITY here restricted to vi-

cinity of Rio de Janeiro (Guanabara) (BM; see Belkin, 1968:26). Bionomics: [Larvae

probably in rain pools in wooded areas].

229. Psorophora (J.) arribalzagae (Giles, 1902) [?= discrucians]. Type: Holotype

female, Sao Paulo (Sao Paulo), A. Lutz (BM; see Belkin, 1968:25). Bionomics:

[As for 228. discrucians].

230. Psorophora (J.) forceps Cerqueira, 1939. Type: Holotype male, Jacarepagua,

Rio de Janeiro (Guanabara) (IOC). Bionomics: [Larvae probably in rain pools in

wooded areas].

231. Psorophora (J.) lanei Shannon & Cerqueira, 1943. Type: Holotype female,

Maracaju (Mato Grosso) (LU; not in IOC). Bionomics: [Larvae probably in rain

pools in wooded areas].

Belkin et al: Topotypic Brazilian Mosquitoes 19

232. Psorophora (J.) lutzii (Theobald, 1901). Type: Lectotype female, Parque

do Museu [Quinta da Boa Vista], Rio de Janeiro (Guanabara), 5 Nov 1899°¢€.

Moreira (BM; designation by Belkin, 1968:27). Bionomics: [Larvae in rain pools

in wooded areas]. |

233. Psorophora (Grabhamia) apicalis (Theobald, 1903); neoapicalis (Theobald,

1910), new name [?= cingulata]. Type: Lectotype male, Rio de Janeiro (Guana-

bara), A. Lutz (BM; designation by Belkin, 1968:25). Bionomics: [Larvae in wood-

land pools, hoofprints and rarely in artificial containers].

234. Psorophora (G.) scutipunctata (Lutz & Neiva, 1911) [= confinnis|. Type:

Syntypes [?], 1 or 2 females (tube 1060) and slide of wing (2347), without data;

type locality [Rio] Tiete, immediately above Itapura (Sao Paulo), Jan 1909 (IOC;

only 2 syntypes mentioned in original description but 3 in tube, 2 mounted on

1 pin). Bionomics: [Larvae probably in rain pools].

235. Psorophora perterrens (Walker, 1856) [nomen dubium]. Type: Holotype

female, described from South America; TYPE LOCALITY here restricted to vici-

nity of Rio de Janeiro (Guanabara) (NE; see Stone, 1957:337-338).

236. Aedes (Ochlerotatus) aenigmaticus Cerqueira & Costa, 1946. Type: Holo-

type male, Maracaju (Mato Grosso), Jan-Feb 1938, R.C. Shannon (LU; not in IOC).

Bionomics: [Larvae probably in rain pools].

237. Aedes (O.) albifasciatus (Macquart, 1838). Type: Holotype female, Brazil,

locality not specified, 1833, C. Gaudichaud; TYPE LOCALITY here restricted to

vicinity of Rio de Janeiro (Guanabara) (MNHP; see Belkin, 1968:4). Bionomics:

[Larvae in temporary ground pools]. ,

238. Aedes (O.) crinifer (Theobald, 1903). Type: Lectotype female, Sao Paulo

(Sao Paulo), A. Lutz (BM; designation by Belkin, 1968:4). Bionomics: [Larvae

in grassy rain pools and flooded grassy margins of creeks].

239. Aedes (O.) fulvus (Wiedemann, 1828). Type: Holotype female, Brazil, lo-

cality not specified, Freireiss; TYPE LOCALITY here restricted to Salvador (Bahia)

(SNG). Bionomics: [Larvae in forest rain pools].

240. Aedes (O.) flavicosta (Walker, 1856) [= fulvus|. Type: Holotype female,

Amazon region; TYPE LOCALITY here restricted to Manaus (Amazonas) (BM).

Bionomics: [As for 239. fulvus].

241. Aedes (O.) jacobinae Serafim & Davis, 1933. Type: Holotype male apparent-

ly lost, only genitalia slide remaining, Rio d’Ouro, in center of Jacobina (Bahia),

elev. 450 m, Dec 1931, J. Serafim (USNM; see Stone and Knight, 1956a:220).

Bionomics: Larvae in streamside rock pools.

242, Aedes (O.) lepidus Cerqueira & Paraense, 1945. Type: Holotype male, Belo

Horizonte (Minas Gerais), Nov 1943 (LU; not in IOC). Bionomics: [Larvae in arti-

ficial containers and rockholes, Cerqueira, 1957].

243. Aedes (O.) pennai Antunes & Lane, 1938. Type: Holotype male (970) with

genitalia on 3 slides (185,186,187), Cabreuva (Sao Paulo), Apr 1937, A.R.R. (FH,

810). Bionomics: Larvae in forest rain pools. —

244. Aedes (O.) perventor Cerqueira & Costa, 1946, Type: Holotype male, Man-

garatiba (Rio de Janeiro), Dec 1938, R.C. Shannon (LU; not in IOC). Bionomics:

[Larvae in crabholes, material in FH].

245. Aedes (O.) rhyacophilus Lima, 1933. Type: Syntypes males, females with

larval exuviae and pupae, Vale do Canaa (Espirito Santo), 1932, J. Serafim; TYPE

LOCALITY here restricted to Sao Joao de Petropolis (LU; not in IOC). Bionomics:

Larvae in rockholes.

246. Aedes (O.) scapularis (Rondani, 1848). Type: Female(s), Brazil, locality

20 3 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

not specified; TYPE LOCALITY here restricted to vicinity of Rio de Janeiro (Gua-

nabara) (LU; see Belkin, 1968:7). Bionomics: [Larvae in grassy rain pools].

247. Aedes (O.) serratus (Theobald, 1901). Type: Lectotype male with attached

genitalia mount, Parque do Museu [Quinta da Boa Vista], Rio de Janeiro (Guana-

bara), 5 Nov 1899, C. Moreira (BM; designation by Belkin, 1968:7). Bionomics:

[Larvae in grassy rain pools].

248. Aedes (O.) mathisi (Neveu-Lemaire, 1902) [?= serratus]. Type: Syntypes

2 females, Cunani (Amapa) [as French Guiana], Jan 1901, Mathis (NE; see Belkin,

1968:6). Bionomics: [As for 247. serratus].

249. Aedes (F.) braziliensis Gordon & Evans, 1922. Tape: Holotype male (10.1/

463), Macapa, near Manaus (Amazonas), 8 Dec 1921, R.M. Gordon (BM). Biono-

mics: Larvae in hollow tree stump..

250. Aedes (Finlaya) fluviatilis (Lutz, 1904). Type: LECTOTYPE by present

designation, the female holotype of tripunctatus (Theobald, 1907), Rio Grande

near Franca (Sao Paulo), 23 Sept 1903, A. Lutz (BM; see Belkin, 1968:5). Biono-

mics: [Larvae in stream bed rockholes].

251. Aedes (F.) mediomaculatus (Theobald, 1907) [?= fluviatilis]. Type: Lecto-

type male, Para [Belem] (Para), E.A. Goeldi (BM; designation by Belkin, 1968:6).

Bionomics: [Probably as for 250. fluviatilis].

252. Aedes (F.) tripunctatus (Theobald, 1907) [?= fluviatilis]. Type: Holotype

female, Rio Grande near Franca (Sao Paulo), 23 Sept 1903, A. Lutz (BM; see Belkin,

1968: 8). Bionomics: [Larvae probably as for 250. Suvietilied:

*253. Aedes (F.) leucomelas (Lutz, 1904); leucocelaenus Dyar & Shannon, 1924,

new name. Type: LECTOTYPE by present designation, female marked by Theobald

“Stegomyia silvestris Lutz Type’, Franca (Sao Paulo), 23 Sept 1903 (BM; see Bel-

kin, 1968:5-6). Bionomics: [Larvae in treeholes and bamboo internodes].

254. Aedes (F.) leucophoebus Galindo, Carpenter & Trapido, 1953. Type: Holo-

type male (1763.2) with slides of genitalia (2204) and associated larval and pupal

skins (2205), Feijo (Acre), 4 Aug 1949 (FH,10377). Bionomics: [Larvae in tree-

holes].

| 255, Aedes (F.) terrens (Walker, 1856). Type: Holotype male with attached geni-

talia mount; described from South America, TYPE LOCALITY here restricted to

vicinity of Rio de Janeiro (Guanabara) (BM). Bionomics: [Larvae in treeholes].

*256. Aedes (F.) oswaldi (Lutz, 1904) [= terrens]. Type: LECTOTYPE by pre-

sent designation, female marked by Theobald “Stegomyia cruzi Lutz Type’’; lo-

cality label here reinterpreted as “[F. do] Bonito, S[erra] d[a] Bocaina [Sao

Paulo]”’, type locality of a tabanid described by A. Lutz (see Fairchild, 1961:207);

22 May 1903 (BM; see Belkin, 1968:6). Bionomics: [Larvae in treeholes] .

257. Aedes (Howardina) aureolineatus Berlin, 1969. Type: Holotype female, Pi-

aja (Bahia), May 1931 (USNM). Bionomics: [Larvae probably in treeholes].

258. Aedes (H.) fulvithorax (Lutz, 1904). Type: Holotype female, Ponte Ipe-

Arcado (Goias) (NE). Bionomics: [Larvae in treeholes and bamboo].

259, Aedes (Stegomyia) toxorhynchus (Macquart, 1838) [= aegypti]. Type: Ho-

lotype female (10), Brazil, locality not specified, C. Gaudichaud, 1833; TYPE LO-

CALITY here restricted to vicinity of Rio de Janeiro (Guanabara) (MNHP;2602/

33; see Belkin, 1968:8). Bionomics: [Larvae in artificial containers].

260. Aedes (S.) exagitans (Walker, 1856) [= aegypti]. Type: Holotype female

(37), Para [Belem] (Para) (BM). Bionomics: [Larvae in artificial containers] .

261. Haemagogus (Longipalpifer) tropicalis Cerqueira & Antunes, 1938. Type:

Holotype male, Curralinho (Para), Jan-May 1936, H.W. Kumm and A. Rabello

Belkin et al: Topotypic Brazilian Mosquitoes 21

(LU; not in IOC). Bionomics: Larvae in treeholes.

262. Haemagogus (Stegoconops) baresi Cerqueira, 1960. Type: Holotype male

(1607.3) with pupal slide (316;2961) but genitalia slide (332) missing, Igarape do

Taruma, Manaus (Amazonas), 6 Dec 1956, C. Elias (FH,15105). Bionomics: Larvae

in treeholes in dark forest.

*263. Haemagogus (S.) capricornii Lutz, 1904. Type: Neotype female reared from

egg, Horto Florestal, Serra da Cantareira (Sao Paulo), Apr 1944 (LU, not in IOC or

FH; designation by Cerqueira and Lane, 1945:286). Bionomics: [Larvae in tree-

holes].

264. Culex (Lutzia) brasiliae (Dyar, 1923) [= bigoti]. Type: Lectotype male with

genitalia slide (1778), Sao Paulo (Sao Paulo), A. Lutz (USNM; designation by Stone

and Knight, 1957a:44). Bionomics: [Larvae in permanent or semipermanent ground

waters]. |

265. Culex (C.) abnormalis Lane, 1936. Type: Holotype male (289) with geni-

talia on slide (289;184a), Coronel Ponce (Mato Grosso), 17 July-27 Sept 1934

(FH,361). Bionomics: Larvae in turbid water next to a creek and in hoofprints

with clean water, in full sun.

266. Culex (C.) acharistus Root, 1927. Type: Lectotype fragmentary male (64-1)

with genitalia on slide, Agua Limpa, near Juiz de Fora (Minas Gerais), 27 Mar 1925,

F.M. Root (USNM; designation by Stone and Knight, 1957a:42). Bionomics: Lar-

vae in marshy expansions of mountain streams and in side pools of a small rapid

river.

267. Culex (C.) airozai Lane, 1945. Type: Holotype male with attached genitalia

mount, Rio Parauari (Amazonas), Mar 1937, C. Worontzow (FH,4297). Bionomics:

[Larvae probably in permanent or semipermanent ground waters].

268. Culex (C.) brami Forattini, Rabello & Lopes, 1967. Type: Holotype male

(15844) with genitalia on slide (3109), Boraceia, municipio Salesopolis (Sao Paulo),

Nov 1965, E.X. Rabello (FH). Bionomics: Pupae in peridomestic artificial con-

tainers. |

269. Culex (C.) carcinoxenus Castro, 1932. Type: LECTOTYPE by present desig-

nation, male “‘cotype’’, with genitalia on slide (4186), Bertioga (Sao Paulo), July

1931, G.M. de Oliveira Castro (IOC; no material in IBSP where “typos” (pre-

sumably 1 male and 1 female) were originally deposited without designation of a

holotype). Bionomics: Larvae in crabholes (Cardisoma guanhumi)..

270. ‘Culex (C.) corniger Theobald, 1903. Type: Lectotype male with attached

genitalia mount, Para [Belem] (Para), E.A. Goeldi (BM; designation by Belkin,

1968:15). Bionomics: [Larvae in ground pools, stream bed rock pools, treeholes,

bamboo, artificial containers and flower bracts].

271. Culex (C.) deanei Correa & Ramalho, 1959. Type: Holotype male (S.2-6)

with genitalia and larval and pupal skins (SPM-62) on | slide (2576), Campo de

Marte, Santana, Sao Paulo (Sao Paulo), R.R. Correa and G.R. Ramalho (FH,

13189). Bionomics: Larvae in ground pools.

272. Culex (C.) bilineatus Theobald, 1903 [?= dolosus]. Type: Lectotype male

with attached genitalia mount; TYPE LOCALITY here restricted to Sao Paulo (Sao

Paulo), A. Lutz (BM; designation by Belkin, 1968:13). Bionomics: [Larvae in per-

manent or semipermanent ground waters].

273. Culex (C.) foliaceus Lane, 1945. Type: Holotype male (5643) with geni-

talia on slide (1288), Ares, near Sao Jose de Mipibu (Rio Grande do Norte), Oct

1935 (FH,5926). Bionomics: [Larvae in crabholes, Stone, 1950:239].

274. Culex (C.) forattinii Correa & Ramalho, 1959. Type: Holotype male (197.6)

22 : Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

with genitalia on slide (2575), Santa Cruz do Rio Pardo (Sao Paulo), Feb 1952

(FH,13188). Bionomics: Larvae in a cemetery vase.

275. Culex (C.) lygrus Root, 1927. Type: Lectotype male (115.2) with genitalia

slide, Mage (Rio de Janeiro), 21 June 1925, F.M. Root (USNM; designation by

Stone and Knight, 1957a:53). Bionomics: Larvae in small ditches and pools with

abundant vegetation.

276. Culex (C.) mauesensis Lane, 1945. Type: Holotype male with attached geni-

talia mount, Maues (Amazonas), Feb 1937, C. Worontzow (FH,4885). Bionomics:

[Larvae probably in permanent or semipermanent ground waters].

277. Culex (C.) oswaldoi Forattini, 1965. Type: Holotype male (12) with geni-

talia on slide (RB62-290), Macaphyba, Natal (Rio Grande do Norte), 23 July 1943,

MacCreary (USNM,67550). Bionomics: Larva in quarry hole.

278. Culex (C.) paramaxi Duret, 1968. Type: Holotype male (672), Engenheiro

Dolabela (Minas Gerais), 6 May 1964 (A). Bionomics: [Larvae probably in per-

manent or semipermanent ground waters].

279. Culex (C.) aestuans Wiedemann, 1828 [= quinquefasciatus|. Type: Lecto-

type male with genitalia slide, Brazil, locality not specified; TYPE LOCALITY here

restricted to Salvador (Bahia) (NMW; designation by Stone, 1958:186). Bionomics:

[Larvae in artificial containers and polluted ground waters].

280. Culex (C.) renatoi Lane & Ramalho, 1960. Type: Holotype male (1372-9)

with genitalia and larval and pupal skins on slide (2423), Bairro Sao Miguel Paulista,

Sao Paulo (Sao Paulo), 5 Jan 1956, F. Rosario (FH,12058). Bionomics: Larvae

in terrestrial bromeliads.

281. Culex (C.) spinosus Lutz, 1905. By be! LECTOTYPE by present designation,

male mounted on 2 slides, Brazil, Dr. Lutz; type locality Sao Paulo (Sao Paulo)

(BM, see Belkin, 1968:20; no material of original series in IOC). Bionomics: Larvae.

in leaf axils of Eriocaulon vaginatum and Eryngium alvofolium.

282. Culex (C.) virgultus Theobald, 1901(nomen dubium). Type: Syntypes 2

males, Parque do Museu [Quinta da Boa Vista] , Rio de Janeiro (Guanabara), 5 Nov

1899, C. Moreira (LU; see Belkin, 1968:21). Bionomics: [Larvae in permanent or

- semipermanent ground pools]. |

283. Culex (Melanoconion) gordoni Evans, 1924 [= albinensis|]. Type: Holotype

male (16.1/463) with attached wing mount and 3 genitalia slides, the Bosque, Man-

aus (Amazonas), 29 Dec 1921, R.M. Gordon (BM). Bionomics: Larvae in a ground

pool.

284. Culex (Mel.) andricus Root, 1927. Type: Holotype male, near Lassance

(Minas Gerais), 13 May 1925, F.M. Root (USNM). Bionomics: Larvae in small -

pond full of vegetation.

285. Culex (Mel.) aureonotatus Duret & Barreto, 1956. Type: Holotype male,

Fazenda Monte Alegre, Ribeirao Preto (Sao Paulo), 27 Dec 1954, M.P. Barreto

and J.P. Duret (FMRP). Bionomics: [Larvae probably in permanent or semiper-

manent ground waters].

286. Culex (Mel.) bahiensis Duret, 1969. Type: Holotype male (2427), Urucuca

(Bahia), Aug 1953, J.P. Duret (A). Bionomics: [Larvae probably in permanent or

semipermanent ground waters].

287. Culex (Mel.) innominatus Evans, 1924 [= bastagarius]. Type: Lectotype

male (D3) with attached genitalia mount, River Amazon, to or from Manaus, 1915,

A.A. Clark; TYPE LOCALITY here restricted to Itacoatiara (Amazonas) (BM; desig-

nation by Belkin, 1968:16). Bionomics: [Larvae probably in permanent or semi-

permanent ground waters].

Belkin et al: Topotypic Brazilian Mosquitoes 23

288. Culex (Mel.) bequaerti Dyar & Shannon, 1925. Type: Holotype male, Soro-

roca, Rio Branco (Roraima), 1 Sept 1924, J.C. Bequaert (USNM,27745). Bionomics:

[Larvae probably in permanent or semipermanent ground waters].

289. Culex (Mel.) bifoliolatus Duret & Barreto, 1956. Type: Holotype male, Fa-

zenda Monte Alegre, Ribeirao Preto (Sao Paulo), 2 June 1953, M.P. Barreto and

J.P. Duret (FMRP). Bionomics: [Larvae probably in permanent or semipermanent

ground waters].

290. Culex (Mel.) mojuensis Duret & Damasceno, 1955 [= breviculus|. Type:

Holotype male (Br49,E4), Oriboca, Rio Guajara (Para), 23 Aug 1953, R.G. Damas-

ceno and J.P. Duret (Duret). Bionomics: [Larvae probably in permanent or semi-

permanent ground waters].

291. Culex (Mel.) chrysothorax (Peryassu, 1908). Type: LECTOTYPE bn: pre-

sent designation, male with genitalia slide (2331), Copacabana, Rio de Janeiro

(Guanabara), Oct 1907 (IOC,2033). Bionomics: Larvae probably in ground pools

only and not found in bromeliads also as stated by Peryassu.

292. Culex (Mel.) contei Duret, 1968. Type: Holotype male (2226), Sao Miguel

do Guama (Para), 9 July 1963, J.P. Duret (A). Bionomics: [Larvae probably in

permanent or semipermanent sround waters].

293. Culex (Mel.) cristovai Duret, 1968. Type: Holo male (1883), Caracarai

(Roraima), 7 July 1964 (A). Bioapance: [Larvae probably in permanent or semi-

permanent ground waters].

294. Culex (Mel.) exedrus Root, 1927 [= dunni]. Type: aaeet: male (30.1),

head missing, Porto das Caixas (Rio de Janeiro), 24 Feb 1925, F.M. Root (USNM;

designation by Stone and Knight, 1957a:49). Bionomics: Larvae in dense aquatic

vegetation in rivers, lagoons and ponds.

295. Culex (Mel.) dyius Root, 1927. Type: Holotype male genitalia only, adult

lost, probably coastal lowlands (Rio de Janeiro), May or June 1925, F.M. Root

(USNM). Bionomics: [Larvae probably in permanent or semipermanent ground

waters].

296. Culex (Mel.) manaosensis Evans, 1924 [= eastor]. Type: Holotype male,

wharf, Manaus (Amazonas), 5 Dec 1923, A.A. Clark (BM). Bionomics: [Larvae

probably in swamps].

297. Culex (Mel.) ernanii Duret, 1968. Type: Holotype male (1730), Boa Vista

(Roraima), 4 July 1964, J.P. Duret (A). Bionomics: [Larvae probably in permanent

or semipermanent ground waters].

298. Culex (Mel.) evansae Root, 1927. Type: Lectotype male (34) with geni-

talia slide (34a), Mage (Rio de Janeiro), 26 Feb 1925, N.C. Davis (USNM; desig-

nation by Stone and Knight, 1957a:49). Bionomics: Larvae in small ‘‘jungle”’ pools.

299. Culex (Mel.) fasciolatus (Lutz, 1904). Type: Syntypes 2 females (Lutz,

1905), wooded mountains near Sao Paulo (Sao Paulo) (possibly in Lutz collection in

IOC). Bionomics: Larvae in swamps.

300. Culex (Mel.) faurani Duret, 1968. Type: Holotype male (1940), near Manaus

(Amazonas), 28 June 1963, J.P. Duret (A). Bionomics: [Larvae probably in per-

manent or semipermanent ground waters].

301. Culex (Mel.) flochi Duret, 1969 [probably not Melanoconion]. Type: Holo-

type male (2283), Rio Moju, Belem (Para), 21 Aug 1953, J.P. Duret and R.G. Dama-

sceno (A). Bionomics: [Larvae probably in bromeliads].

302. Culex (Mel.) humilis Theobald, 1901. Type: Lectotype male, genitalia ap-

parently lost, Sao Paulo (Sao Paulo), A. Lutz (BM; designation by Belkin, 1968:16).

Bionomics: [Larvae in stream bed pools].

24 | Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

303. Culex (Mel.) inadmirabilis Dyar, 1928. Type: Holotype male, Sao Paulo

(Sao Paulo), A. Lutz (USNM,40776). Bionomics: [Larvae probably in permanent

or semipermanent ground waters].

304. Culex (Mel.) indecorabilis (Theobald, 1903). Type: Lectotype female, Para

[Belem] (Para), H.E. Durham (BM; designation by Belkin, 1968:16). Bionomics:

Larvae probably in permanent or semipermanent ground waters].

305. Culex (Mel.) cenus Root, 1927 [= intrincatus]. Type: Lectotype male

(115-1) with genitalia slide, Mage (Rio de Janeiro), 21 June 1925 (USNM,40527;

designation by Stone and Knight, 1957a:45). Bionomics: Larvae in woodland pools,

roadside ditch and side eddies of a river.

306. Culex (Mel.) isabelae Duret, 1968. Type: Holotype male (1925), Caracarai

(Roraima), 3 July 1964, J.P. Duret (A). Bionomics: [Larvae probably in permanent

or semipermanent stound waters] . |

307. Culex (Mel.) johnnyi Duret, 1968. Type: Holotype male (723), Rio Preto,

municipio Joao Goulard (Amazonas), 18 July 1964, J.P. Duret (A). Bionomics:

[Larvae probably in permanent or semipermanent sround waters].

308. Culex (Mel.) kerri Duret, 1968. Type: Holotype male (1858), Rio Preto,

municipio Joao Goulard (Aarons): 17 July 1964, J.P. Duret (A). Bionomics:

[Larvae probably in permanent or semipermanent ground waters].

309. Culex (Mel.) lugens Lutz, 1905. Type: Syntypes male(s) and female(s), 1 of

each remaining in collection (2610), Lagoa (Sao Paulo), 3 Feb 1904, A. Lutz (IOC;

in box 6). Bionomics: Larvae in swamps.

310. Culex (Mel.) nigrescens (Theobald, 1907). Type: Holotype male, Santo

Amaro, Sao Paulo (Sao Paulo), 1 Nov 1900, A. Lutz (BM). Bionomics: [Larvae

probably in permanent or semipermanent ground waters].

311. Culex (Mel.) clarki Evans, 1924 [= nigrescens]. Type: Lectotype male (C4)

with attached wing mount and genitalia slide, River Amazon (Amazonas), 1915,

A.A. Clark; TYPE LOCALITY here restricted to Manaus (Amazonas) (BM; desig-

nation by Belkin, 1968:14). Bionomics: [As for 310. nigrescens].

312. Culex (Mel.) nigricorpus (Theobald, 1901). Type: Lectotype female, Itacoa-

tiara (Amazonas), 7 Feb 1896, E.E. Austen (BM; designation by Belkin, 1968:18).

Bionomics: [Larvae probably in permanent or semipermanent ground waters].

313. Culex (Mel.) nigrimacula Lane & Whitman, 1943 [probably not Melano-

conion|. Type: Holotype male, vicinity of Rio de Janeiro (Guanabara), Sept 1940,

L. Whitman (LU). Bionomics: Larvae in broad-leaved bromeliads.

314. Culex (Mel.) ocellatus Theobald, 1903 [probably not Melanoconion]. Type:

Holotype male, Sao Paulo (Sao Paulo), A. Lutz (BM; see Belkin, 1968:18). Bio-

nomics: Larvae in bromeliads.

315. Culex (Mel.) automartus Root, 1927 [= ocellatus]. Type: Holotype male,

Botanical Garden, Rio de Janeiro (Guanabara), 24 May 1925, F.M. Root (USNM).

Bionomics: Pupa in bromeliad.

316. Culex (Mel.) oedipus Root, 1927. Type: Lectotype male (8-1) with geni-

talia slide, Mage (Rio de Janeiro), 4 Feb 1925, F.M. Root (USNM; designation by

Rozeboom and Komp, 1950:94). Bionomics: Larvae in “‘jungle”’ pools.

317. Culex (Mel.) plectoporpe Root, 1927 [= phlogistus]. Type: Lectotype male

(109-1), Bangu, Rio de Janeiro (Guanabara), 11 June 1925, F.M. Root (USNM;

designation by Stone and Knight, 1957a:55). Bionomics: Larvae in ditch in small

pools full of grass and water weeds.

318. Culex (Mel.) putumayensis Matheson, 1934. Type: Holotype male with

genitalia slide, Amazon River, 7 Aug 1920, J.C. Bradley; TYPE LOCALITY here

Belkin et al: Topotypic Brazilian Mosquitoes 25

restricted to Santo Antonio do Ica (Amazonas) (USNM,50353). Bionomics: [Lar-

vae in permanent or semipermanent ground waters].

319. Culex (Mel.) rachoui Duret, 1968. Type: Holotype male (2258), Parago-

minas, municipio Capim (Para), 24 June 1964, J.P. Duret (A). Bionomics: [ Larvae

probably in permanent or semipermanent sround waters].

320. Culex (Mel.) serratimarge Root, 1927. Type: Holotype male, Sant’Anna

(Rio de Janeiro), 27 Apr 1925 (USNM). Bionomics: Larvae in a “qungle” pool.

321. Culex (Mel.) silvai Duret, 1968. Type: Holotype male (85), Caracarai (Ro-

raima), 6 July 1964, J.P. Duret (A). ea es [Larvae probably in permanent

or semipermanent ground waters].

322. Culex (Mel.) theobaldi (Lutz, 1904). Type: LECTOTYPE by present de-

signation, female (next to large pin) on mount with another female, Lagoa (Sao

Paulo), 6 Mar 1904, A. Lutz (BM; see Belkin, 1968:20-21). Bionomics: [Larvae

in shaded roadside borrow pit].

323. Culex (Mel.) chrysothorax (Newstead & Thomas, 1910) [= theobaldi].

Type: Lectotype female (160), inner Flores swamp, Pensador, near Manaus (Ama-

zonas), 12 July 1906 (BM; designation by Belkin, 1968:14). Bionomics: [As for

322. theobaldi|.

324. Culex (Mel.) thomasi Evans, 1924. Type: Holotype male (8.1) with attached

wing mount and 4 slides of genitalia, Manaus (Amazonas), 1910, H.W. Thomas

(BM). Bionomics: Larvae in swamp water from “‘Amatory”’ [?].

325. Culex (Mel.) trilobulatus Duret & Darreto, 1956. Type: Holotype aaie.

Rio Tamandua, Ribeirao Preto (Sao Paulo), Nov 1954, M.P. Barreto and J.P. Buret

(FMRP). Bionomics: [Larvae probably in permanent or semipermanent ground wa-

ters].

326. Culex (Mochlostyrax) megapus Root, 1927 [= alogistus|. Type: Holotype

male destroyed, only genitalia slide remaining, mountains near Angra dos Reis (Rio

de Janeiro), Jan 1925, N.C. Davis (USNM). Bionomics: Larva in ‘“‘jungle’’ pool.

327. Culex (Mochl.) galvaoi Duret, 1968. Type: Holotype male (2414), Chere

(Rio de Janeiro), 15 July 1953, J.P. Duret (A). Bionomics: [Larvae probably in

permanent or semipermanent ground waters].

328. Culex (Mochl.) innovator Evans, 1924. Type: Lectotype male (D4) with

attached wing mount and 4 genitalia slides, River Amazon, to or from Manaus,

1915, A.A. Clark; TYPE LOCALITY here restricted to Itacoatiara (Amazonas)

(BM; designation by Belkin, 1968:16). Bionomics: [Larvae probably in permanent

or semipermanent ground waters].

329. Culex (Mochl.) palaciosi Duret, 1968. Type: Holotype male (1817), Boa

Vista (Roraima), 8-9 July 1964, J.P. Duret (A). Bionomics: [Larvae probably in

permanent or semipermanent ground waters].

330. Culex (Microculex) albipes Lutz, 1904. Type: Holotype female, [Ilha de]

Itaparica (Bahia) (NE; no specimens in IOC or BM). Bionomics: Larva in bromeliads.

331. Culex (Micr.) aphylactus Root, 1927. Type: Lectotype male (79) with only

genitalia slide remaining, Serra dos Orgaos, woods near Rio Soberbo at crossing of

railroad from Mage to Teresopolis (Rio de Janeiro), 1 May 1925, F.M. Root (USNM;

designation by Stone and Knight, 1957a:43). Bionomics: Larvae in bromeliads.

332. Culex (Micr.) aureus Lane & Whitman, 1951. Type: Holotype male, Rio de

Janeiro (Guanabara), Aug 1940, L. Whitman (LU). Bionomics: Larvae in fallen

epiphytic tank bromeliad.

333. Culex (Micr.) carioca Lane & Whitman, 1951. Type: Holotype male, Rio de

Janeiro (Guanabara), Sept 1940, L. Whitman (LU). Bionomics: Larvae in tank bro-

26 ! Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

meliad.

334, Culex (Micr.) trychnus Root, 1927 [= consolator|. Type: Lectotype male

(79-2) with genitalia on slide, Serra dos Orgaos, woods near Rio Soberbo at crossing

of railroad between Mage and Teresopolis (Rio de Janeiro), 1 May 1925, F.M. Root

(USNM;; designation by Stone and Knight, 1957a:57). Bionomics: Larvae in brome-

liads. | |

335. Culex (Micr.) davisi Kumm, 1933. Type: Males, females, larvae, Salvador

(Bahia), 1931 (LU). Bionomics: Larvae in bromeliads.

336. Culex (Micr.) dubitans Lane & Whitman, 1951. Type: Holotype male, Rio

de Janeiro (Guanabara), Oct or Dec 1940, L. Whitman (LU). Bionomics: Larvae

in tank bromeliad.

337. Culex (Micr.) gairus Root, 1927. Type: Lectotype male (22c) with slides

of male genitalia and associated larval and pupal skins, Botanical Garden, Rio de

Janeiro (Guanabara), 15 Feb 1925, F.M. Root (USNM; designation by Stone and

Knight, 1957a:50). Bionomics: Larvae in bromeliads.

338. Culex (Micr.) hedys Root, 1927. Type: Holotype male, destroyed except

for genitalia slide, Angra dos Reis (Rio de Janeiro), Jan 1925, N.C. Davis (USNM).

Bionomics: Larvae in bromeliads.

339. Culex (Micr.) imitator Theobald, 1903. Type: Holotype male with attached

genitalia mount, Sao Paulo (Sao Paulo), A. Lutz (BM). Bionomics: Larvae in brome-

liads.

*340. Culex (Micr.) argenteoumbrosus (Theobald, 1907) [= imitator]. Type: Lec-

totype male with genitalia slide, Rio de Janeiro (Guanabara), Apr 1903, E.A. Goeldi

(BM; designation by Belkin, 1968:13). Bionomics: [Probably as for 339. imitator].

341. Culex (Micr.) retrosus Lane & Whitman, 1951 [= ssp. of imitator]. Type:

Holotype male, Rio de Janeiro (Guanabara), Sept 1939 or Oct 1940, L. Whitman

(LU). Bionomics: Larvae in terrestrial tank bromeliads.

342. Culex (Micr.) fuscatus Lane & Whitman, 1951 [= ssp. of inimitabilis]. Type:

Holotype male, Rio de Janeiro (Guanabara), Oct 1940, L. Whitman (LU). Bio-

nomics: Larvae in epiphytic tank bromeliad.

343. Culex (Micr.) intermedius Lane & Whitman, 1951. Type: Holotype male,

Rio de Janeiro (Guanabara), Nov 1940, L. Whitman (LU). Bionomics: Larvae in

tank bromeliads.

344, Culex (Micr.) lanei Coutinho & Forattini, 1962.. Type: Holotype male

(1B(6)7), with attached genitalia mount and slide of larval and pupal skins (2941,

marked cotype), Bertioga, municipio Santos (Sao Paulo), Aug 1961, O.P. Forattini

(FH,15062). Bionomics: Larvae in artificial containers placed in a forest.

345, Culex (Micr.) microphyllus Root, 1927. Type: Lectotype male (92-1) with

genitalia on slide, Mage (Rio de Janeiro), 26 May 1925, F.M. Root (USNM; de-

signation by Stone and Knight, 1957a:53). Bionomics: Larvae in bromeliads. |

346. Culex (Micr.) neglectus Lutz, 1904. Type: Lectotype male with genitalia

on slide (783; 108-34), Serra da Cantareira (Sao Paulo), 12-II-04, A. Lutz (USNM:

designation by Lane in Lane and Whitman, 1951:364). Bionomics: Larvae in bam-

boo.

347. Culex (Micr.) pleuristriatus Theobald, 1903. Type: Lectotype female, Sao

Paulo (Sao Paulo), A. Lutz (BM; designation by Belkin, 1968:19). Bionomics: [ Lar-

vae in terrestrial tank bromeliads] .

348. Culex (Micr.) reducens Lane & Whitman, 1951. Type: Holotype male, Rio

de Janeiro (Guanabara), Sept 1940, L. Whitman (LU). Bionomics: [Larvae pro-

bably in bromeliads].

Belkin et al: Topotypic Brazilian Mosquitoes 27

349. Culex (Micr.) shopei Forattini & Toda, 1966. Type: Holotype male (46U12)

with slides of male genitalia (3107) and larval and pupal skins (3107), Utinga, Belem

(Para), 4 Mar 1965, A. Toda (FH,15841 and 15842). Bionomics: Larvae in brome-

liads.

350. Culex (Micr.) worontzowi Pessoa & Galvao, 1936. Type: Holotype male

(1,Nt5), Tabatinguera (Sao Paulo), C. Worontzow (FMSP). Bionomics: Larvae in

bromeliads.

351. Culex (Aedinus) accelerans Root, 1927. Type: Holotype male, Porto das

Caixas (Rio de Janeiro), 15 Apr 1925, F.M. Root (USNM). Bionomics: Larvae in

dense vegetation in lagoon connected to river.

*352, Culex (Aed.) amazonensis (Lutz, 1905). Type: Syntypes males and females,

Amazon River below Manaus (Amazonas), Lindenberg (LU; not in IOC). Bionomics:

[Larvae in swamps, margins of lakes and permanent pools].

353. Culex (Aed.) hildebrandi Evans, 1923 [= amazonensis|. Type: Holotype

male (1/467) with slide of wing and 2 slides of genitalia, River Amazon on S.S.

‘Hildebrand’? on way to Manaus, 1922, A.A. Clark; TYPE LOCALITY here re-

stricted to Itacoatiara (Amazonas) (BM). Bionomics: [Probably as for 352. ama-

zonensis | .

354. Culex americanus (Neveu-Lemaire, 1902) [momen dubium; possibly = ama-

zonensis|. Type: Syntypes 4 females, Cunani (Amapa) [as French Guiana], Jan

1901, Mathis (NE; see Belkin, 1968:12-13). Bionomics: [Probably as for 352. ama-

zonensis | .

355. Culex (Aed.) clastrieri Casal & Garcia, 1968. Type: Holotype male, Belem

(Para), 29 Nov 1959, Eber and Evangelista (Casal). Bionomics: [Larvae probably

in swamps].

356. Culex (Anoedioporpa) belemensis Duret & Damasceno, 1955. Type: Holo-

type male (Br15,E10), Belem (Para), 19 Aug 1953, R.G. Damasceno and J.P. Duret

(Duret). Bionomics: [Larvae probably in treeholes].

357. Culex (Anoed.) canaanensis Lane & Whitman, 1943. Type: Holotype male,

Sao Joao de Petropolis, Vale do Canaa (Espirito Santo), Apr or July, 1940, L.

Whitman (LU). Bionomics: [Larvae probably in treeholes].

358. Culex (Anoed.) damascenoi Duret, 1969. Type: Holotype male (2275), Rio

Preto, municipio Joao Goulard (Amazonas), 18 July 1964, J.P. Duret (A). Bio-

nomics: [Larvae probably in treeholes]. _

359. Culex (Anoed.) luteopleurus (Theobald, 1903). Type: Holotype female,

[Belem] (Para), H.E. Durham (BM). Bionomics: [Larvae probably in treeholes].

360. Culex (Anoed.) originator Gordon & Evans, 1922. Type: Lectotype male

(13.2/463) with genitalia on 2 slides, 0.5 mi in forest, Macapa, near Manaus (Ama-

zonas), 21 Dec 1921, R.M. Gordon (BM; designation by Belkin, 1968:18). Bio-

nomics: Larvae in treehole, “‘Carapana uba’’.

361. Culex (Carrollia) anduzei Cerqueira & Lane, 1944. Type: Holotype male,

Rio Maues (Amazonas), Feb 1937, C. Worontzow (FH,4099). Bionomics: [Larvae

probably in treeholes, bamboo or plant material on ground].

362. Culex (Car.) antunesi Lane & Whitman, 1943. Type: Holotype male, Sao

Joao de Petropolis, Vale do Canaa (Espirito Santo), Apr or May 1940, L. Whitman

(LU). Bionomics: Larvae in bamboo internodes.

*363. Culex (Car.) iridescens (Lutz, 1905). Type: Neotype male with genitalia

on slide (222), Serra da Cantareira (Sao Paulo), Apr 1938, M. Sanches, E. Coimbra

and H. Guimaraes (FH,953; contrary to interpretation of Belkin, 1968:16, the

neotype designation of Antunes and Ramos, 1939:380-381 is probably correct as

28 > “Geviiinc: Merci diet: a6x6. aon

the Lutz material in IOC may not be part of type series). Bionomics: Larvae in

bamboo.

364. Culex (Car.) soperi Antunes & Lane, 1937. Type: Holotype male (886),

genitalia on 2 slides (99,100), Perus (Sao Paulo), 23 Apr 1937 (FH,721). Bionomics:

[Larvae in bamboo internodes] .

365. Culex (Car.) wilsoni Lane & Whitman, 1943. Type: Holotype male, Sao

Joao de Petropolis, Vale do Canaa (Espirito Santo), Apr 1940, L. Whitman (LU).

Bionomics: Larvae in bamboo internodes.

366. Culex pallipes Robineau-Desvoidy, 1827 [nomen dubium]. Type: Female(s),

Brazil, locality not specified; TYPE LOCALITY here restricted to vicinity of Rio de

Janeiro (Guanabara) (NE; see Belkin, 1968:19).

367. Culex molestus Kollar, 1832 [nomen dubium]. Type: Male(s) and female(s),

Rio de Janeiro (Guanabara) (NE).

368. Corethrella confusa Lane, 1939 [?= appendiculata|. Type: Holotype male

(1080) with genitalia on slide (279), Inhumas (Goias), 19 Apr 1935, J. Paternostro

(FH). Bionomics: Pupa in treehole. .

369. Corethrella bromelicola Lane, 1939. Type: ‘Holotype female (874) with

pupal skin on point and part of larval skin on slide (203), Poco Grande, Juquia

(Sao Paulo), Nov 1938, J. Lane (FH). Bionomics: Larva in epiphytic bromeliad.

370. Corethrella cardosoi Lane, 1942. Type: Holotype male (3297) with attached

genitalia mount, Casa Grande (Sao Paulo), Dec 1940 (FH). Bionomics: [Larvae

probably in bromeliads or.ground waters].

371. Corethrella edwardsi Lane, 1942. Type: Holotype female (3334), locality

not specified (Mato Grosso), July 1939; TYPE LOCALITY here restricted to Sa-

lobra (FH,1206). Bionomics: [Larvae probably in ground waters or bromeliads].

372. Corethrella flavitibia Lane, 1939. Type: Holotype female (1137), Juquia

(Sao Paulo), Jan 1939, J. Lane (FH,1067). Bionomics: [Larvae in bromeliads].

373. Corethrella fulva Lane, 1939. Type: Holotype male (868) with attached

genitalia mount, Agua Fria, Perus (Sao Paulo), June 1937, Ramalho and Vieira

(FH). Bionomics: Larvae in epiphytic bromeliads. |

374. Corethrella infuscata Lane, 1939. Type: Holotype female (1153), Juquia

(Sao Paulo), 20 Jan 1939, J. Lane (FH,1028). Bionomics: [Larvae in bromeliads] .

~ 375. Corethrella kummi Lane, 1942. Type: Holotype female, locality not speci-

fied (Bahia), 1931, H.W. Kumm; TYPE LOCALITY restricted to vicinity of Salvador

(BM). Bionomics: [Larvae probably in ground waters or bromeliads].

376. Corethrella lopesi Lane, 1942. Type: Holotype male (3282), Japuiba, Angra

dos Reis (Rio de Janeiro), Mar 1940, H. Sousa Lopes and J. Lane (FH). Bionomics:

[Larvae probably in bromeliads or around waters].

377. Corethrella pillosa Lane, 1939. Type: Holotype male (872) with genitalia

on slide (202), Poco Grande, Tuquia (Sao Paulo), 12 Dec 1938 (FH). Bionomics:

[Larvae probably in bromeliads or ground waters].

378. Corethrella selvicola Lane, 1939. Type: Holotype male (1146), Juquia (Sao

Paulo), 16-20 Jan 1939, J. Lane (FH,1030). Bionomics: [Larvae in running water

and epiphytic bromeliads, Lane and Cerqueira, 1958b:562].

379. Corethrella striata Lane, 1942. Type: Holotype female (3314), Palmeira

(Sao Paulo), Mar 1941, J.O. Coutinho (FH). Bionomics: [Larvae probably in brome-

liads or ground waters].

380. Corethrella tarsata Lane, 1942. Type: Holotype male with attached genitalia

mount, Camacari (Bahia), 1931, H.W. Kumm (BM). Bionomics: ee probably

in bromeliads)

Belkin et al: Topotypic Brazilian Mosquitoes 29

381. Corethrella travassosi Lane, 1942. Type: Holotype female (1206), Salobra

(Mato Grosso), July 1939 (FH,3266). Bionomics: [Larvae probably in bromeliads

or ground waters].

382. Corethrella vittata Lane, 1939. Type: Holotype male (1042) with genitalia

Slide (268), Juquia (Sao Paulo), 16-19 Jan 1939, J. Lane (FH). Bionomics: [Larvae

probably in bromeliads].

383. Corethrella whitmani Lane, 1942. Type: Holotype female (3311), locality

not specified (Espirito Santo), Aug 1940, L. Whitman; TYPE LOCALITY here re-

stricted to Sao Joao de Petropolis (FH). Bionomics: [Larvae in bromeliads].

384. Lutzomiops alticola (Lane, 1939). Type: Holotype male (378-M-18) with

genitalia slide (208), Campos do Jordao (Sao Paulo), elev. 1700 m, 13 Jan 1936,

F. Lane (FH,881). Bionomics: [Larvae probably in bromeliads].

385. Lutzomiops amazonicus (Lane, 1939). Type: Holotype female, Porto Velho,

Rio Madeira (Rondonia, as Amazonas), Mar-Apr 1931, R.C. Shannon (USNM). Bio-

nomics: [Larvae probably in bromeliads or ground waters].

386. Lutzomiops barrettoi (Lane, 1942). Type: Holotype male (461), Casa

Grande (Sao Paulo), Sept 1940, M.P. Barretto (FH,3367). Bionomics: [Larvae pro-

bably in bromeliads or ground waters].

387. Lutzomiops coutinhoi (Lane, 1942). Type: Holotype male (3365) with at-

tached genitalia mount, Palmeira (Sao Paulo), Mar 1941, J.0. Coutinho (FH). Bio-

nomics: [Larvae probably in bromeliads or ground waters].

388, Lutzomiops niger (Lane, 1939) [= davisi]. Type: Holotype male (875),

only | leg left on pin, genitalia (204) on slide together with those of paratype

(877), Maracaju (Mato Grosso), May 1937, R.C. Shannon (FH). Bionomics: [Lar-

vae in ground waters].

389. Lutzomiops iridescens (Lane, 1939). Type: Holotype female (1148), Juquia

(Sao Paulo), 19 Jan 1939, J. Lane (FH,1046). Bionomics: [Larvae in running water,

Lane and Cerqueira, 1958b:565].

390. Lutzomiops juquianus (Lane, 1939). Type: Holotype female (873), Poco

Grande, Juquia (Sao Paulo), 14 Dec 1938, J. Lane (FH). Bionomics: [Larvae in

ground pools].

391. Lutzomiops lutzi (Lane, 1942). Type: Holotype female (3362), Jaragua

(Sao Paulo), Sept 1940, M.P. Barretto (FH). Bionomics: [Larvae probably in bro-

meliads or ground waters]. |

392, Lutzomiops manaosensis Lane & Cerqueira, 1958. Type: Holotype male

(406-13) with associated larval and pupal skins on slide, genitalia not located, near

Manaus (Amazonas), N.L. Cerqueira (FH,13193). Bionomics: Larvae in swamps

and running water.

393. Lutzomiops nigrescens (Lane, 1942). Typé: Holotype female (3363), Mage

(Rio de Janeiro), 18 Mar 1940, R.C. Shannon (FH). Bionomics: [Larvae probably

in bromeliads or ground waters].

394. Sayomyia braziliensis (Theobald, 1901). Type: Holotype female, repre-

sented by slide of 1 wing and 1 leg, Brazil, locality not specified, A. Lutz; TYPE

LOCALITY here restricted to Sao Paulo (Sao Paulo) (BM). Bionomics: [Larvae

probably in ponds, lakes or reservoirs] .

395. Sayomyia antunesi (Lane, 1939) [= braziliensis]. Type: Holotype male

(1150), Juquia (Sao Paulo), Jan 1939, J. Lane (FH,1027). Bionomics: [As for

394, braziliensis |.

396. Sayomyia souzai (Lane, 1939). Type: Holotype male (1083) with genitalia

Slide (221), Santarem (Para), June 1931, R.C. Shannon (FH). Bionomics: [Larvae

30 | Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

in shallow forest ground pools].

397. Edwardsops brevisector (Edwards, 1931). Type: Holotype female, “ton board

S.S. Jerome”, Manaus (Amazonas), 24-27 April 1901 (BM). Bionomics: [Larvae

probably in permanent or semipermanent ground waters].

398. Edwardsops magnificus (Lane, 1942). Type: Holotype female (3223), Salo-

bra (Mato Grosso), 31 Jan 1941, F. Lane (FH). Bionomics: [Larvae probably in

permanent or semipermanent ground waters].

*399. Edwardsops unicolor (Lane, 1942). Type: Holotype male (1216) with geni-

talia mount, Salobra, bank of Rio Miranda (Mato Grosso), July 1939 (FH,3203).

Bionomics: [Larvae probably in permanent or semipermanent ground waters].

400. Dixella chapadensis (Lane, 1939). Type: Holotype male (845) with geni-

talia on slide with another specimen, Portinha, Chapada region (Mato Grosso), 27

Aug, J. Lane (FH). Bionomics: [Larvae probably in permanent or semipermanent

ground waters].

401. Dixella limai (Santos, 1940). Type: Adult(s), larvae, pupae, Uberaba (Minas

Gerais) (Univ. Minas Gerais). Bionomics: Larvae in small streams.

402. Dixella paulistana (Lane, Forattini & Rabello, 1955). Type: Holotype male

(8094) with larval and pupal skins on slide, Campo Experimental da Agua Funda,

Sao Paulo (Sao Paulo), 5 Nov 1954, E.X. Rabello (FH). Bionomics: Larvae in a

rather shallow, sunlit pond with turbid water; in muddy margins with sparse emer-

gent vegetation. |

403. Dixella torrentia (Lane, 1939). Type: Holotype male (847), genitalia intact,

Salto do Poco Grande, Juquia (Sao Paulo), 13 Dec 1938, J. Lane (FH). Bionomics:

[Larvae probably in stream margins].

404. Dixella wygodzinskyi (Lane, 1945). Type: LECTOTYPE by present desig-

nation, female (5653), marked as holotype by Lane but not so indicated in original

publication, Vila Capivari, Campos do Jordao (Sao Paulo), elev. 1700 m, Apr 1935,

P. Wygodzinsky (FH). Bionomics: Larvae in epiphytic bromeliads in Araucaria for-

est.

AUTHORS

Amaral, A.D. Franco. See Galvao for the 2 species described by Galvao and

Amaral.

Antunes, P.C.A. The only Brazilian species described by Antunes were: 2 as senior

author with Lane, 243. Aedes (O.) pennai and 364. Culex (Car.) soperi, whose types

are in FH; 1 with Lane as senior author, 200. Coquillettidia (R.) shannoni, whose

holotype is also in FH; and | with Cerqueira as senior author, 261. Haemagogus

(L.) tropicalis, whose location is unknown (LU).

Barretto (also Barreto), M. Pereira. Of the 3 species described by Barretto with

Coutinho as junior author, all with designated holotypes, we have found only 205.

Mansonia (M.) pessoai, whose holotype is in FH (see section on depositories). The

location of the holotypes of the other 2 species: 202. Mansonia (M.) cerqueirai and

208. Mansonia (M.) wilsoni, is unknown (LU).

See also Duret for 3 species described by Duret and Barretto, and Galvao for 1

species described by Ayroza Galvao and Barretto.

Berlin, Olavil George William. The holotype of the only Brazilian species des-

cribed by Berlin is in USNM: 257. Aedes (H.) aureolineatus.

Belkin et al: Topotypic Brazilian Mosquitoes 31

Bonne-Wepster, Jean and Cornelis Bonne. The holotype of 158. Phoniomyia tri-

partita, the only Brazilian species described by these authors, is non-existent (NE).

It is represented by fig. 2 in Dyar (1919); see Belkin, 1968:24.

Bourroul, Celestino. The type material of the only species described by Bour-

roul is non-existent (NE): 77. Toxorhynchites (L) mariae. See Belkin (1968:51) for

other names attributed to Bourroul in the past.

Carpenter, Stanley J. See Galindo for the | aheuins from Brazil described at Gal-

indo, Carpenter and Trapido.

Casal, Osvaldo H. The holotype of 355, Culex (Aed.) clastrieri described by Casal

with Garcia as junior author is in the private collection of Casal and will presumably

be eventually deposited in BA.

Castro, Gustavo M. de Oliveira. We have designated a lectotype, in IOC, for 1 of

the 2 species described by Castro: 269. Culex (C.) carcinoxenus. The location of

the holotype of 162. Limatus flavisetosus is unknown (LU), we did not find it in

IOC where it was supposedly deposited.

Causey, Ottis R. The holotype of the 1 Brazilian species described by Causey as

sole author is in USNM: 57. Anopheles (N.) dunhami. Of the 2 species described by

Causey, Deane and Deane, we have designated a lectotype, in FMP, for: 48. Ano-

pheles (N.) galvaoi. No material of the other species, based on eggs, 5. Chagasia roze-

boomi, has been located and it is safe to consider that its type is non-existent (NE).

We have restricted the type locality of this species to the vicinity of Sao Benedito,

the locality in the state of Ceara where other Causey material was collected. The

fourth species: 38. Anopheles (N.) sawyeri, was described by Causey with Deane,

Deane and Sampaio as junior authors; its holotype is in USNM.

Cerqueira, Nelson L. The holotypes of all 8 Brazilian species described by Cer-

queira as sole author are in existence: | in IOC, 230. Psorophora (J.) forceps; and

7 in FH: 173. Sabethes (S.) forattinii; 174. Sabethes (S.) lanei; 179. Sabethes (S.)

shannoni; 180. Sabethes (S.) spixi, 183. Sabethes (Sabethoides) tridentatus; 227.

Psorophora (J.) amazonica; 262. Haemagogus (S.) baresi.

The location of the holotype of 261. Haemagogus (L.) tropicalis, ies Lae by

Cerqueira with Antunes as junior author, is unknown (LU); it was not found in

IOC or FH.

The location of the holotypes of 236. Aedes (O.) aenigmaticus and 244. Aedes

(O.) perventor, described by Cerqueira with Costa as junior author, is unknown

(LU); they were not found in IOC or FH.

The holotype of 361. Culex (Car.) anduzei, described by Cerqueira with Lane as

junior author, is in FH.

The location of the holotype of 242. Aedes (O.) lepidus, described by Cerqueira

and Paraense, is unknown (LU); it was not found in IOC or FH.

Cerqueira was also the junior author of 1 species with Correa (see), 4 species

with Del Ponte (see), 34 species with Lane (see) and 1 species with Shannon (see).

Cerqueira’s list of Amazonian species (1961) is the most useful source of informa-

tion on the bionomics and distribution of Brazilian mosquitoes.

Chagas, Carlos. We have designated lectotypes for 2 of the 4 species described

by Chagas, both in IOC: 39. Anopheles (N.) braziliensis (type locality restricted

to vicinity of Lassance on the basis of the statement in Root, 1926:704); 196.

Coquillettidia (R.) juxtamansonia. Type material of 6/. Anopheles (N.) parvus may

32. Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

also be in IOC but since the only specimen remaining in the collection bears no data

we hesitate to designate it as lectotype; the type locality is stated to be Oliveira

(Minas Gerais) by Galvao, 1941:537 and is one of the localities mentioned for this

species by Peryassu (1908:60). We have not been able to locate (LU) any original

material of 53. Anopheles (N.) nigritarsis whose type locality we have determined

to be Oliveira also on the basis of the citation of this locality in Peryassu (1908:60).

Christophers, S. Rickard. The holotype of the only Brazilian species described by

Christophers, 16. Anopheles (A.) amazonicus, is in BM. We have restricted its ori-

ginal indefinite type locality (River Amazon) to Manaus.

Correa, Renato R. Correa is the sole author of only 2 nominal species: the type

material of 10. Anopheles (A.) geometricus is non-existent (NE); the holotype of

67. Anopheles (K.) montemor, originally in SPM, is now in FH.

The holotype of the 1 species described by Correa and Cerqueira, 68, Anopheles

(K.) laneanus, is in FH.

The holotypes of all 5 species described by Correa and Ramalho are in FH: &.

Anopheles (A.) anchietai; 148. Phoniomyia galvaoi; 151. Phoniomyia lopesi; 271.

Culex (C.) deanei; 274, Culex (C.) forattinii,

The holotype of 32. Anopheles (N.) imperfectus, described by Correa and Ramos

and originally in SPM, is now in FH.

Correa was also the junior author with Galvao and Lane of 42. Anopheles (N.)

paulistensis. The location of the type material of this species is not known (LU).

Costa, Almir F : For the 2 species described by Costa with Cerqueira as senior au-

thor, see under the latter.

Coutinho, Jose Oliveira. The holotype of 344. Culex (Micr.) lanei described by

Coutinho and Forattini is in FH. Coutinho also proposed the replacement name

costalimai for 19. Anopheles (A.) limai (FH).

See also Barretto for 3 species described by Barretto and Coutinho, and Lane for

2 species described by Lane and Coutinho.

Cruz, Oswaldo Goncalves. We have designated a lectotype for only 1 of the 4 spe-

cies described by Oswaldo Cruz (in IOC): 225. Psorophora (P.) genumaculata. Au-

thentic original material of 50. Anopheles (N.) lutzii is in the IOC collection but it

requires careful study before designation of a lectotype. Also in IOC is a slide with

2 wings labelled Manguinhosia lutzi; this may be the only remaining original material

of 22. Anopheles (A.) lutzi but this also requires study. The location of the type ma-

terial of 2. Chagasia neivae remains unknown (LU), it was not found in IOC where it

should be but Belkin’s search was not exhaustive. The localities of the species des-

cribed by Oswaldo Cruz are given in Peryassu (1908).

Damasceno, Reinaldo G. See Duret for 2 species described by Duret and Damas-

ceno, and Galvao for 3 species described by Galvao and Damasceno.

Davis, Nelson C. The holotype of the only Brazilian species described by Davis as

sole author is in USNM: 27. Anopheles (A.) shannoni. See also Serafim for 1 species

described by Serafim with Davis as junior author.

Deane, Leonidas M. and M.P. Deane. See Causey.

Del Ponte, Eduardo. In the world catalog (Stone, Knight and Starcke, 1959:87),

2 Brazilian species, 136. Wyeomyia (D.) rooti and 139. Wyeomyia (D.) subcom-

plosa, are credited to Del Ponte (1939) as sole author in a table (Del Ponte, 1939:

Belkin et al: Topotypic Brazilian Mosquitoes 33

541) listing coded character states of females without mention of type specimens.

Stone, Knight and Starcke (loc. cit.) indicate the state of Goias as the type locality

of these nominal species, apparently on the basis of the initial sentence in Del

Ponte’s paper (1939:535), which however merely states that the preparation of Del

Ponte’s work was initiated at the time of his study of the mosquitoes of Goias.

There are a number of unfortunate errors in Del Ponte’s paper (1939), among

them: (1) a reference to a prior non-existent description of rooti (p. 536) by Del

Ponte and Cerqueira (1938), (2) no reference to undulata described by these au-

thors, and (3) a discrepancy in 2 important character states attributed to kerri in the

table with those appearing in the original description of Del Ponte and Cerqueira

(1938:228,229). Fortunately these errors do not affect the availability of rooti

Del Ponte, 1939. There is no doubt that type material of rooti Del Ponte, 1939 was

in the hands of Lane and Cerqueira (1942:690) when they described delpontei in

the belief that the former name had not been proposed (Lane and Cerqueira, 1957).

In IOC there is a notation by Costa Lima that the type of rooti was loaned to the

Rockefeller Foundation, which supported in part the studies of Lane and Cerqueira

~ (1942:473). In spite of Lane and Cerqueira’s statement (1957) that they did not

describe delpontei until material additional. to the rooti of Del Ponte came to hand,

we believe that the specimen which they designated as the holotype of delpontei was

one of the original specimens of rooti. The date and locality of capture of this speci-

men agree with others mentioned by Del Ponte and Cerqueira (1938) and it was

probably collected by Gastao Cesar. Since Del Ponte (1939) did not designate a

holotype, we have designated here as lectotype of 136. Wyeomyia (D.) rooti, Del

- Ponte, 1939 the same specimen as the holotype of 137. Wyeomyia (D.) delpontei

Lane & Cerqueira, 1942 (IOC). The latter nominal species was synonymized by Lane

and Cerqueira (1957) with rooti Del Ponte, 1939 when they were informed by Alan

Stone of the validity of Del Ponte’s description. At the same time Lane and Cer-

queira (1957) proposed the name alani to replace the preoccupied specific name of

Wyeomyia (A.) rooti described by them in 1942.

The character states given by Del Ponte for 139. Wyeomyia (D.) subcomplosa

are identical to those indicated for complosa. We believe that Del Ponte intended

to propose subcomplosa for the Brazilian species identified earlier as complosa

Dyar, 1928 by Del Ponte and Cerqueira (1938:234-235). The latter authors pointed

out some differences in the male genitalia of their complosa from the description

and figure by Dyar (1928:74, fig. 45). Therefore, we consider that all the females

listed under complosa by Del Ponte and Cerqueira are syntypes of subcomplosa, Un-

fortunately the male of complosa of Del Ponte and Cerqueira, with the distinctive

genitalia, cannot be included in the type series of subcomplosa as this sex is not

mentioned in Del Ponte (1939). The location of the female syntypes of subcomplosa

is unknown (LU) at present but it is very likely that some of these specimens will be

found in FH, IOC or USNM. To our knowledge, subcomplosa has not been described

or recognized as a distinct species since the original mention of the name by Del

Ponte (1939). Its taxonomic identity remains uncertain.

The holotypes of 3 of the 4 species described by Del Ponte with Cerqueira as jun-

ior author are in IOC: 124. Wyeomyia (D.) cesari; 128. Wyeomyia (D.) kerri; 141.

Wyeomyia (D.) undulata. We did not find the holotype of 135. Wyeomyia (D.)

brucei; in IOC there is only the “‘allotype” designated by Lane and Cerqueira (1942:

599); its location is considered unknown (LU) for the present but it may be lost.

Duret, Jose Pedro. The holotypes of all 15 Brazilian species described up to 1970

by Duret as sole author are all in the Duret private collection: 278. Culex (C.) para-

34 | Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

maxi; 286. Culex (Mel.) bahiensis; 292. Culex (Mel.) contei; 293. Culex (Mel.) cris-

tovai; 297. Culex (Mel.) ernanii; 300. Culex (Mel.) faurani; 301, Culex (Mel.) flochi;

306. Culex (Mel.) isabelae; 307. Culex (Mel.) johnnyi; 308. Culex (Mel.) kerri; 319.

Culex (Mel) rachoui; 321. Culex (Mel.) silvai; 327. Culex (Mochl.) galvaoi; 329.

Culex (Mochl.) palaciosi; 358, Culex (Anoed.) damascenoi.

The holotypes of all 3 species described by Duret with Barretto as junior author

are in FMRP: 285, Culex (Mel.) aureonotatus; 289, Culex (Mel.) bifoliolatus; 325.

Culex (Mel.) trilobulatus.

The holotypes of the 2 species described by Duret with Damasceno as junior au-

thor are in the Duret private collection: 290. Culex (Mel.) mojuensis; 356. Culex

‘(Anoed.) belemensis.

Dyar, Harrison G. The holotypes or lectotypes of all 5 species described from

Brazil by Dyar as sole author are in USNM: 102. Trichoprosopon (R.) trichopus;

132. Wyeomyia (D.) mystes; 197. Coquillettidia (R.) hypocindyna; 264. Culex (L.)

brasiliae; 303. Culex (Mel.) inadmirabilis. Dyar also proposed the replacement names

remipusculus for 177. Sabethes (S.) purpureus Peryassu, 1908 (IOC) and melano-

nymphe for 188. Sabethes (Sabethinus) albiprivatus (BM).

The location of the holotype of the 1 Brazilian species described by Dyar with

Knab as junior author is unknown (LU): 84. Toxorhynchites (L.) ambiguus. This

species was never seen by the authors and was based on a single specimen mentioned

by Wiedemann (see Belkin, 1968:32). This specimen may possibly be in NMW. Dyar

and Knab also proposed the replacement names peryassui for 22. Anopheles (A.)

lutzi (210C), cruzii for 66. Anopheles (K.) lutzii Theobald, 1901 (BM), and wiede-

manni for 81. Toxorhynchites (L.) ferox (Wiedemann, 1828) (SNG and NMW).

The holotypes of all 4 Brazilian species described by Dyar with Shannon as junior

author are in USNM: 24. Anopheles (A.) celidopus; 182. Sabethes (Sabethoides)

glaucodaemon, 204. Mansonia (M.) indubitans; 288. Culex (Mel.) bequaerti. Dyar

and Shannon also proposed the replacement name Jeucocelaenus for 253. Aedes

(F.) leucomelas (BM).

Edwards, Frederick W. The 2 holotypes and 1 lectotype of the only 3 Brazilian

species described by Edwards are in BM: 166. Sabethes (S.) longfieldae; 169, Sa-

bethes (S.) argyronotum; 397. Edwardsops brevisector.

Evans, Alwen M. The holotypes or lectotypes of all 7 Brazilian species described

by Miss Evans as sole author are in BM: 283. Culex (Mel) gordoni; 287. Culex

(Mel.) innominatus (type locality here restricted); 296. Culex (Mel) manaosensis;

311, Culex (Mel.) clarki (type locality here restricted); 324. Culex (Mel) thomasi;

328. Culex (Mochli.) innovator (type locality here restricted); 353. Culex (Aed.)

hildebrandi (type locality here restricted). The restricted type locality for 3 of the

species noted above is Itacoatiara when the original description or labels indicated

River Amazon to or from Manaus. For the fourth species, clarki, which was stated

to be collected on the River Amazon, the restriction is to Manaus.

See also Gordon for 6 species described by Gordon and Evans.

Ferreira, Orlando. See Lima for the 1 species described by Lima, Guitton and

Ferreira.

Fonseca, Flavio da. The holotypes of the 2 nominal species described by Fonseca

and Ramos are in FH: 19. Anopheles (A.) limai; 20. Anopheles (A.) costai.

Forattini, Oswaldo Paulo. The holotype of the 1 Brazilian species credited to

Belkin et al: Topotypic Brazilian Mosquitoes 35

Forattini as sole author is in USNM: 277. Culex (C.) oswaldoi, The holotypes of the

2 Brazilian species described by Forattini as senior author are in FH: 268. Culex

(C.) brami Forattini, Rabello & Lopes; 349. Culex (Micr.) shopei Forattini & Toda.

See also Coutinho for 1 species described by Coutinho and Forattini; and Lane

for 1 species described by Lane and Forattini and 1 species described by Lane,

Forattini and Rabello.

Gabaldon, Arnoldo. See Rozeboom for 1 Brazilian species described by Roze-

boom and Gabaldon.

Galindo, Pedro. The holotype of 254. Aedes (F.) leucophoebus, the only Bra-

zilian species described by Galindo, with Carpenter and Trapido as junior authors,

is in FH.

Galvao, A.L. Ayroza. Type material of 2 of the 3 species described by Ayroza

Galvao as sole author have been located in FH: the holotype of 9. Anopheles (A.)

bustamentei and the lectotype, here designated, of 26. Anopheles (A.) rachoui. The

location of the type material of 65. Anopheles (N.) chagasi is unknown (LU); the

type locality is here restricted to Manaus, | of 3 localities mentioned in the original

description. |

The holotypes of the 2 species described by Ayroza Galvao and Amaral are in

FMSP (see section on depositories): 34. Anopheles (N.) antunesi and 49. Anopheles

(N.) lanei.

The holotype of 70. Anopheles (L.) pseudotibiamaculatus described by Ayroza

Galvao and Barretto is also in FMSP (see section on depositories).

Syntypes of the 3 species described by Ayroza Galvao and Damasceno were pro-

bably originally in FMSP as stated for 60. Anopheles (N.) konderi. However, Belkin

could not locate any original material of this species which is recorded in this col-

lection and it is probably lost. There is some material in this collection of 31. Ano-

pheles (N.) marajoara that could probably be designated as the lectotype. The type

locality of this species is here restricted to the vicinity of Cachoeira do Arari, the

site of collection of the following species. No material or record of the third species,

33. Anopheles (N.) domesticus, was located in FMSP; for the present the location of

its type material is unknown (LU).

Of the 4 species described by Ayroza Galvao and Lane, the holotype of 40. Ano-

pheles (N.) pessoai was found in FMSP. The only type material of 54. Anopheles

(N.) noroestensis found by Belkin was in FH and possibly FMSP; the lectotype here

designated is represented by the male genitalia slide in FH. The type locality of this

species is Lussanvira (Galvao and Lane, 1938:176,177) and not Tiete as stated by

Stone, Knight and Starcke (1959:33). No type material of 30. Anopheles (N.)

limai was found, only slides of stomachs in FMSP; it is considered non-existent

(NE). We have designated a lectotype (USNM) for 59. Anopheles (N.) metcalfi,

which was based on tarsimaculatus of Root (1926).

No material of 42. Anopheles (N.) paulistensis described by Ayroza Galvao, Lane

and Correa, was found by Belkin; the location of the type material remains un-

known (LU).

See Pessoa for | species described by Pessoa and Galvao.

Garcia, Miguel. See Casal.

Giles, George M. The holotype of the only species described from Brazil by Giles

is in BM: 229. Psorophora (J.) arribalzagae.

36 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

Gordon, Rupert M. The holotypes or lectotypes of all 6 species described from

Brazil by Gordon with Evans as junior author are in BM: 76. Toxorhynchites (L.)

horei; 133. Wyeomyia (D.) negrensis; 165. Sabethes (S.) amazonicus; 209. Urano-

taenia albitarsis; 249, Aedes (F.) braziliensis; 360. Culex (Anoed.) originator.

Guimaraes, Lindolpho da Rocha. See Lane for the | species described by Lane

and Guimaraes.

Guitton, Neide. See Lima for the | ee described by Lima, Guitton and

Ferreira.

Knab, Frederick. See Dyar.

Kollar, Vincenz. The type feel of 367. Culex molestus, a nomen dubium, is

non-existent (NE).

Komp, William H.W. The holotype of 35. Anopheles (N.) emilianus, the only

species described by Komp from Brazil, is in USNM.

~ Kumm, Henry W. The location of the type material is unknown (LU) for the only

species described from Brazil by Kumm: 335. Culex (Micr.) davisi.

_ Lane, John. Holotypes (in 1 case the lectotype) of 41 of the 43 species described

from Brazil by Lane as sole author have been located. In USNM is the holotype of

385. Lutzomiops amazonicus and in BM the holotypes of 375. Corethrella kummi

(type locality restricted) and 380. Corethrella tarsata. The FH contains the types

of 38 species: 96. Trichoprosopon (R.) luederwaldti; 108. Wyeomyia (W.) downsi;

210. Uranotaenia davisi; 212, Uranotaenia burkii; 214. Uranotaenia mathesoni; 265.

Culex (C.) abnormalis; 267. Culex (C.) airozai; 273. Culex (C.) foliaceus; 276, Culex

(C.) mauesensis; 368. Corethrella confusa; 369. Corethrella bromelicola; 370. Core-

thrella cardosoi; 371. Corethrella edwardsi (type locality restricted); 372. Core-

thrella flavitibia; 373. Corethrella fulva; 374. Corethrella infuscata; 376. Corethrella

lopesi; 377. Corethrella pillosa; 378. Corethrella selvicola; 379. Corethrella striata;

381. Corethrella travassosi; 382. Corethrella vittata; 383. Corethrella whitmani (type

locality restricted); 384. Lutzomiops alticola; 386. Lutzomiops barrettoi; 387. Lut-

zomiops coutinhoi; 388. Lutzomiops niger; 389. Lutzomiops iridescens; 390. Lutzo-

miops juquianus; 391. Lutzomiops lutzi; 393. Lutzomiops nigrescens; 395. Lutzo-

miops antunesi; 396. Sayomyia souzai; 398. Edwardsops magnificus; 399. Edward-

sops unicolor; 400. Dixella chapadensis; 403. Dixella torrentia; 404. Dixella wygod-

zinskyi (lectotype). The location of the type material (holotypes not designated) '

of the following 2 species is unknown (LU), we did not find any material in FH or

IOC: 109. Wyeomyia (W.) leucotarsis and 119. Wyeomyia (C.) mattinglyi (type

locality restricted).

The holotype of the 1 species described by Lane with Antunes as junior author

is in FH: 200. Coquillettidia (R.) shannoni.

The holotypes (in 2 cases lectotypes) of all 34 species described by Lane with

Cerqueira as junior author have been located. We have designated a lectotype

(USNM) for 146. Phoniomyia edwardsi. In FH are the types of 117. Wyeomyia

(C.) dyari (lectotype) and 392, Lutzomiops manaosensis. Holotypes of 31 species

are in IOC: 90. Trichoprosopon (T.) obscurum; 92. Trichoprosopon (T.) soaresi;

97. Trichoprosopon (C.) walcotti; 98. Trichoprosopon (R.) castroi; 100. Tricho-

prosopon (R.) edwardsianum; 101. Trichoprosopon (R.) humboldti; 104. Tricho-

prosopon (R.) reversum; 105. Trichoprosopon (R.) simile; 106. Trichoprosopon

(R.) theobaldi; 110. Wyeomyia (W.) limai; 115. Wyeomyia (W.) sabethea; 118.

Belkin et al: Topotypic Brazilian Mosquitoes 37

Wyeomyia (C.) kummi; 121. Wyeomyia (A.) rooti (alani, new name); 122. Wyeo-

myia (D.) airosai; 126. Wyeomyia (D.) finlayi; 127. Wyeomyia (D.) howardi; 129.

Wyeomyia (D.) knabi; 137. Wyeomyia (D.) delpontei; 138. Wyeomyia (D.) shan-

noni; 140. Wyeomyia (D.) tarsata; 143. Phoniomyia bonnei; 144. Phoniomyia davisi,

147. Phoniomyia flabellata; 153. Phoniomyia palmata; 156. Phoniomyia neivai, 157.

Phoniomyia theobaldi; 167. Sabethes (S.) batesi; 185. Sabethes (Sabethinus) fabri-

cii; 186. Sabethes (Sabethinus) lutzianus; 189. Sabethes (Sabethinus) soperi; 190.

Sabethes (Sabethinus) whitmani. There is some confusion regarding the holotypes

of 143. Phoniomyia bonnei; 144. Phoniomyia davisi; 156. Phoniomyia neivai and

157, Phoniomyia theobaldi. There are specimens of all 4 species labelled as holo-

types in both IOC and FH. We consider that in all cases the true holotypes are in

IOC and that the specimens in FH were so marked by Lane before he undertook

his joint work with Cerqueira and inadvertently the type labels were not removed

later.

The holotypes of the 2 species described by Lane with Coutinho as junior author

are in FH: 195. Coquillettidia (R.) hermanoi; 199. Coquillettidia (R.) neivai.

The holotypes of 145. Phoniomyia diabolica, described with Forattini as junior

author, and 402. Dixella paulistana, described with Forattini and Rabello as joint

authors, are in FH.

Syntypes of 142. Phoniomyia antunesi, described by Lane with Guimaraes as

junior author, are in FH. We have not designated a lectotype for this species since

the type material requires careful study.

The holotype of 280. Culex (C.) renatoi, described by Lane with Ramalho as

junior author, is in FH.

We did not locate the holotypes of the 11 species described by Lane with Whit-

man (LU). They were presumably originally in the Rockefeller Foundation Labora-

tory at IOC: 313. Culex (Mel) nigrimacula; 332. Culex (Micr.) aureus; 333. Culex

(Micr.) carioca; 336. Culex (Micr.) dubitans; 341, Culex (Micr.) retrosus; 342. Culex

(Micr.) fuscatus; 343, Culex (Micr.) intermedius; 348. Culex (Micr.) reducens; 357.

Culex (Anoed.) canaanensis; 362. Culex (Car.) antunesi; 365, Culex (Car.) wilsoni.

In addition to the description of the above 95 species, Lane participated as jun-

ior author in the description of 8 species: 2 species with Antunes: 243. Aedes (O.)

pennai and 364. Culex (Car.) soperi, whose holotypes are FH; 1 species with Cer-

queira: 361. Culex (Car.) anduzei (FH); 4 species with Ayroza Galvao: 30. Ano-

pheles (N.) limai; 40. Anopheles (N.) pessoai; 54. Anopheles (N.) noroestensis and

59, Anopheles (N.) metcalfi, for the type material of these species see Galvao; and

1 species, 42. Anopheles (N.) paulistensis, with Ayroza Galvao and Correa, the lo-

cation of whose type is unknown (see Galvao). |

Laveran, Charles-Louis Alphonse. Type material of 160. Limatus curvirostris, the

only species described from Brazil by Laveran, is non-existent (NE).

Lima, Angelo da Costa. Type material of all but 1 of the 10 Brazilian nominal

species described by Costa Lima as sole author are in the IOC collection. The fol-

lowing 6 species are represented by holotypes or lectotypes: J//. Anopheles (A.)

evandroi; 93. Trichoprosopon (L.) brevipes; 114. Wyeomyia (W.) pintoi; 203. Man-

sonia (M.) chagasi; 220. Orthopodomyia townsendi; 221. Orthopodomyia sampaioi.

The following 3 species are represented by syntypes that need careful study before

designating lectotypes: 21. Anopheles (A.) minor; 78. Toxorhynchites (L.) pusil-

lus; 111. Wyeomyia (W.) lutzi. Belkin did not find any type material of 245. Aedes

(O.) rhyacophilus in IOC where it was deposited; its location is considered unknown

38 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

(LU) for the present as it may have been borrowed. We have restricted the type lo-

cality of this species to the vicinity of Sao Joao de Petropolis, one of the cities in

the valley of Canaa mentioned in the original description.

Costa Lima was the senior author with Guitton and Ferreira of 1 species, 71.

Toxorhynchites (A.) catharinensis, whose holotype is in IOC.

Judging by the notation on the record cards, Costa Lima was responsible for the

reorganization of the mosquito collection at the Instituto Oswaldo Cruz. The speci-

mens were apparently transferred from small insect boxes into individual tubes. The

majority of them are without labels on the pins and with fragmentary data only on

the tubes or with brief notation on the record cards. The transfer was not com-

pleted, the remainder of the specimens are in 10 small boxes in 2 drawers (see

Lutz).

Lopes, Oscar de Souza. See Forattini for the 1 species described by Forattini,

Rabello and Lopes.

- Lutz, Adolpho. After examining Lutz material in European museums, Belkin

(1968:48-51) discussed the involved problem of the determination of the type

specimens of the nominal species described by this author. In July and August

1969 Belkin found that the only remaining authentic type ‘material of Lutz in

Brazil was in the Instituto Oswaldo Cruz in Rio de Janeiro. This material is in

general in poor condition and often without labels on the pins. Part of it was appar-

ently studied by A. da Costa Lima and incorporated by him into the general col-

lection of the institute, usually with only the notation that the specimens were from

the old collection of the institute; the majority of these specimens have no labels at

all on the pins. The remainder of the Lutz material, apparently not previously

studied, is stored separately in 15 small insect boxes, 1-9 in one drawer and 10-15

in another. These specimens are all numbered (2550-2664) but not all numbers in

this series are present and it is evident that some specimens were removed from these

boxes, probably by Costa Lima. No record cards could be located in the institute for

the numbers on the specimens remaining in the small boxes. A typed species label

precedes the series of specimens of each species. Some specimens bear labels with

locality and data, others lack labels. The data on the labels of only a few specimens

indicate that they were undoubtedly part of the original type series of some of

Lutz’s species.

Since Lutz did not designate holotypes for any of his species described from more

than 1 specimen and since apparently all the remaining type material has now been

located in the BM, IOC or USNM collections, we have designated here lectotypes for

all his species except as noted below. These lectotypes have been designated with

due regard to the original data, agreement with original description as well as the

current interpretation of the involved species.

Lutz was the sole author of 33 nominal species. Type material of the following

is in existence. In the BM collection, 13 species: 79. Toxorhynchites (L.) solstitialis;

91. Trichoprosopon (T.) pailidiventer; 112. Wyeomyia (W.) medioalbipes; 113. Wye-

omyia (W.) oblita (syntype; also IOC); 134. Wyeomyia (D.) personata,; 164. Sabe-

thes (S.) albiprivatus; 184. Sabethes (Sabethinus) aurescens; 187. Sabethes (Sabe-

thinus) intermedius; 250. Aedes (Finlaya) fluviatilis; 253. Aedes (F.) leucomelas;

256, Aedes (F.) oswaldi; 281, Culex (C.) spinosus; 322, Culex (Mel.) theobaldi.

In the USNM collection, the lectotype of 346. Culex (Micr.) neglectus, In the IOC

collection, 3 species: 107, Wyeomyia (W.) arthrostigma,; 116. Wyeomyia (W.) ser-

rata; 218. Orthopodomyia albicosta, The following 6 species are apparently repre-

Belkin et al: Topotypic Brazilian Mosquitoes 39

sented by type material in IOC but lectotype designations have not been made

either because of fragmentary data or because of the need of careful study of the

specimens: 6. Anopheles (S.) lineatus; 86. Trichoprosopon (T.) compressum; 95,

Trichroprosopon (S.) longipalpis; 113. Wyeomyia (W.) oblita (also BM); 299, Cu-

lex (Mel.) fasciolatus; 309. Culex (Mel.) lugens. The neotype of 363. Culex (Car.) iri-

descens is in FH. The types of the following 3 species, stated to be in IOC by Lane

(1953) and Stone, Knight and Starcke (1959) were not found in this collection in

1969: 1. Chagasia fajardi; 263. Haemagogus (S.) capricornii; 352. Culex (Aed.) ama-

zonensis,; their location is unknown (LU) at present. No type material of the follow-

ing 7 species has been found by Belkin or other workers in the past anywhere and is

considered non-existent (NE): 74. Toxorhynchites (A.) neglectus; 88. Trichoproso-

pon (T.) splendens; 120. Wyeomyia (M.) leucostigma; 123. Wyeomyia (D.) bourrouli,

125. Wyeomyia (D.) confusa; 258. Aedes (H.) fulvithorax; 330, Culex (Micr.) albipes,

Lutz also described 4 species with A. Neiva as junior author. Type material of all

these species is apparently in IOC, the holotype of 15. Anopheles (A.) mattogrossen-

sis and probably the syntypes of 75. Toxorhynchites (L.) bambusicola; 82. Toxor-

hynchites (L.) posticatus and 234. Psorophora (G.) scutipunctata. Lectotypes for

the latter 3 species are not designated at this time because of the need of further

study.

Several species described by Lutz, and by Theobald from Lutz material, did not

have a locality specified in the original description and were reported from one or

more states in Brazil. In those instances where labels on extant original material

did not indicate a specific locality we have used the distribution data on Lutz

material in Peryassu (1908:59-75) as an indication of the type locality. All records

from Sao Paulo without a specific locality we have interpreted as being from the

city of Sao Paulo or its vicinity.

Dr. Bertha Lutz, the daughter of Adolpho Lutz, has informed us that the Fazenda

do Bonito, Serra da Bocaina, belonged to her father and uncle and that it was in the

possession of the family before 1900. It is probably therefore that our interpretation

of the place of capture of the lectotype of 256. Aedes (F.) oswaldi is correct.

Macquart, P. Justin M. All 3 species described by Macquart from Brazil, without

locality specified, were probably based on material obtained by C. Gaudichaud from

the “Uranie’”’ world expedition. We are restricting the type locality of these species

to the vicinity of Rio de Janeiro, the only area in Brazil visited by this expedition.

The holotypes of 237. Aedes (O.) albifasciatus and 259, Aedes (S.) toxorhynchus

are in MNHP; the type material of 223. Psorophora (P.) pilipes is non-existent (NE).

Martini, Erich C.W. The holotype of 4. Chagasia stigmopteryx, the only species

described by Martini from Brazil and formerly believed to be non-existent, has been

located in DEI through the courtesy of G. Morge.

Matheson, Robert. The holotype of 318. Culex (Mel.) putumayensis, the only

Brazilian species described by Matheson, is in USNM. The type locality, stated to be

the Amazon River near Peru, is here restricted to Santo Antonio do Ica (Amazonas),

near the mouth of the Putumayo River (Rio Ica).

Neiva, Arthur. Belkin did not find the type material of the 2 species described by

Neiva as sole author that was presumed to be in IOC: 28. Anopheles (A.) tibiamacu-

latus and 168. Sabethes (S.) belisarioi. However, Belkin’s search was not exhaustive

and for the present we consider that the location of these types is unknown (LU).

The types 2 of the 4 species described by Neiva and Pinto are in IOC: 62, Ano-

40 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

pheles (N.) rondoni and 64. Anopheles (N.) cuyabensis. We have designated lecto-

types for both species and restricted the type locality of rondoni. We found no

type material of 63. Anopheles (N.) triannulatus or 215, Uranotaenia noctivaga in

IOC where it should be, but consider its location unknown (LU) pending exhaus-

tive search in this collection.

Neiva was the junior author of 4 other species with Adolpho Lutz (see). He also

proposed the new name adolphoi for 66. Anopheles (K.) lutzii Theobald, 1901

whose lectotype is in BM. -

Neveu-Lemaire, Maurice. Type material is non-existent (NE) of the 2 nominal

species described by Neveu-Lemaire from Brazil (as French Guiana): 248. Aedes

(O.) mathisi and 354. Culex americanus (nomen dubium).

Newstead, Robert. The lectotypes of 219. Orthopodomyia longipalpis and 323.

Culex (Mel.) chrysothorax, described by Newstead and Thomas, are in BM.

Paraense, Wladimir Lobato. See Cerqueira for the | species described by Cer-

queira and Paraense. |

Peryassu, Antonio Goncalves. Apparently all the extant type material of the 16

nominal species described by Peryassu is in IOC or in the Museu Nacional in Quinta

da Boa Vista in Rio de Janeiro. The material in the latter institution was discovered

by Belkin in 1969. Careful study of the material in both institutions is needed be-

fore lectotype designations can be made since several Peryassu species were com-

posite ones. Therefore we have designated lectotypes here only when the specimens

agreed with the current interpretation of Peryassu species. At this time the definite

types of only the following 4 nominal species are recognized, all in IOC: 177. Sa-

bethes (S.) purpureus; 191. Coquillettidia (R.) albicosta; 226. Psorophora (J.) albi-

genu; 291, Culex (Mel.) chrysothorax. Apparently authentic original material of the

following 6 nominal species is either in IOC or MusNac as indicated: 13. Anopheles

(A.) intermedius (IOC); 25. Anopheles (A.) pseudomaculipes (IOC); 29. Anopheles

(N.) allopha (MusNac); 58. Anopheles (N.) oswaldoi (MusNac); 85. Toxorhynchites

(L.) fluminensis (IOC); 194. Coquillettidia (R.) chrysonotum (IOC and MusNac).

No authentic original material of following 6 nominal species was found in either

institution but since an exhaustive search was not made it is possible that some is

extant and therefore the location is recorded as unknown (LU) for the present:

3. Chagasia maculata; 18. Anopheles (A.) rockefelleri; 23. Anopheles (A.) alagoanii;

69. Anopheles (L.) gilesi; 178. Sabethes (S.) quasicyaneus; 216. Uranotaenia argen-

teopennis.

More or less specific localities were mentioned by Peryassu in the original descrip-

tions. We have restricted the type locality of 69. Anopheles (L.) gilesi to the vicinity

of Lassance, the area in the valley of the Rio das Velhas where C. Chagas collected

anophelines in 1907 (Root, 1926:704). The type locality of 18. Anopheles (A.)

rockefelleri was not specified; we have restricted it to the vicinity of the city of Rio

de Janeiro, where Peryassu was working at the time of the description of this spe-

cies.

Peryassu’s major work on the Culicidae of Brazil (1908) is an important source

of information on the species described from Brazil prior to its date of publication,

particularly the section dealing with the distribution of the species (p. 59-75). Ap-

parently all the material then in IOC (at that time, Instituto de Manguinhos) was

examined and recorded by Peryassu, including the species described by Adolpho

Lutz, Carlos Chagas, Oswaldo Cruz and Arthur Neiva.

Belkin et al: Topotypic Brazilian Mosquitoes 41

Pessoa, Samuel Barnsley. The holotype of 350. Culex (Micr.) worontzowi des-

cribed by Pessoa-and Galvao is in FMSP.

Pinto, Cesar. See Neiva for the 4 species described by Neiva and Pinto.

Prado, Alcides. The holotypes of both species described by Prado are in IB: 192.

Coquillettidia (R.) albifera; 211. Uranotaenia ditaenionota.

Rabello, Ernesto Xavier. See Forattini for 1 species described by Forattini, Ra-

bello and Lopes; and Lane for | species described by Lane, Forattini and Rabello.

Ramalho, Gabriel R. See Lane for 1 species described by Lane and Ramalho, and

Correa for 5 species described by Correa and Ramalho.

Ramos, Alberto da Silva. The location of the type material is unknown (LU) for

the 1 species described by Ramos as sole author: 36. Anopheles (N.) guarujaensis.

See Correa for 1 species described by Correa and Ramos; and Fonseca for 2 spe-

cies described by Fonseca and Ramos.

Robineau-Desvoidy, Andre Jean-Baptiste. All the original culicid material of Robi-

neau-Desvoidy is non-existent (NE; see Belkin, 1968:52). The 4 species described

from Brazil did not have a locality specified, we are restricting it to the vicinity of.

Rio de Janeiro (Guanabara) for all these: 37. Anopheles (N.) argyritarsis; 171. Sa-

bethes (S.) locuples; 222. Psorophora (P.) tibialis; 366. Culex pallipes. |

Rondani, Camillo. The location of the type material is unknown (LU) for the 2

species described by Rondani from Brazil, without specified locality. We have re-

stricted the type locality for both species to the vicinity of Rio de Janeiro (Guana-

bara): 87. Trichoprosopon (T.) digitatum, 246. Aedes (O.) scapularis.

Root, Francis Metcalf. Holotypes or lectotypes of all 24 nominal species des-

cribed by Root from Brazil are in USNM: 12. Anopheles (A.) fluminensis; 41. Ano-

pheles (N.) darlingi; 43. Anopheles (N.) strodei; 131. Wyeomyia (D.) melanoides;

149. Phoniomyia incaudata; 154. Phoniomyia pilicauda; 266. Culex (C.) acharistus;

275. Culex (C.) lygrus; 284. Culex (Mel.) andricus; 294. Culex (Mel.) exedrus; 295.

Culex (Mel.) dyius; 298. Culex (Mel.) evansae; 305. Culex (Mel.) cenus; 315. Culex

(Mel.) automartus; 316. Culex (Mel.) oedipus; 317. Culex (Mel.) plectoporpe; 320.

Culex.(Mel.) serratimarge; 326. Culex (Mochl.) megapus; 331. Culex (Micr.) aphy-

lactus; 334. Culex (Micr.) trychnus; 337. Culex (Micr.) gairus; 338. Culex (Micr.)

hedys; 345. Culex (Micr.) microphyllus; 351, Culex (Aed.) accelerans.

All these species are based on material collected in the vicinity of the city Rio de

Janeiro and in the states of Rio de Janeiro and Minas Gerais from April to June

1925, primarily by Root himself.

Rozeboom, Lloyd E. The holotype of the 1 species described from Brazil by

Rozeboom, with Gabaldon as junior author, is in USNM: 56. Anopheles (N.) goeldii.

Sampaio, M.M. See Causey.

Santos, O.B. dos. The type material of 40/. Dixella limai, stated to be in Univ.

Minais Gerais was not examined by us.

Serafim, Jose Jr. The holotype of 241. Aedes (O.) jacobinae, described by Sera-

fim and Davis, is in USNM.

Shannon, Raymond C. The holotypes of all 3 Brazilian species described by

Shannon as sole author are in USNM: 7. Anopheles (S.) lewisi (thomasi, new name).

42 Contrib. Amer. Ent. Inst., vol. 7 no. 5, 1971

52. Anopheles (N.) guarani; 198. Coquillettidia (R.) lynchi.

The holotype of 231. Psorophora (J.) lanei, described by Shannon and Cerqueira,

was not found in IOC and its location is unknown (LU).

See Dyar for 4 species described by Dyar and Shannon.

Stone, Alan. The holotypes of the 2 Brazilian species described by Stone are in

USNM: 89. Trichoprosopon (T.) townsendi; 99. Trichoprosopon (R.) cerqueirai.

Theobald, Frederick V. Holotypes or lectotypes of 42 of the 44 nominal species

described by Theobald from Brazil are extant and all are in the BM collection: 14.

Anopheles (A.) maculipes; 17. Anopheles (A.) mediopunctatus; 51, Anopheles (N.)

niger; 66. Anopheles (K.) lutzii; 72. Toxorhynchites (A.) purpureus,; 80. Toxor-

hynchites (L.) chrysocephalus; 94. Trichoprosopon (S.) fluviatile; 103. Trichopro-

sopon (R.) lunatum; 130. Wyeomyia (D.) luteoventralis; 150. Phoniomyia longi-

rostris; 152, Phoniomyia pallidoventer; 155. Phoniomyia quasilongirostris; 159. Lim-

atus durhamii; 161. Limatus paraensis; 163. Sabethes (S.) albiprivus; 170. Sabethes

(S.) nitidus; 176. Sabethes (S.). purpureus; 181. Sabethes (Sabethoides) confusus;

188. Sabethes (Sabethinus) albiprivatus; 193. Coquillettidia (R.) arribalzagae; 201.

Mansonia (M.) amazonensis; 206. Mansonia (M.) pseudotitillans; 213. Uranotaenia

geometrica; 217. Uranotaenia pallidoventer; 232. Psorophora (J.) lutzii; 233. Psoro-

phora (G.) apicalis (neoapicalis, new name); 238. Aedes (O.) crinifer; 247. Aedes

(O.) serratus; 251, Aedes (F.) mediomaculatus; 252. Aedes (F.) tripunctatus; 270.

Culex (C.) corniger; 272. Culex (C.) bilineatus; 302. Culex (Mel.) humilis; 304.

Culex (C.) indecorabilis; 310. Culex (Mel.) nigrescens; 312. Culex (Mel.) nigricorpus;

314. Culex (Mel.) ocellatus; 339. Culex (Micr.) imitator; 340. Culex (Micr.) argen-

teoumbrosus; 347. Culex (Micr.) Oe SE a) 359, Culex (Anoed.) luteopleurus;

394. Sayomyia braziliensis.

The holotype of 175, Sabethes (S.) lutzii may be in the IOC collection. The type

material of 282. Culex (C.) virgultus, originally in BM, has not been located ae

see Stone, 1957:341-342; Belkin, 1968:21).

There has been some asntiian regarding several nominal species described as new

by Theobald (1907) with supposed Lutz manuscript names. In the Addenda (p.

[630]) in the same work Theobald gives references to Lutz’s descriptions (1905,

incorrectly as 1906) of these species as well as to his own. Therefore we consider

that Theobald did not propose new nominal species and merely redescribed Lutz’s

species, in spite of the fact that later (Theobald, 1910) he attributed 2 of these spe-

cies to himself and the remainder to Lutz in Theobald, 1907. Stone, Knight and

Starcke (1959) did not consider the Theobald descriptions as valid proposals of new

taxa except for Sabethinus aurescens. Belkin (1968:30) followed the above authors

and listed aurescens Theobald, 1907 among the validly proposed nominal taxa.

We see no reason for this exception and therefore are eliminating this name also.

The majority of the Brazilian species described by Theobald were based on mat-

erial sent by Adolpho Lutz. In instances where the localities were not specified for

this material we have followed the same practice in restricting them as with species

described by Adolpho Lutz himself (see). We have also restricted type localities

for some species based on material from other sources as follows. E.E. Austen col-

lected 201. Mansonia (M.) amazonensis aboard a ship on the Amazon; the locality

is restricted here to one of the ports mentioned, Gurupa. The type material of 72.

Toxorhynchites (A.) purpureus was collected by H.W. Bates in the Amazon region;

its locality is restricted to Manaus. H.E. Durham (1902) collected all his material

in Para, the contemporary name for the city of Belem; accordingly the type lo-

Belkin et al: Topotypic Brazilian Mosquitoes 43

cality of all the 9 species described by Theobald from Durham material is restricted

to this city. The same restriction is made for the type locality of species based on

E.A. Goeldi material stated to be from: Para.

Thomas, Harold Wolferstan. See Newstead for the 2 Brazilian nominal species

described by Newstead and Thomas.

Toda, Amazonia. See Forattini for 1 species described by Forattini and Toda.

Trapido, Harold. See Galindo for the 1 species described from Brazil by Galindo,

Carpenter and Trapido.

Unti, Ovidio. Belkin did not find type material of the 5 nominal species or sub-

species described by Unti in any of the Brazilian institutions and therefore we con-

sider that these types are non-existent (NE): 44. Anopheles (N.) ramosi; 45. Ano-

pheles (N.) arthuri; 46. Anopheles (N.) artigasi; 47. Anopheles (N.) albertoi; 55.

Anopheles (N.) ayrozai.

Walker, Francis. The holotypes or lectotypes of all but 1 of the 7 species des-

cribed from Brazil or South America without locality specified are in BM: 207.

Mansonia (M.) titillans; 224. Psorophora (P.) scintillans; 228. Psorophora (J.) dis-

crucians; 240. Aedes (O.) flavicosta; 255. Aedes (F.) terrens; 260. Aedes (S.) exagi-

tans. The holotype of 235. Psorophora perterrens is non-existent (NE).

Since the locality of Para [Belem] was given for 2 of the above species, we have

restricted the type locality to the vicinity of Belem for another species, fitillans,

which was described from Brazil. For flavicosta, stated to be from the Amazon Re-

gion, we have designated Manaus as the type locality. We have restricted the type

locality to the vicinity of the city of Rio de Janeiro for 3 species, discrucians, per-

terrens and terrens, which were stated to be from South America.

Whitman, Loring. See Lane for 11 species described by Lane and Whitman.

Wiedemann, Christian R.W. Type material of all 6 species described by Wiede-

mann from Brazil is in existence: in ZMC, 172. Sabethes (S.) remipes (holotype);

in SNG, 73. Toxorhynchites (A.) trichopygus (syntypes), 239. Aedes (O.) fulvus

(holotype); in NMW, SJ. Toxorhynchites (L.) ferox (syntypes, also in SNG), 83.

Toxorhynchites (L.) violaceus (lectotype), 279. Culex (C.) aestuans.

In only one instance, violaceus, was a specific locality (Bahia) indicated, how-

ever we have restricted the type localities of all the other species to the vicinity

of Salvador (Bahia), the current name of the same city.

DEPOSITORIES

A (Author’s collection) - Private collections; see Duret.

BM (British Museum (Nat. Hist.), Cromwell Road, London, S.W. 7). The BM col-

lection contains the type of the largest number of topotypic Brazilian species (83):

| species described by Christophers; 3 by Edwards, 7 by Evans, 1 by Giles, 6 by

Gordon & Evans, 2 by Lane, 13 by Lutz, 2 by Newstead & Thomas, 42 by Theo-

bald and 6 by Walker. Except as noted all of these are represented by holotypes or

previously designated lectotypes: 14. Anopheles (A.) maculipes; 16. Anopheles (A.)

amazonicus (type locality restricted); 17. Anopheles (A.) mediopunctatus; 51, Ano-

pheles (N.) niger; 66. Anopheles (K.) lutzii (cruzii and adolphoi, new names); 72.

44 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

Toxorhynchites (A.) purpureus (type locality restricted); 76. Toxorhynchites (L.)

horei; 79. Toxorhynchites (L.) solstitialis (new lectotype); 80. Toxorhynchites (L.)

chrysocephalus; 91. Trichoprosopon (T.) pallidiventer (new lectotype); 94. Tricho-

prosopon (S.) fluviatile; 103. Trichoprosopon (R.) lunatum; 112. Wyeomyia (W.)

medioalbipes (new lectotype); 113. Wyeomyia (W.) oblita (syntype; also IOC), 130.

Wyeomyia (D.) luteoventralis; 133. Wyeomyia (D.) negrensis; 134. Wyeomyia (D.)

personata (new lectotype); 150. Phoniomyia longirostris; 152. Phoniomyia palli-

doventer; 155. Phoniomyia quasilongirostris; 159. Limatus durhamii; 161. Limatus

paraensis; 163. Sabethes (S.) albiprivus; 164. Sabethes (S.) albiprivatus (new lecto-

type); 165. Sabethes (S.) amazonicus; 166. Sabethes (S.) longfieldae; 169. Sabethes

(S.) argyronotum; 170. Sabethes (S.) nitidus; 176. Sabethes (S.) purpureus; 181.

Sabethes (Sabethoides) confusus; 184. Sabethes (Sabethinus) aurescens (new lecto-

type); 187. Sabethes (Sabethinus) intermedius (new lectotype); 188. Sabethes (Sabe-

thinus) albiprivatus (melanonymphe, new name); 193. Coquillettidia (R.) arribal-

zagae; 201. Mansonia (M.) amazonensis (type locality restricted); 206. Mansonia

(M.) pseudotitillans; 207. Mansonia (M.) titillans (type locality restricted); 209.

Uranotaenia albitarsis; 213. Uranotaenia geometrica; 217. Uranotaenia pallidoven-

ter; 219. Orthopodomyia longipalpis; 224. Psorophora (P.) scintillans; 228. Psoro-

Phora (J.) discrucians (type locality restricted); 229. Psorophora (J.) arribalzagae;

232. Psorophora (J.) lutzii; 233. Psorophora (G.) apicalis (neoapicalis, new name);

238, Aedes (O.) crinifer; 240. Aedes (O.) flavicosta (type locality restricted); 247.

Aedes (O.) serratus; 249. Aedes (F.) braziliensis; 250. Aedes (F.) fluviatilis (new

lectotype); 251. Aedes (F.) mediomaculatus; 252. Aedes (F.) tripunctatus; 253.

Aedes (F.) leucomelas (leucocelaenus, new name; new lectotype); 255. Aedes (F.)

terrens (type locality restricted); 256. Aedes (F.) oswaldi (new lectotype); 260.

Aedes (S.) exagitans; 270. Culex (C.) corniger; 272. Culex (C.) bilineatus (type

locality restricted); 281. Culex (C.) spinosus (new lectotype); 283. Culex (Mel)

gordoni; 287. Culex (Mel.) innominatus (type locality restricted); 296. Culex (Mel)

manaosensis; 302. Culex (Mel.) humilis; 304. Culex (Mel.) indecorabilis; 310. Cu-

lex (Mel.) nigrescens; 311. Culex (Mel.) clarki (type locality restricted); 312. Culex

(Mel.) nigricorpus; 314. Culex (Mel.) ocellatus; 322. Culex (Mel) theobaldi (new

lectotype); 323. Culex (Mel.) chrysothorax; 324. Culex (Mel.) thomasi; 328. Culex

(Mochl.) innovator (type locality restricted); 339. Culex (Micr.) imitator; 340. Cu-

lex (Micr.) argenteoumbrosus,; 347. Culex (Micr.) pleuristriatus; 353. Culex (Aed.)

hildebrandi (type locality restricted); 359. Culex (Anoed.) luteopleurus; 360. Cu-

lex (Anoed.) originator; 375. Corethrella kummi (type locality restricted); 380.

Corethrella tarsata; 394. Sayomyia braziliensis (type locality restricted); 397. Ed-

wardsops brevisector. :

The only type material of Brazilian mosquitoes that is missing in the BM col-

lection is for 235. Psorophora perterrens and 282. Culex (C.) virgultus.

Casal (Osvaldo H. Casal, Departamento de Entomologia Sanitaria, Instituto Na-

cional de Microbiologia, Avda. Velez Sarsfield 563, Buenos Aires). The only type

material in the private collection of Casal is that of 355. Culex (Aed.) clastrieri.

DEI (Deutsches Entomologisches Institut; now Institut fur Pflanzenschutzfors-

chung, Zweigstelle Eberswalde, Abteilung Taxonomie der Insekten, Schicklerstrasse

5, 13 Eberswalde, DDR). The holotype of 4. Chagasia stigmopteryx Martini, 1932,

formerly believed to be non-existent, has been located in DEI by G. Morge.

Duret (Jose P. Duret, Venezuela 2130, Buenos Aires). The holotypes of 15 spe-

cies described by Duret and 2 species described by Duret and Damasceno are in the

Belkin et al: Topotypic Brazilian Mosquitoes 45

private collection of Duret: 278. Culex (C.) paramaxi; 286. Culex (Mel.) bahiensis;

290. Culex (Mel.) mojuensis; 292. Culex (Mel.) contei; 293. Culex (Mel.) cristovai;,

297. Culex (Mel.) ernanii; 300. Culex (Mel.) faurani; 301, Culex (Mel.) flochi; 306.

Culex (Mel.) isabelae; 307. Culex (Mel.) johnnyi; 308. Culex (Mel.) kerri; 319. Cu-

lex (Mel) rachoui; 321. Culex (Mel) silvai; 327. Culex (Mochl.) galvaoi; 329. Cu-

lex (Mochl.) palaciosi; 356. Culex (Anoed.) belemensis; 358. Culex (Anoed.) damas-

cenoi.

FH (Faculdade de Higiene e Saude Publica, Av. Dr. Arnaldo, 715, Caixa Postal

8099, Sao Paulo, S.P.). The FH collection is in excellent condition, but the type

material is not segregated. The type material originally in SPM has been transferred

to FH. At present the collection contains type material of 76 species: 2 species

described by Antunes & Lane, 1 by Barretto & Coutinho, 7 by Cerqueira, | by

Cerqueira & Lane, | by Correa, 1 by Correa & Cerqueira, 5 by Correa & Ramalho,

1 by Correa & Ramos, 1 by Coutinho & Forattini, 2 by Fonseca & Ramos, | by

- Forattini, Rabello & Lopes, 1 by Forattini & Toda, | by Galindo, Carpenter & Tra-

pido, 2 by Galvao, 1 by Galvao & Lane, 38 by Lane, 1 by Lane & Antunes, 2 by

Lane & Cerqueira, 2 by Lane & Coutinho, | by Lane & Forattini, 1 by Lane, Forat-

tini & Rabello, 1 by Lane & Guimaraes, | by Lane & Ramalho, and 1 by Lutz. Ex-

cept as noted, all the species are represented by holotypes or previously designated

lectotypes: 8. Anopheles (A.) anchietai; 9. Anopheles (A.) bustamentei; 19. Ano-

pheles (A,) limai (costalimai, new name) 20. Anopheles (A.) costai; 26. Anopheles

(A.) rachoui (new lectotype); 32. Anopheles (N.) imperfectus; 54. Anopheles (N.)

noroestensis (new lectotype; possibly partly in FMSP),; 67. Anopheles (K.) monte-

mor; 68. Anopheles (K.) laneanus; 96. Trichoprosopon (I.) luederwaldti; 108. Wyeo-

myia (W.) downsi; 117. Wyeomyia (C.) dyari; 142. Phoniomyia antunesi (syntypes),

145. Phoniomyia diabolica; 148. Phoniomyia galvaoi; 151. Phoniomyia lopesi; 173.

Sabethes (S.) forattinii; 174. Sabethes (S.) lanei; 179. Sabethes (S.) shannoni; 180.

Sabethes (S.) spixi; 183. Sabethes (Sabethoides) tridentatus; 195, Coquillettidia (R.)

hermanoi; 199. Coquillettidia (R.) neivai; 200. Coquillettidia (R.) shannoni; 205.

Mansonia (M.) pessoai; 210. Uranotaenia davisi; 212. Uranotaenia burkii; 214. Ura-

notaenia mathesoni; 227. Psorophora (J.) amazonica; 243. Aedes (O.) pennai; 254.

Aedes (F.) leucophoebus; 262. Haemagogus (S.) baresi; 265. Culex (C.) abnormalis;

267. Culex (C.) airozai; 268. Culex (C.) brami; 271, Culex (C.) deanei; 273. Culex

(C.) foliaceus; 274. Culex (C.) forattinii; 276. Culex (C.) mauesensis; 280. Culex

(C.) renatoi; 344. Culex (Micr.) lanei; 349. Culex (Micr.) shopei; 361. Culex (Car.)

anduzei; 363. Culex (Car.) iridescens (neotype),; 364. Culex (Car.) soperi; 368. Core-

thrella confusa; 369. Corethrella bromelicola; 370. Corethrella cardosoi; 371. Core-

thrella edwardsi (type locality restricted); 372. Corethrella flavitibia; 373. Core-

thrella fulva; 374. Corethrella infuscata; 376. Corethrella lopesi; 377. Corethrella

pillosa; 378. Corethrella selvicola; 379. Corethrella striata; 381. Corethrella trava-

ssosi; 382. Corethrella vittata; 383. Corethrella whitmani (type locality restricted);

384. Lutzomiops alticola; 386. Lutzomiops barrettoi; 387. Lutzomiops coutinhoi;

388. Lutzomiops niger; 389. Lutzomiops iridescens; 390. Lutzomiops juquianus;

391, Lutzomiops lutzi; 392. Lutzomiops. manaosensis; 393. Lutzomiops nigrescens;

395, Sayomyia antunesi; 396. Sayomyia souzai, 398. Edwardsops magnificus; 399.

Edwardsops unicolor; 400. Dixella chapadensis; 402. Dixella paulistana; 403. Dix-

ella torrentia; 404. Dixella wygodzinskyi (new lectotype).

As noted under Lane in the authors section, we consider. that the specimens in

FH labelled as holotypes of the following species are only paratypes: 143. Phonio-

46 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

myia bonnei; 144. Phoniomyia davisi; 156. Phoniomyia neivai and 157. Phoniomyia

theobaldi. |

Type material of 109. Wyeomyia (W.) leucotarsis, stated to be in FH by Stone,

Knight and Starcke (1959:79), is not in this collection.

FMRP (Departamento de Parasitologia, Faculdade de Medicina, Ribeirao Preto,

S.P.). The holotypes of the 3 species described by Duret and Barretto, 285, Culex

(Mel) aureonotatus, 289. Culex (Mel) bifoliolatus and 325. Culex (Mel.) trilobula-

tus, were not examined by Belkin but J.P. Duret assurred him that they are in

FMRP. . af

FMSP (Departamento de Parasitologia, Faculdade de Medicina, Sao Paulo, S.P.).

It appears that the mosquito collection in FMSP has not been taken care of in sev-

eral years. It is in deplorable condition and unless prompt action is taken the re-

maining type material will be lost. The only type material found by Belkin in 1969

was for the following 7 species: 31. Anopheles (N.) marajoara Galvao & Damasceno

(doubtful syntypes); 34. Anopheles (N.) antunesi Galvao & Amaral (holotype); 40.

Anopheles (N.) pessoai Galvao & Lane (holotype); 48. Anopheles (N.) galvaoi Cau-

sey, Deane & Deane (lectotype by present designation); 49. Anopheles (N.) lanei

Galvao & Amaral (holotype); 70. Anopheles (L.) pseudotibiamaculatus Galvao &

Barretto (holotype); 350. Culex (Micr.) worontzowi Pessoa & Galvao (holotype).

Type material of the following 2 species, originally stated to be in FMSP, was not

found; its location is considered unknown (LU) for the present, although it seems

probable that it has been lost: 60. Anopheles (N.) konderi Galvao & Damasceno;

208. Mansonia (M.) wilsoni Barretto & Coutinho.

A male of 54. Anopheles (N.) noroestensis Galvao & Lane, 1937 in FMSP (343)

may be the specimen from which the genitalia, designated as lectotype of this spe-

cies (in FH), was removed.

IB (Instituto Butantan, Butanta, Sao Paulo, S.P.). The small mosquito collection

in IB is in good condition. The specimens have been renumbered but there is a cross

reference to the original numbers. The few Adolpho Lutz specimens do not include

any type material. The only type material in IB consists of the holotypes of the 2

species described by Prado: 192. Coquillettidia (R.) albifera and 211. Uranotaenia

ditaenionota.

IBSP (Instituto Biologico, Sao Paulo, S.P.). The mosquito collection in the Ani-

mal Parasitology section of the institute is in poor condition; its register has been

lost. Fortunately some type material of the only species originally deposited in this

collection, 269. Culex (C.) carcinoxenus, was found in IOC (see).

IOC (Instituto Oswaldo Cruz, Av. Brasil, Manguinhos, Rio de Janeiro, GB.). The

mosquito collection in IOC is the oldest in Brazil. As indicated in the authors

section under Lima and Lutz, apparently the specimens were at one time pinned in |

small boxes. This arrangement may date back to Lutz or to Peryassu or Neiva. Later

some of the specimens from the old collection were transferred to individual tubes,

probably by Costa Lima. Others are still in the original boxes. Additions to the col-

lection were put in individual tubes except for the types of sabethines described

by Lane and Cerqueira which are in unit trays. We have not been able to find in

IOC some of the types stated to have been deposited in this collection by the orig-

inal authors with indication of collection number. We did not find a register of

the collection numbers in IOC. The only records available to us were cards arranged

in a taxonomic order. It is apparent that IOC has had a very liberal policy of loan-

Belkin et al: Topotypic Brazilian Mosquitoes 47

ing type specimens. In a few instances notations were made on the cards but in

other cases it seems that the cards were either removed from the file or no record

of the loan was entered. We did not locate the collection of the Rockefeller Founda-

tion Laboratory which was originally in IOC.

We found apparently authentic type material of 77 topotypic Brazilian nominal

species in IOC: 1 species described by Castro; 1 by Cerqueira; 3 by Chagas; 3 by

Cruz; 1 by Del Ponte; 3 by Del Ponte and Cerqueira; 31 species by Lane and Cer-

queira; 9 by Lima; 1 by Lima, Guitton and Ferreira; 9 by Lutz; 4 by Lutz and Neiva;

2 by Neiva and Pinto; 8 by Peryassu; 1 by Theobald. Because of insufficient data on

the specimens and need for further study of the material we have refrained from

designating lectotypes or positively identifying the holotypes of several species as

indicated in the following list; the species without notation are represented by

holotypes: 6. Anopheles (S.) lineatus (possibly holotype); 11. Anopheles (A.) evan-

droi; 13. Anopheles (A.) intermedius (possibly syntype),; 15. Anopheles (A.) matto-

grossensis; 21, Anopheles (A.) minor (syntypes); 22. Anopheles (A.) lutzi (pery-

assui, new name; possibly syntypes); 25. Anopheles (A.) pseudomaculipes (possi-

bly syntype); 39. Anopheles (N.) braziliensis (syntypes); 50. Anopheles (N.) lutzii

(syntypes),; 61. Anopheles (N.) parvus (possibly syntypes),; 62. Anopheles (N.) ron-

doni (new lectotype); 64. Anopheles (N.) cuyabensis (new lectotype); 71. Toxor-

hynchites (A.) catharinensis; 75. Toxorhynchites (L.) bambusicola (syntypes); 78.

Toxorhynchites (L.) pusillus (syntypes); 82. Toxorhynchites (L.) posticatus (syn-

types); 85. Toxorhynchites (L.) fluminensis (possibly syntype); 86. Trichoprosopon

(T.) compressum (syntypes); 90. Trichoprosopon (T.) obscurum; 92. Trichoproso-

pon (T.) soaresi; 93. Trichoprosopon (L.) brevipes (new lectotype); 95. Trichopro-

sopon (S.) longipalpis (possibly syntype); 97. Trichoprosopon (C.) walcotti; 98.

Trichoprosopon (R.) castroi; 100. Trichoprosopon (R.) edwardsianum; 101. Tri-

choprosopon (R.) humboldti; 104. Trichoprosopon (R.) reversum; 105. Tricho-

prosopon (R.) simile; 106. Trichoprosopon (R.) theobaldi; 107. Wyeomyia (W.)

arthrostigma (new lectotype),; 110. Wyeomyia (W.) limai; 111. Wyeomyia (W.) lutzi

(syntypes); 113. Wyeomyia (W.) oblita (syntype; also BM); 114. Wyeomyia (W.)

pintoi; 115. Wyeomyia (W.) sabethea; 116. Wyeomyia (W.) serrata (new lectotype),

118. Wyeomyia (C.) kummi; 121. Wyeomyia (A.) rooti (alani, new name); 122.

Wyeomyia (D.) airosai; 124. Wyeomyia (D.) cesari; 126. Wyeomyia (D.) finlayi;

127. Wyeomyia (D.) howardi; 128. Wyeomyia (D.) kerri; 129. Wyeomyia (D.)

knabi; 136. Wyeomyia (D.) rooti (new lectotype); 137. Wyeomyia (D.) delpontei;

138. Wyeomyia (D.) shannoni; 140. Wyeomyia (D.) tarsata; 141, Wyeomyia (D.)

undulata; 143. Phoniomyia bonnei; 144. Phoniomyia davisi; 147. Phoniomyia fla-

bellata; 153. Phoniomyia palmata; 156. Phoniomyia neivai; 157. Phoniomyia theo-

baldi; 167. Sabethes (S.) batesi; 175. Sabethes (S.) lutzii (possibly holotype); 177.

Sabethes (S.) purpureus (remipusculus, new name; new lectotype); 185. Sabethes

(Sabethinus) fabricii; 186. Sabethes (Sabethinus) lutzianus; 189. Sabethes (Sabe-

thinus) soperi; 190. Sabethes (Sabethinus) whitmani; 191. Coquillettidia (R.) albi-

costa (new lectotype); 194. Coquillettidia (R.) chrysonotum (syntypes, also in Mus

Nac); 196. Coquillettidia (R.) juxtamansonia (new lectotype),; 203. Mansonia (M.)

chagasi; 218. Orthopodomyia albicosta (new lectotype); 220. Orthopodomyia town-

sendi; 221. Orthopodomyia sampaioi; 225. Psorophora (P.) genumaculata (new

lectotype); 226. Psorophora (J.) albigenu; 230. Psorophora (J.) forceps; 234. Psoro-

phora (G.) scutipunctata (possibly syntypes); 269. Culex (C.) carcinoxenus (new

lectotype); 291. Culex (Mel.) chrysothorax (new lectotype); 299. Culex (Mel.) fas-

ciolatus (possibly syntypes); 309. Culex (Mel.) lugens (syntypes).

48 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

Type material of the following species, stated to be in IOC, was not located in this

collection; it was probably borrowed and not returned or misplaced: 1. Chagasia

fajardi; 28. Anopheles (A.) tibiamaculatus; 63. Anopheles (N.) triannulatus; 119.

Wyeomyia (C.) mattinglyi; 135. Wyeomyia (D.) brucei; 162. Limatus flavisetosus;

168. Sabethes (S.) belisarioi; 215. Uranotaenia noctivaga; 236. Aedes (O.) aenig-

maticus; 242. Aedes (O.) lepidus; 244. Aedes (O.) perventor; 245. Aedes (O.) rhya-

cophilus; 261. Haemagogus (L.) tropicalis; 352. Culex (Aed.) amazonensis.

For other material possibly in IOC see LU under the following senior authors:

Chagas; Castro; Cerqueira; Cruz; Del Ponte; Lane; Lima; Lutz; Neiva; Peryassu;

Shannon. |

LU (Location unknown). Listed here are 49 nominal species whose types have

not been located but may be extant. They are arranged according to senior authors

with indication of depositories where additional search should be made before their

type material can be declared to be non-existent.

Barretto & Coutinho; possibly in FMSP: 202. Mansonia (M.) cerqueirai; 208.

Mansonia (M.) wilsoni.

Castro; should be in IOC: 162. Limatus flavisetosus.

Cerqueira & Antunes; Cerqueira & Costa; Cerqueira & Paraense; should be in

IOC: 236. Aedes (O.) aenigmaticus; 242. Aedes (O.) lepidus; 244. Aedes (O.) per-

ventor; 261. Haemagogus (L.) tropicalis.

Chagas; should be in IOC: 53. Anopheles (N.) nigritarsis.

Cruz; should be in IOC: 2. Chagasia neivae.

Del Ponte; Delponte & Cerqueira; should be in IOC or FH: 135. Wyeomyia

(D.) brucei; 139. Wyeomyia (D.) subcomplosa.

Dyar & Knab; possibly in NMW: 84. Toxorhynchites (L.) ambiguus.

Galvao; Galvao & Damasceno; Galvao, Lane & Correa; should be in FMSP: 33.

Anopheles (N.) domesticus; 42. Anopheles (N.) paulistensis; 60. Anopheles (N.)

konderi; 65, Anopheles (N.) chagasi.

Kumm; possibly in IOC or USNM: 335. Culex (Micr.) davisi.

Lane; should be in IOC or FH: 109. Wyeomyia (W.) leucotarsis; 119. Wyeomyia

(C.) mattinglyi. | $ |

Lane & Whitman; originally in Rockefeller Foundation Laboratory at IOC: 313.

Culex (Mel.) nigrimacula; 332, Culex (Micr.) aureus; 333. Culex (Micr.) carioca;

336. Culex (Micr.) dubitans; 341. Culex (Micr.) retrosus; 342. Culex (Micr.) fusca-

tus; 343. Culex (Micr.) intermedius; 348. Culex (Micr.) reducens; 357. Culex

(Anoed.) canaanensis; 362. Culex (Car.) antunesi; 365. Culex (Car.) wilsoni. 3

Lima; should be in IOC: 245, Aedes (O.) rhyacophilus.

Lutz; should be in IOC: J. Chagasia fajardi; 263. Haemagogus (S.) capricornii

(neotype); 352. Culex (Aed.) amazonensis.

Neiva; Neiva & Pinto; should be in IOC: 28. Anopheles (A.) tibiamaculatus; 63.

Anopheles (N.) triannulatus; 168. Sabethes (S.) belisarioi; 215. Uranotaenia noc-

tivaga.

Peryassu; should be in IOC or MusNac: 3. Chagasia maculata; 18. Anopheles

(A.) rockefelleri; 23. Anopheles (A.) alagoanii; 69. Anopheles (L.) gilesi; 178. Sa-

bethes (S.) quasicyaneus; 216. Uranotaenia argenteopennis.

Ramos; possibly in SPM: 36. Anopheles (N.) guarujaensis.

Rondani; possibly in Bologna (see Belkin, 1968:53): 87. Trichoprosopon (T.)

digitatum,; 246. Aedes (O.) scapularis.

Shannon & Cerqueira; should be in IOC: 231. Psorophora (J.) lanei.

Belkin et al: Topotypic Brazilian Mosquitoes : 49

Theobald; should be in BM: 282. Culex (C) virgultus.

MNHP (Museum National d’Histoire Naturelle, Laboratoire d’Entomologie Gen-

Macquart are in MNHP: 237. Aedes (O.) albifasciatus and 259. Aedes (S.) toxo-

rhynchus.

Museu Nacional, Rio de Janeiro (Quinta da Boa Vista, Rio de Janeiro, GB.).

Syntypes of 3 Peryassu species were found in this collection: 29. Anopheles (N.)

allopha; 58. Anopheles (N.) oswaldoi; 194. Coquillettidia (R.) chrysonotum. For

additional species possibly to be found in this collection see LU under Peryassu.

NE (Non-existent). Type material of the following 27 species, listed by authors,

has not been located and it is reasonably certain that it is non-existent. 3

Bonne-Wepster & Bonne, originally in USNM: 158. Phoniomyia tripartita,

Bourroul: 77. Toxorhynchites (L.) mariae.

Causey, Deane & Deane: 5. Chagasia rozeboomi.

Correa: 10. Anopheles (A.) geometricus.

Galvao & Lane; originally in FMSP: 30. Anopheles (N.) limai.

Kollar: 36 7. Culex molestus.

Laveran: 160. Limatus curvirostris. !

Lutz; originally probably in IOC: 74. Toxorhynchites (A.) neglectus; 88. Tri-

choprosopon (T.) splendens; 120. Wyeomyia (M.) leucostigma; 123. Wyeomyia (D.)

bourrouli; 125. Wyeomyia (D.) confusa; 258. Aedes (H.) fulvithorax; 330. Culex

(Micr.) albipes.

Macquart; originally in MNHP: 223. Psorophora (P.) pilipes.

Neveu-Lemaire; originally in FMP: 248. Aedes (O.) mathisi; 354. Culex ameri-

canus.

Robineau-Desvoidy; originally some in MNHP: 37. Anopheles (N.) argyritarsis;

171. Sabethes (S.) locuples; 222. Psorophora (P.) tibialis; 366. Culex pallipes.

Unti: 44. Anopheles (N.) ramosi; 45. Anopheles (N.) arthuri; 46. Anopheles (N.)

artigasi; 47, Anopheles (N.) albertoi; 55. Anopheles (N.) ayrozai.

Walker; originally in BM: 235. Psorophora perterrens.

NMW (Naturhistorisches Museum, Burgring 7, Wien 1). Type material of only 3

Brazilian species is in NMW: 81. Toxorhynchites (L.) ferox (wiedemanni, new name;

syntypes; also in SNG); 83. Toxorhynchites (L.) violaceus (lectotype); 279. Culex

(C.) aestuans (lectotype); all described by Wiedemann.

SNG (Natur-Museum and Forschungs-Institut Senckenberg, Senckenberg-Anlage

25, 6 Frankfurt 1). Type material of only 3 Brazilian species is in SNG: 73. Toxo-

rhynchites (A.) trichopygus (syntypes); 81. Toxorhynchites (L.) ferox (wiedemanni,

new name; syntypes; also in NMW), 239. Aedes (O.) fulvus; all described by Wiede-

mann. |

SPM (Servico de Profilaxia da Malaria, Sao Paulo, S.P.). All the type material for-

merly in the SPM collection has been transferred to FH. See also LU under Ramos.

Universidade do Minas Gerais (Belo Horizonte). The exact location of the type

of 401. Dixella limai, described by Santos, is unknown.

USNM (U.S. National Museum of Natural History, Wash., D.C. 20560). Holo-

types or lectotypes of 51 topotypic Brazilian nominal species are in USNM: 1 spe-

cies described by Berlin; 1 by Causey; 1 by Causey, Deane, Deane & Sampaio;

50 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

1 by Davis; 5 by Dyar; 4 by Dyar & Shannon; | by Forattini; 1 by Galvao & Lane;

1 by Komp; | by Lane; 1 by Lane & Cerqueira; 1 by Lutz; 1 by Matheson; 24 by

Root; 1 by Rozeboom & Gabaldon; 1 by Serafim and Davis; 3 by Shannon; 2 by

Stone. 7. Anopheles (S.) lewisi (thomasi, new name); 12. Anopheles (A.) flumi-

nensis; 24, Anopheles (A.) celidopus; 27. Anopheles (A.) shannoni; 35. Anopheles

(N.) emilianus; 38. Anopheles (N.) sawyeri; 41. Anopheles (N.) darlingi; 43. Ano-

pheles (N.) strodei; 52. Anopheles (N.) guarani; 56. Anopheles (N.) goeldii; 57.

Anopheles (N.) dunhami; 59. Anopheles (N.) metcalfi; 89. Trichoprosopon (T.)

townsendi; 99, Trichoprosopon (R.) cerqueirai; 102. Trichoprosopon (R.) trichopus;

131. Wyeomyia (D.) melanoides; 132. Wyeomyia (D.) mystes; 146. Phoniomyia ed-

wardsi; 149. Phoniomyia incaudata; 154. Phoniomyia pilicauda; 182. Sabethes (Sa-

bethoides) glaucodaemon; 197. Coquillettidia (R.) hypocindyna; 198. Coquillettidia

(R.) lynchi; 204. Mansonia (M.) indubitans; 241. Aedes (O.) jacobinae; 257. Aedes

(H.) aureolineatus; 264. Culex (L.) brasiliae; 266. Culex (C.) acharistus; 275, Culex

(C.) lygrus; 277. Culex (C.) oswaldoi; 284. Culex (Mel.) andricus; 288. Culex (Mel.)

bequaerti; 294. Culex (Mel.) exedrus; 295. Culex (Mel.) dyius; 298. Culex (Mel)

-evansae; 303. Culex (Mel.) inadmirabilis; 305. Culex (Mel.) cenus; 315. Culex (Mel.)

automartus; 316, Culex (Mel.) oedipus; 317. Culex (Mel.) plectoporpe; 318. Culex

(Mel.) putumayensis; 320. Culex (Mel.) serratimarge; 326. Culex (Mochl.) megapus;

331. Culex (Micr.) aphylactus; 334. Culex (Micr.) trychnus; 337. Culex (Micr.)

gairus; 338. Culex (Micr.) hedys; 345, Culex (Micr.) microphyllus; 346. Culex

(Micr.) neglectus; 351, Culex (Aed.) accelerans; 385, Lutzomiops amazonicus.

ZMC (Universitetets Zoologiske Museum, Universitetsparken 15, Kobenhavn).

The only Brazilian culicid type material in ZMC is the holotype of Wiedemann’s

172. Sabethes (S.) remipes.

LOCALITIES

ACRE

Feijo: 254. Aedes (F.) leucophoebus.

Rio Branco: 48. Anopheles (N.) galvaoi.

ALAGOAS

Maceio: 23. Anopheles (A.) alagoanii.

AMAPA

Cunani: 248. Aedes (O.) mathisi; 354. Culex americanus.

AMAZONAS

Coari: 60. Anopheles (N.) konderi.

Itacoatiara: 287. Culex (Mel.) innominatus; 312. Culex (Mel.) nigricorpus; 328.

Culex (Mochl.) innovator; 353. Culex (Aed.) hildebrandi.

Manaus and vicinity: 16. Anopheles (A.) amazonicus; 65. Anopheles (N.) chagasi;

72. Toxorhynchites (A.) purpureus; 76. Toxorhynchites (L.) horei (Macapa); 88.

Trichoprosopon (T.) splendens; 133. Wyeomyia (D.) negrensis (Macapa), 165. Sa-

bethes (S.) amazonicus (Macapa); 173. Sabethes (S.) forattinii (km 23, route 17);

174. Sabethes (S.) lanei (km 23, route 17); 175. Sabethes (S.) lutzii; 179. Sabethes

Belkin et al: Topotypic Brazilian Mosquitoes | 51

(S.) shannoni (Igarape do Leao); 180. Sabethes (S.) spixi (km 23, route 17); 183. Sa-

bethes (Sabethoides) tridentatus (Igarape de Bolivia); 209. Uranotaenia albitarsis

(Macapa), 219. Orthopodomyia longipalpis; 227. Psorophora (J.) amazonica (Iga-

rape do Taruma),;240. Aedes (O.) flavicosta; 249, Aedes (F.) braziliensis (Macapa);

262. Haemagogus (S.) baresi (Igarape do Taruma); 283. Culex (Mel.) gordoni (Bos-

que); 296. Culex (Mel.) manaosensis; 300. Culex (Mel.) faurani; 311, Culex (Mel.)

clarki; 323. Culex (Mel) chrysothorax (inner Flores swamp, Pensador); 324. Culex

(Mel.) thomasi; 352. Culex (Aed.) amazonensis; 360. Culex (Anoed.) originator

(Macapa), 392. Lutzomiops manaosensis; 397. Edwardsops brevisector.

Maues: 276. Culex (C.) mauesensis.

Rio Maues: 361. Culex (Car.) anduzéi.

Rio Parauari: 267. Culex (C.) airozai.

Rio Preto, municipio Joao Goulard: 307. Culex (Mel.) johnnyi; 308. Culex (Mel.)

kerri; 358. Culex (Anoed.) damascenoi. ;

Santo Antonio do Ica: 318. Culex (Mel.) putumayensis.

Tefe: 57. Anopheles (N.) dunhami; 102. Trichoprosopon (T.) trichopus.

BAHIA

Camacari: 380. Corethrella tarsata.

Caravelas: 97. Trichoprosopon (Ct.) walcotti.

Gois Calmon (Muriqueira): 127. Wyeomyia (D.) howardi; 147. Phoniomyia fla-

bellata.

Itaparica, Ilha de: 77. Toxorhynchites (L.) mariae; 330. Culex (Micr.) albipes.

Jacobina: 241. Aedes (O.) jacobinae.

Piraja: 189. Sabethes (Sabethinus) soperi; 257. Aedes (H.) aureolineatus.

Salvador (Bahia) and vicinity: 7. Anopheles (S.) lewisi; 73. Toxorhynchites (A.)

trichopygus; 81, Toxorhynchites (L.) ferox; 83. Toxorhynchites (L.) violaceus; 84.

Toxorhynchites (L.) ambiguus; 112. Wyeomyia (W.) medioalbipes; 119. Wyeomyia

(C.) mattinglyi; 139. Wyeomyia (D.) subcomplosa [see also Cuiaba (Mato Grosso);

Belem and Curralinho (Para); Sergipe]; 172. Sabethes (S.) remipes; 210. Uranotae-

nia davisi; 239. Aedes (O.) fulvus; 279. Culex (C.) aestuans; 335. Culex (Micr.)

davisi; 375. Corethrella kummi.

Santo Amaro (municipio), Acude Sao Bento: 26. Anopheles (A.) rachoui.

Urucuca: 286. Culex (Mel.) bahiensis.

CEARA

Sao Benedito: 5. Chagasia rozeboomi; 38. Anopheles (N.) sawyeri.

ESPIRITO SANTO

Santa Teresa: 122. Wyeomyia (D.) airosai.

Sao Joao de Petropolis (Vale do Canaa): 92. Trichoprosopon (T.) soaresi; 190,

Sabethes (Sabethinus) whitmani; 245, Aedes (O.) rhyacophilus; 357. Culex (Anoed.)

canaanensis; 362. Culex (Car.) antunesi; 365. Culex (Car.) wilsoni; 383. Corethrella

whitmani.

Vale do Rio Doce (see also Baixada Fluminense, state of Rio de Janeiro): 58.

Anopheles (N.) oswaldoi; 194. Coquillettidia (R.) chrysonotum.

GOIAS

Anapolis: 140. Wyeomyia (D.) tarsata.

52 : ~ Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

Inhumas: 368. Corethrella confusa.

Ponte Ipe-Arcado: 258. Aedes (H.) fulvithorax.

GUANABARA

Rio de Janeiro and vicinity: 13. Anopheles (A.) intermedius (see also Xerem,

state of Rio de Janeiro); 18. Anopheles (A.) rockefelleri; 29. Anopheles (N.) aie

pha (see also state of Rio de Janeiro, Baixada Fluminense); 37. Anopheles (N.)

argyritarsis; 50. Anopheles (N.) lutzii (asoa Rodrigo de Freitas); 66. Anopheles

(K.) lutzii; 78. Toxorhynchites (L.) pusillus (Alto da Boa Vista); 85. Toxorhyn-

chites (L.) fluminensis; 87. Trichoprosopon (T.) digitatum; 93. Trichoprosopon (L.)

brevipes (Laranjeiras); 99. Trichoprosopon (R.) cerqueirai; 101, Trichoprosopon

(R.) humboldti (Paineiras); 103. Trichoprosopon (R.) lunatum (Maua); 111. Wyeo-

myia (W.) lutzi (Alto da Boa Vista); 121. Wyeomyia (A.) rooti; 132. Wyeomyia (D.)

mystes; 143. Phoniomyia bonnei; 149. Phoniomyia incaudata; 150.. Phoniomyia

-longirostris; 152. Phoniomyia pallidoventer; 153. Phoniomyia palmata, 155, Phonio-

myia quasilongirostris (Maua) ; 157. Phoniomyia theobaldi; 160. Limatus curviro-

stris; 171. Sabethes (S.) locuples; 176. Sabethes (S.) purpureus; 215, Uranotaenia

noctivaga (Gavea); 216. Uranotaenia argenteopennis (see also Baixada Fluminense,

state of Rio de Janeiro); 221. Orthopodomyia sampaioi (Tijuca); 222. Psorophora

(P.) tibialis; 223. Psorophora (P.) pilipes; 228. Psorophora (J.), discrucians; 230.

Psorophora (J.) forceps (Jacarepagua); 232. Psorophora (J.) lutzii (Quinta da Boa

Vista); 233. Psorophora (G.) apicalis; 235. Psorophora perterrens; 237. Aedes (O.)

albifasciatus; 246. Aedes (O.) scapularis; 247. Aedes (O.) serratus (Quinta da Boa

Vista); 255. Aedes (F.) terrens; 259. Aedes (S.) toxorhynchus; 282. Culex (C.) vir-

gultus (Quinta da Boa Vista); 291. Culex (Mel.) chrysothorax (Copacabana); 313.

Culex (Mel.) nigrimacula; 315. Culex (Mel.) automartus (Jardim Botanico); 31 7.

Culex (Mel) plectoporpe (Bangu); 332. Culex (Micr.) aureus; 333. Culex (Micr.)

carioca; 336. Culex (Micr.) dubitans; 337. Culex (Micr.) gairus (Jardim Botanico);

340. Culex (Micr.) argenteoumbrosus; 341, Culex (Micr.) retrosus; 342. Culex (Micr.)

fuscatus; 343. Culex (Micr.) intermedius; 348. Culex (Micr.) reducens; 366. Culex

pallipes; 367. Culex molestus.

MATO GROSSO

Acampamento dos Morros Azues: 195. Coquillettidia (R.) hermanoi.

Boa Esperanca (see also Pocinho): 109. Wyeomyia (W.) leucotarsis.

Coronel Ponce: 212. Uranotaenia burkii; 265. Culex (C.) abnormalis.

-~Cuiaba: 124. Wyeomyia (D.) cesari; 128. Wyeomyia (D.) kerri; 135. Wyeomyia

(D.) brucei; 136. Wyeomyia (D.) rooti; 137. Wyeomyia (D.) delpontei; 139. Wye-

omyia (D.) subcomplosa [see also Salvador (Bahia); Belem and Curralinho (Para);

Sergipe]; 141. Wyeomyia (D.) undulata; 200. Coquillettidia (R.) shannoni.

Ladario, on Rio Paraguai: 62. Anopheles (N.) rondoni.

Lagoa de Mandiore [as Manicore and Mandicore]: 15, Anopheles (A.) matto-

grossensis.

Maracaju: 202. Mansonia (M.) cerqueirai; 231, Psorophora (J.) lanei; 236. Aedes

(O.) aenigmaticus; 388. Lutzomiops niger.

Melguierra, Ribeirao Amolar, headwaters of Rio Paraguai south of Diamantino:

166. Sabethes (S.) longfieldae; 169. Sabethes (S.) argyronotum.

Pocinho (see also Boa Esperanca): 109. Wyeomyia (W.) leucotarsis.

Portinha, Chapada region: 400. Dixella chapadensis.

Salobra, on Rio Miranda: 371. Corethrella edwardsi; 381. eavenieela travassoSsi;

Belkin et al: Topotypic Brazilian Mosquitoes 53

398. Edwardsops magnificus; 399. Edwardsops unicolor. |

Sao Joao (Fazenda), right bank of Rio Cuiaba: 63. Anopheles (N.) triannulatus,

64, Anopheles (N.) cuyabensis.

MINAS GERAIS

Belo Horizonte: 242. Aedes (O.) lepidus.

- Bicudo(s), Rio: 22. Anopheles (A.) lutzi; 168. Sabethes (S.) belisarioi; 203. Man-

sonia (M.) chagasi. 7

Cambuquira: 3. Chagasia maculata.

Engenheiro Dolabela: 278. Culex (C.) paramaxi.

Juiz de Fora and vicinity: 2. Chagasia neivae; 43. Anopheles (N.) strodei (near

Agua Limpa station); 177. Sabethes (S.) purpureus; 196. Coquillettidia (R.) juxta-

mansonia; 266. Culex (C.) acharistus (Agua Limpa).

Lassance and vicinity, Vale do Rio das Velhas: 39. Anopheles (N.) braziliensis;

69. Anopheles (L.) gilesi; 284. Culex (Mel.) andricus.

Oliveira: 28. Anopheles (A.) tibiamaculatus; 53. Anopheles (N.) nigritarsis; 61.

Anopheles (N.) parvus.

Uberaba, near Belo Horizonte: 401. Dixella limai.

PARA

Belem (Para) and vicinity: 27. Anopheles (A.) shannoni; 35. Anopheles (N.)

emilianus; 130. Wyeomyia (D.) luteoventralis; 139. Wyeomyia (D.) subcomplosa

[see also Curralinho; Salvador (Bahia); Cuiaba (Mato Grosso); Sergipe]; 159. Limatus

durhamii; 161. Limatus paraensis; 170. Sabethes (S.) nitidus; 178. Sabethes (S.)

quasicyaneus (Utinga); 181. Sabethes (Sabethoides) confusus; 193. Coquillettidia

(R.) arribalzagae; 198. Coquillettidia (R.) lynchi; 204. Mansonia (M.) indubitans;

207. Mansonia (M.) titillans; 217. Uranotaenia pallidoventer; 224. Psorophora (P.)

scintillans; 251. Aedes (F.) mediomaculatus; 260. Aedes (S.) exagitans; 270, Culex

(C.) corniger; 301, Culex (Mel) flochi (Rio Moju); 304. Culex (Mel.) indecorabilis;

349. Culex (Micr.) shopei (Utinga); 355. Culex (Aed.) clastrieri; 356. Culex (Anoed.)

belemensis; 359. Culex (Anoed.) luteopleurus.

Boa Vista, Rio Tapajos: 56. Anopheles (N.) goeldii; 89. Trichoprosopon (T.)

townsendi; 220. Orthopodomyia townsendi.

Breves: 206. Mansonia (M.) pseudotitillans.

Cachoeira do Arari, Ilha do Marajo: 3/1. Anopheles (N.) marajoara; 33. Ano-

pheles (N.) domesticus.

Curralinho: 118. Wyeomyia (C.) kummi; 139. Wyeomyia (D.) subcomplosa (Rio

Canaticu) [see also Belem; Salvador (Bahia); Cuiaba,(Mato Grosso); Sergipe]; 26J.

Haemagogus (L.) tropicalis. |

- Guama (Sao Miguel do): 292. Culex (Mel.) contei.

Gurupa: 201. Mansonia (M.) amazonensis.

Oriboca, Rio Guajara: 290. Culex (Mel.) mojuensis.

Paragominas, municipio Capim: 319. Culex (Mel.) rachoui.

Santarem and vicinity: 100. Trichoprosopon (R.) edwardsianum (Belterra),; 396.

Sayomypia souzai.,

PARANA

Curitiba: 205. Mansonia (M.) pessoai.

Iguacu: 52. Anopheles (N.) guarani.

54 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

Londrina: 110. Wyeomyia (W.) limai; 156. Phoniomyia neivai.

RIO DE JANEIRO (STATE)

Angra dos Reis and vicinity: 326. Culex (Mochl.) megapus; 338. Culex (Micr.)

hedys; 376. Corethrella lopesi (Japuiba).

Baixada Fluminense, coastal lowlands: 29. Anopheles (N.) allopha (see also Rio

de Janeiro, Guanabara),; 58. Anopheles (N.) oswaldoi (see also Vale do Rio Doce,

Espirito Santo); 194. Coquillettidia (R.) chrysonotum (see also Vale do Rio Doce,

Espirito Santo); 216. Uranotaenia argenteopennis (see also Rio de Janeiro, Guana-

bara); 295. Culex (Mel.) dyius.

Cachoeira: 129. Wyeomyia (D.) knabi.

Chere: 327. Culex (Mochl.) galvaoi.

Imbarie (Estrella): 27. Anopheles (A.) minor.

Itaperuna: 12. Anopheles (A.) fluminensis.

Mage: 104. Trichoprosopon (R.) reversum; 106. Trichoprosopon (R.) theobaldi;

131. Wyeomyia (D.) melanoides; 275. Culex (C.) lygrus; 298. Culex (Mel.) evansae;

305. Culex (Mel.) cenus; 316. Culex (Mel.) oedipus; 345. Culex (Micr.) micro-

phyllus; 393. Lutzomiops nigrescens.

Mangaratiba: 90. Trichoprosopon (T.) obscurum; 138. Wyeomyia (D.) shannoni;

144. Phoniomyia davisi; 244. Aedes (O.) perventor.

Mendes: 114. Wyeomyia (W.) pintoi.

Orgaos, Serra dos; woods near Rio Soberbo at crossing of railroad from Mage |

to Teresopolis: 331. Culex (Micr.) aphylactus; 334. Culex (Micr.) trychnus.

Petropolis: 75. Toxorhynchites (L.) bambusicola; 82. Toxorhynchites (L.) posti-

catus; 86. Trichoprosopon (T.) compressum (see also Sao Paulo, Sao Paulo), 117.

Wyeomyia (C.) dyari.

Porto das Caixas and vicinity: 41. Anopheles (N.) darlingi (Caxiribu); 59. Ano-

pheles (N.) metcalfi; 146. Phoniomyia edwardsi; 154. Phoniomyia pilicauda; 294.

Culex (Mel.) exedrus; 351. Culex (Aed.) accelerans.

Sant’Anna: 320. Culex (Mel.) serratimarge.

Sao Bento: J 1. Anopheles (A.) evandroi.

Teresopolis: 98. Trichoprosopon(R.) castroi; 115. Wyeomyia (W.) sabethea.

Tingua: 167. Sabethes (S.) batesi; 185. Sabethes (Sabethinus) fabricii; 186. Sa-

bethes (Sabethinus) lutzianus.

~ Xerem: 13. Anopheles (A.) intermedius (see also Rio de Janeiro, Guanabara),

25. Anopheles (A.) pseudomaculipes; 126. Wyeomyia (D.) finlayi; 191. Coquil-

lettidia (R.) albicosta.

RIO GRANDE DO NORTE

Ares, near Sao Jose de Mipibu: 273. Culex (C.) foliaceus.

Natal, Macaphyba: 277. Culex (C.) oswaldoi.

RONDONIA

Porto Velho: 385. Lutzomiops amazonicus.

RORAIMA

Boa Vista: 297. Culex (Mel.) ernanii; 329, Culex (MochlL.) palaciosi.

Caracarai: 293, Culex (Mel.) cristovai; 306. Culex (Mel.) isabelae; 321. Culex

Belkin et al: Topotypic Brazilian Mosquitoes 55

(Mel.) silvai.

Carmo: 24. Anopheles (A.) celidopus.

San Alberto [?]: 182. Sabethes (Sabethoides) glaucodaemon.

Sororoca: 288. Culex (Mel.) bequaerti.

SANTA CATARINA

“Brusque: 71. Toxorhynchites (A.) catharinensis.

Ribeirao Pequena, municipio Laguna: 9. Anopheles (A.) bustamentei.

SAO PAULO

Avare, Fazenda Jose Euphrasio: 96. Trichoprosopon (I.) luederwaldti.

Bertioga, municipio Santos: 269. Culex (C.) carcinoxenus; 344, Culex (Micr.)

lanei.

Bocaina, Serra da; Fazenda do Bonito: 256. Aedes (F.) oswaldi.

Boraceia, municipio Salesopolis: 268. Culex (C.) brami.

Cabreuva: 243. Aedes (O.) pennai.

Campos do Jordao and vicinity: 34. Anopheles (N.) antunesi (Emilio Ribas);

49. Anopheles (N.) lanei (Emilio Ribas); 68. Anopheles (K.) laneanus (1600 m),

105. Trichoprosopon (R.) simile; 142. Phoniomyia antunesi; 384, Lutzomiops alti-

cola (1700 m); 404. Dixella wygodzinskyi (Vila Capivari, 1700 m).

Canaan: 226. Psorophora (J.) albigenu. |

Cantareira and vicinity: 5/. Anopheles (N.) niger; 134. Wyeomyia (D.) personata;

184, Sabethes (Sabethinus) aurescens; 188. Sabethes (Sabethinus) albiprivatus; 218.

Orthopodomyia albicosta (Serra da Cantareira); 263. Haemagogus (S.) capricornii

(Horto Florestal); 346. Culex (Micr.) neglectus (Serra da Cantareira); 363. Culex

(Car.) iridescens (Serra da Cantareira).

Caraguatatuba: 67. Anopheles (K.) montemor. ?

Casa Grande, municipio Mogi das Cruzes: 70. Anopheles (L.) pseudotibiamacu-

latus; 370. Corethrella cardosoi; 386. Lutzomiops barret toi.

Cubatao: 213. Uranotaenia geometrica.

Cubatao, Serra do: 162. Limatus flavisetosus.

Diabo, Serra do; rio Cuiaba, municipio Venceslau: 145. Phoniomyia diabolica.

Franca: 253. Aedes (F.) leucomelas. See also Rio Grande.

Guaratingueta: 55. Anopheles (N.) ayrozai.

Guaruja and vicinity: 10. Anopheles (A.) geometricus; 36. Anopheles (N.) guaru-

jaensis; 148. Phoniomyia galvaoi (Parada 24 de Outubro); 151. Phoniomyia lopesi

(Parada 24 de Outubro).

Itaici (Estacao de): 123. Wyeomyia(D.) bourrouli.

Itapura, Rio Tiete immediately above: 234. Psorophora(G.) scutipunctata.

Jaragua: 391. Lutzomiops lutzi.

Juquia and vicinity: 199. Coquillettidia (R.) neivai; 214, Uranotaenia mathesoni;,

369. Corethrella bromelicola (Poco Grande), 372. Corethrella flavitibia; 374. Core-

thrella infuscata; 377. Corethrella pillosa (Poco Grande); 378. Corethrella selvicola;

382. Corethrella vittata; 389. Lutzomiops iridescens; 390. Lutzomiops juquianus

bile Grande); 395. Sayomyia antunesi; 403. Dixella torrentia (Salto do Poco

rande).

Lagoa: 309. Culex (Mel.) lugens; 322. Culex (Mel.) theobaldi.

Lorena: 44. Anopheles (N.) ramosi.

Lussanvira: See Pereira Barreto.

56 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

Mirassol: 108. Wyeomyia (W.) downsi.

Palmeira: 379. Corethrella striata; 387. Lutzomiops coutinhoi. ,

Pereira Barreto (Lussanvira, Novo Oriente): 42. Anopheles (N.) paulistensis; 54.

Anopheles (N.) noroestensis.

Perus: 364. Culex (Car.) soperi; 373. Corethrella fulva (Agua Fria).

Ribeirao Preto: 285. Culex (Mel) aureonotatus (Fazenda Monte Alegre), 289.

Culex (Mel) bifoliolatus (Fazenda Monte Alegre); 325. Culex (Mel.) trilobulatus

(Rio Tamandua).

Rio Grande, near Franca: 250. Aedes (F.) fluviatilis; 252. Aedes (F.) tripunc-

fatus.

Rio Paraiba do Sul, valley of: 45. Anopheles (N.) arthuri; 46. Anopheles (N.)

artigasi; 47. Anopheles (N.) albertoi.

Santa Cruz do Rio Pardo: 274. Culex (C.) forattinii.

Santos: 225. Psorophora (P.) genumaculata.

Sao Paulo and vicinity: 1. Chagasia fajardi; 4. Chagasia stigmopteryx (Butantan),

6. Anopheles (S.) lineatus; 8. Anopheles (A.) anchietai (Cangaiba),; 14. Anopheles

(A.) maculipes; 17. Anopheles (A.) mediopunctatus; 30. Anopheles (N.) limai (Pin-

heiros and Butantan); 40. Anopheles (N.) pessoai (left bank of Rio Pinheiros, Pin-

heiros); 74. Toxorhynchites (A.) neglectus; 79. Toxorhynchites (L.) solstitialis; 80.

Toxorhynchites (L.) chrysocephalus; 86. Trichoprosopon (T.) compressum (see also

Petropolis, Rio de Janeiro); 91. Trichoprosopon (T.) pallidiventer; 94. Trichoproso-

pon (S.) fluviatile; 95. Trichoprosopon (S.) longipalpis; 107. Wyeomyia (W.) arthro-

stigma; 113. Wyeomyia (W.) oblita (Pacaembu); 116. Wyeomyia (W.) serrata (Rio

Frio, Pindamonhangaba); 120. Wyeomyia (M.) leucostigma; 125. Wyeomyia (D.)

confusa (woods near); 158. Phoniomyia tripartita; 163. Sabethes (S.) albiprivus;

164. Sabethes (S.) albiprivatus; 187. Sabethes (Sabethinus) intermedius (near); 192.

Coquillettidia (R.) albifera (Rio Pinheiros, Butantan); 197. Coquillettidia (R.) hypo-

cindyna; 208. Mansonia (M.) wilsoni; 211. Uranotaenia ditaenionota; 229. Psoro-

phora (J.) arribalzagae; 238. Aedes (O.) crinifer; 264. Culex (L.) brasiliae; 271. Cu-

lex (C.) deanei (Campo de Marte, Santana); 272. Culex (C.) bilineatus; 280. Culex

(C.) renatoi (Bairro Sao Miguel Paulista); 281. Culex (C.) spinosus; 299. Culex

(Mel.) fasciolatus (wooded mountains near); 302. Culex (Mel.) humilis; 303. Culex

(Mel.) inadmirabilis; 310. Culex (Mel.) nigrescens (Santo Amaro); 314. Culex (Mel.)

ocellatus; 339. Culex (Micr.) imitator; 347. Culex (Micr.) pleuristriatus; 394. Sayo-

myia braziliensis, 402. Dixella paulistana (Campo Experimental da Agua Funda).

Sao Vicente: 19. Anopheles (A.) limai; 20. Anopheles ( A. ) costai.

Tabatinguera: 350. Culex (Micr.) worontzowi.

Vera Cruz: 32. Anopheles (N.) imperfectus.

SERGIPE

Locality not specified: 139. Wyeomyia (D.) subcomplosa [see also Salvador

(Bahia); Cuiaba (Mato Grosso); Belem and Curralinho (Para) ].

Belkin et al: Topotypic Brazilian Mosquitoes a?

REFERENCES CITED

Antunes, P.C.A. and A.S. Ramos |

1939. Culex (Carrollia) iridescens, bonnei e soperi (Diptera, Culicidae). Bol. Biol.

(n.s.) 4:375-385.

Belkin, J.N.

1968. Mosquito Studies (Diptera, Culicidae). IX. The type specimens of New

World mosquitoes in European museums. Amer. Entomol. Inst., Contrib. 3(4).

69D.

Belkin, J.N., R.X. Schick and S.J. Heinemann

1965. Mosquito Studies (Diptera, Culicidae). V. Mosquitoes originally described

from Middle America. Amer. Entomol. Inst., Contrib. 1(5). 95 p.

1966. Mosquito Studies (Diptera, Culicidae). VI. Mosquitoes originally described

from North America. Amer. Entomol. Inst., Contrib. 1(6). 39 p.

1968. Mosquito Studies (Diptera, Culicidae). XI. Mosquitoes originally described

from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay. Amer. Entomol.

Inst., Contrib. 4(1):9-29. 8 Nov.

Cerqueira, N.L. |

1957. Descricao da larva e pupa de Aedes (Ochlerotatus) lepidus Cerqueira &

Paraense (Diptera:Culicidae, Culicini). Mus. Paraense Emilio Goeldi, Bol. (n.s.)

Zool. 3. 4 p.

1961. Distribuicao geografica dos mosquitos da Amazonia (Diptera, Culicidae,

Culicinae). Rev. Brasil. Entomol. 10:111-168.

Cerqueira, N.L. and J. Lane

1945. Note on Haemagogus capricornii Lutz, 1904 (Diptera, Culicidae). Entomol.

Soc. Wash., Proc. 47:279-288.

Davis,’ D.E. |

1944. Larval habits of some Brazilian mosquitoes. Rev. Entomol. 15:221-235.

Deane, M.P., O.R. Causey and L.M. Deane

1946. Studies on Brazilian anophelines from the Northeast and Amazon regions.

III. An illustrated key by larval characteristics for the identification of thirty-

two species of Anophelini, with descriptions of two larvae. Amer. J. Hyg.,

Monogr. Ser. 18:35-50.

Del Ponte, E.

1939. Identificacion de ‘“‘Sabethini’’ (Dip. Culicidae) por medio de tarjetas perfora-

das. Physis 17:535-541.

Del Ponte, E. and N. Cerqueira

1938. Alguns sabethineos do Brasil. Rev. Entomol. 8:225-237.

Durham, H.E.

1902. Report of the yellow fever expedition to Para. Liverpool Sch. Trop. Med.,

Mem. 4:479-563.

Dyar, H. G.

1919. A revision of the American Sabethini of the Sabethes group by the male

genitalia (Diptera, Culicidae). Insecutor Inscitiae Mens. 7:114-142. 17 Nov.

1928. The mosquitoes of the Americas. Wash., Carnegie Inst. Wash. (Publication

387). 616 D:

Fairchild, G.B.

1961. The Adolpho Lutz collection of Tabanidae (Diptera). I. The described gen-

era and species, condition of the collection, and selection of lectotypes. Inst.

58 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

Oswaldo Cruz, Mem. 59:185-249.

Galvao, A.L. Ayroza

1941. Contribuicao ao conhecimento das especies de Myzorhynchella (Diptera,

Culicidae). Sao Paulo (State), Arq. Zool. 2:505-575.

Galvao, A.L. Ayroza and J. Lane

1938. Notas sobre os Nyssorhynchus de S. Paulo. VI. Revalidacao de Anopheles

(Nyssorhynchus) oswaldoi Peryassu, 1922 e discussao sobre Anopheles (Nys-

sorhynchus) tarsimaculatus Goeldi, 1905. Livro Jubilar do Professor Lauro

Travassors. Rio de Janeiro, p. 169-178. |

Kumm, H.W. and O. Novis ;

1938. Mosquito studies on the Ilha de Marajo, Para, Brazil. Amer. J. Hyg. 27:

498-515.

Lane, J.

1943. Sobre o genero Uranotaenia (Diptera, Culicidae, Qulicini): Rev. Entomol.

14:137-161.

1945. Os sabetineos da America. (Addenda e corrigenda). Rev. Entomol. 16:132-

Lavra

Lane, J. and N.L. Cerqueira

1942, Os sabetineos da America (Diptera, Culicidae). Sao Paulo (State), Arq.

Zool. 3:473-849. 19 Oct.

1957. The validity and change of name of two species of Wyeomyia (Diptera,

Culicidae). Entomol. Soc. Wash., Proc. 59:244,

1958a. The types of Wyeomyia (Wyeomyia) dyari Lane & Cerqueira, 1942 (Dip-

tera, Culicidae). Entomol. Soc. Wash., Proc. 60:135.

1958b. Amazonian Chaoborinae (Diptera, Culicidae). Stud. Entomol. 1:561-566.

Lane, J. and L. Whitman

1951. The subgenus “Microculex”’ in Brazil (Diptera, Culicidae). Rev. Brasil. Biol.

11:341-366.

Lutz, A.

1905. Novas especies de mosquitos do Brasil. Imprensa Med. 13:26-29, 48-52,

65-70, 81-84, 101-104, 125-128, 169-172, 212-214, 269-271, 287- 290, ali

314, 347- 350. 25 Jan- 25 Sept.

Mattingly, P.F.

1958. Lectotypes of mosquitoes (Diptera:Culicidae) in the British Museum. Part

III. Genera Sabethes, Udaya and Aedes (subgenera Paraedes, Cancraedes and

Skusea). Roy. Entomol. Soc. London, Proc. (B) 27:105-108.

Peryassu, A.G.

1908. Os culicidos do Brasil. Rio de Janeiro, Inst. Manguinhos. 407 p.

Peters, T.M.

1968. Mosquito Studies (Diptera, Culicidae). X. Dixinae originally described from

North America. Amer. Entomol. Inst., Contrib. 4(1):1-7. 8 Nov.

Root, F.M.

1926. Studies on Brazilian mosquitoes. L The anophelines of the Nyssorhynchus

group. Amer. J. Hyg. 6:684-717.

Rozeboom, L.E. and W.H.W. Komp

1950. A review of the species of Culex of the subgenus Melanoconion (Diptera,

Culicidae). Entomol. Soc. Amer., Ann. 43:75-114.

Shannon, R. C.

1931. The environment and behaviour of some Brazilian mosquitoes. Entomol.

Soc. Wash., Proc. 33:1-27. |

Belkin et al: Topotypic Brazilian Mosquitoes 59

Stone, A.

1943. [Lectotype designation for Anopheles (N.) darlingi Root, 1926] in Ross,

E.S. and H.R. Roberts. Mosquito Atlas. Part I. p. 30. Phila., Amer. Entomol.

Soc,

1950. The larva of Culex foliaceus Lane (Diptera, Culicidae). Rev. Entomol. ah

237-239.

1957. Corrections in the taxonomy and nomenclature of mosquitoes (Diptera,

Culicidae). Entomol. Soc. Wash., Proc. 58:333-344.

1958. The identity of Culex aestuans Wiedemann (Diptera, Culicidae). Entomol.

Soc. Wash., Proc. 60:186.

1961. A synoptic catalog of the mosquitoes of the world, supplement I. (Diptera:

Culicidae). Entomol. Soc. Wash., Proc. 63:29-52.

1963. A synoptic catalog of the mosquitoes of the world, supplement IH. (Dip-

tera: Culicidae). Entomol. Soc. Wash., Proc. 65:117-140.

1966a. Family Dixidae. Sao Paulo, Univ. Sao Paulo, Mus. Zool. Cat. Diptera

Amer. S. of USA. 10. 4p.

1966b. Family Chaoboridae. Sao Paulo, Univ. Sao Paulo, Mus. Zool. Cat. Dip-

tera Amer. S. of USA. 11. 8p.

1967. A synoptic catalog of the mosquitoes of the world, supplement ILI. (Dip-

tera: Culicidae). Entomol. Soc. Wash., Proc. 69:197-224.

1970. A synoptic catalog of the mosquitoes of the world, supplement IV. Ento-

mol. Soc. Wash., Proc. 72-137-171.

Stone, A and K.L. Knight

1956a. Type specimens of mosquitoes in the United States National Museum: I,

The genus Aedes (Diptera, Culicidae). Wash., Acad. Sci., J. 46:213-228.

1956b. Type specimens of mosquitoes in the United States National Museum: III,

The genera Anopheles and Chagasia (Diptera, Culicidae). Wash. Acad. Sci., J.

46:276-280.

1957a. Type specimens of mosquitoes in the United States National Museum: IV,

The genus Culex (Diptera, Culicidae). Wash. Acad. Sci., J. 47:42-59.

1957b. Type specimens of mosquitoes in the United States National Museum, V:

The Sabethini (Diptera, Culicidae). Wash. Acad. Sci., J. 47:117-126.

Stone, A., K.L. Knight and H. Starcke

1959. A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae).

Wash., Entomol. Soc. Amer. (Thomas Say Found., vol. 6). 358 p.

Theobald, F.V.

1907. A monograph of the Culicidae or mosquitoes. Vol. 4. London, Brit. Mus.

Natur. Hist. 639 p.

1910. A monograph of the Culicidae or mosquitoes. Vol. 5. London, Brit. Mus.

(Natur. Hist.). 646 p.

U.S. Board on Geographic Names

1963. Brazil. Gazetteer No. 71. Wash., Dep. Interior, Office Geogr. 915 p.

Zavortink, T.J. |

1968. Mosquito Studies (Diptera, Culicidae). VIII. A prodrome of the genus

Orthopodomyia. Amer. Entomol. Inst., Contrib. 3(2). 221 p.

ADDENDUM

Belkin, J.N., S.J. Heinemann and W.A. Page

1970. The Culicidae of Jamaica (Mosquito Studies. XXI). Amer. Entomol. Inst.,

Contrib. 6(1). 458p. (Also Inst. Jamaica, Bull. Sci. Ser. 20)

60 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

INDEX TO SCIENTIFIC NAMES

abnormalis (265), Culex, 21, 36, 45, 52

accelerans (351), Culex, 27, 41, 50, 54

acharistus (266), Culex, 21, 41, 50, 53

adolphoi (66), Anopheles (K.), 7, 40, 43

aegypti, Aedes, 20

aenigmaticus (236), Aedes, 19, 31, 48, 52

aestuans (279), Culex, 22, 43, 49, 51

airosai (122), Wyeomyia, 11, 37, 47, 51

airozai (267), Culex, 21, 36, 45, 51

alagoanii (23), Anopheles, 4, 40, 48, 50

alani (121), Wyeomyia, 11, 33,37, 47

albertoi (47), Anopheles, 5, 43, 49, 56

albicosta (191), Coquillettidia, 16, 40, 47, 54

albicosta (218), Orthopodomyia, 18, 38, 47,

albifasciatus (237), Aedes, 19, 39, 49, 52

albifera (192), Coquillettidia, 16, 41, 46, 56

albigenu (226), Psorophora, 18, 40, 47, 55

albinensis, Culex, 22

albipes (330), Culex, 25, 39, 49, 51

albiprivatus (164), Sabethes (S.), 13, 38, 44,

albiprivatus (188), Sabethes (Sabethinus), 15,

34,42, 44, 55

albiprivus (163), Sabethes, 13, 14, 42, 44, 56

albitarsis, Anopheles, 4

- albitarsis (209), Uranotaenia, 17, 36, 44, 51

allopha (29), Anopheles, 4, 40, 49, 52, 54

alogistus, Culex, 25

alticola (384), Lutzomiops, 29, 36, 45, 55

amazonensis (352), Culex, 27, 39, 48, 51

amazonensis (201), Mansonia, 26, 42, 44, 53

amazonica (227), Psorophora, 18, 31, 45, 51

amazonicus (16), Anopheles, 3, 32, 43, 50

amazonicus (385), Lutzomiops, 29, 36, 50, 54

amazonicus (165), Sabethes, 14, 36, 44, 50

ambiguus (84), Toxorhynchites, 8, 34, 48, 51

americanus (354), Culex, 27, 40, 49, 50

anchietai (8), Anopheles, 2, 32, 45, 56

andricus (284), Culex, 22, 41, 50, 53.

anduzei (361), Culex, 27, 31, 37, 45, 51

antunesi (34), Anopheles, 4, 35, 46; 55

antunesi (362), Culex, 27, 37, 48, 51

antunesi (142), Phoniomyia, 12, 37, 45, 55

antunesi (395), Sayomyia, 29, 36, 45, 55

aphylactus (331), Culex, 25, 41, 50, 54

apicalis (233), Psorophora, 19, 42, 44, 52

appendiculata, Corethrella, 28

aquasalis, Anopheles, 4

argenteopennis (216), Uranotaenia, 17, 40, 48,

52, 54

argenteoumbrosus (340), Culex, 26, 42, 44,

SZ |

argyritarsis (37), Anopheles, 5, 41, 49, 52

argyronotum (169), Sabethes, 14, 34, 44, 52

arribalzagae (193), Coquillettidia, 16, 42, 44,

53 |

arribalzagae (229), Psorophora, 18, 35, 44, 56

arthrostigma (107), Wyeomyia, 10, 38, 47, 56

arthuri (45), Anopheles, 5, 43, 49, 56

artigasi (46), Anopheles, 5, 43, 49, 56

aureolineatus (257), Aedes, 20, 30, 50, 51

aureonotatus (285), Culex, 22, 34, 46, 56

-aurescens Lutz (184), Sabethes, 15, 38, 44,

aurescens Theobald (184), Sabethes, 15, 42

aureus (332), Culex, 25, 37, 48, 52

automartus (315), Culex, 24, 41, 50, 52

ayrozai (55), Anopheles, 6, 43, 49, 55

bahiensis (286), Culex, 22, 34, 45, 51

bambusicola (75), Toxorhynchites, 7, 39, 47,

baresi (262), Haemagogus, 21, 31, 45, 51

barrettoi (386), Lutzomiops, 29, 36, 45, 55

bastagarius, Culex, 22

batesi (167), Sabethes, 14, 37, 47, 54

belemensis (356), Culex, 27, 34, 45, 53

belisarioi (168), Sabethes, 14, 39, 48, 53

bequaerti (288), Culex, 23, 34, 50, 54

bigoti, Culex, 21

bifoliolatus (289), Culex, 23, 34, 46, 56

bilineatus (272), Culex, 21, 42, 44, 56

bipartipes, Sabethes, 14

bonnei (143), Phoniomyia, 12, 37, 45, 47, 52

bourrouli (123), Wyeomyia, 11, 39, 49, 55

brami (268), Culex, 21, 35, 45, 55

brasiliae (264), Culex, 21, 34, 50, 56

braziliensis (249), Aedes, 20, 36, 44, 51

braziliensis (39), Anopheles, 5, 31, 47, 53

braziliensis (394), Sayomyia, 29, 42, 44, 56

breviculus, Culex, 23

brevipes (93), Trichoprosopon, 9, 37, 47, 52

brevisector (397), Edwardsops, 30, 34, 44, 51

bromelicola (369), Corethrella, 28, 36, 45, 55

brucei (135), Wyeomyia, 11, 33, 48, 52

burkii (212), Uranotaenia, 17, 36, 45, 52

bustamentei (9), Anopheles 3, 35, 45, 55

calosomata, Uranotaenia, 17

canaanensis (357), Culex, 27, 37, 48, 51

capricornii (263), Haemagogus, 21, 39, 48, 55

Belkin et al: Topotypic Brazilian Mosquitoes 61

carcinoxenus (269), Culex, 21, 31, 46, 47, 55

cardosoi (370), Corethrella, 28, 36, 45, 55

carioca (333), Culex, 25, 37, 48, 52

castroi (98), Trichoprosopon, 9, 36, 47, 54

catharinensis (71), Toxorhynchites, 7, 38, 47,

celidopus (24), Anopheles, 4, 34, 50, 55

cenus (305), Culex, 24, 41, 50, 54

cerqueirai (202), Mansonia, 17, 30, 48, 52

cerqueirai (99), Trichoprosopon, 9, 42, 50, 52

_ cesari (124), Wyeomyia, 11, 33, 47, 52

chagasi (65), Anopheles, 7, 35, 48, 50

chagasi (203), Mansonia, 17, 37, 47, 53

_chapadensis (400), Dixella, 30, 36, 45, 52

chloropterus, Sabethes, 15

chrysocephalus (80), Toxorhynchites, 8, 42,

44, 56

chrysonotum (194), Coquillettidia, 16, 40, 47,

49, 51, 54

chrysothorax (291), Culex, 23, 40, 47, 52

chrysothorax (323), Culex, 25, 40, 44, 51

ciliata, Psorophora, 18

cilipes, Psorophora, 18

cingulata, Psorophora, 19

clarki (311), Culex, 24, 34, 44, 51

clastrieri (355), Culex, 27, 31, 44, 53

complosa, Wyeomyia, 33

compressum (86), Trichoprosopon, 8, 39, 47,

54, 56

confinnis, Psorophora, 19

confusa (368), Corethrella, 28, 36, 45, 52

confusa (125), Wyeomyia, 11, 39, 49, 56

confusus (181), Sabethes, 15, 42, 44, 53

consolator, Culex, 26

contei (292), Culex, 23, 34, 45, 53

corniger (270), Culex, 21, 42, 44, 53

- costai (20), Anopheles, 3, 34, 45, 56

costalimai (19), Anopheles, 3, 34, 45

coutinhoi (387), Lutzomiops, 29, 36, 45, 56

crinifer (238), Aedes, 19, 42, 44, 56

cristovai (293), Culex, 23, 34, 45, 54

cruzii (66), Anopheles (K.), 7, 34, 43

curvirostris (160), Limatus, 13, 37, 49, 52

cuyabensis (64), Anopheles, 6, 7, 40, 47, 53

cyaneus, Sabethes, 14

damascenoi (358), Culex, 27, 34, 45, 51

darlingi (41), Anopheles, 5, 41, 50, 54

davisi (335), Culex, 26, 36, 48, 51

davisi, Lutzomiops, 29

davisi (144), Phoniomyia, 12, 37, 46, 47, 54

davisi (210), Uranotaenia, 17, 36, 45, 51

deanei (271), Culex, 21, 32, 45, 56

delpontei (137), Wyeomyia, 12, 33, 37, 47,

diabolica (145), Phoniomyia, 12, 37, 45, 55

digitatum (87), Trichoprosopon, 8, 41, 48, 52

discrucians (288), Psorophora, 18, 43, 44, 52

ditaenionota (211), Uranotaenia, 17, 41, 46,

BSG |

dolosus, Culex, 21

domesticus (33), Anopheles, 4, 35, 48, 53

downsi (108), Wyeomyia, 10, 36, 45, 56

dubitans (336), Culex, 26, 37, 48, 52

dunhami (57), Anopheles, 6, 31, 50, 51

dunni, Culex, 23

durhamii (159), Limatus, 13, 42, 44, 53

dyari (117), Wyeomyia, 10, 12, 36, 45, 54

dyius (295), Culex, 23, 41, 50, 54

eastor, Culex, 23

edwardsi (371), Corethrella, 28, 36, 45, 52.

edwardsi (146), Phoniomyia, 12, 36, 50, 54

_ edwardsianum (100), Trichoprosopon, 9, 36,

47, 53

eiseni, Anopheles, 3

emilianus (35), Anopheles, 4, 36, 50, 53

ernanii (297), Culex, 23, 34, 45, 54

evandroi (11), Anopheles, 3, 37, 47, 54

evansae, Anopheles, 5

evansae (298), Culex, 23, 41, 50, 54

exagitans (260), Aedes, 20, 43, 44, 53

exedrus (294), Culex, 23, 41, 50, 54

fabricii (185), Sabethes, 15, 37, 47, 54

fajardi (1), Chagasia, 2, 39, 48, 56

fasciolatus (299), Culex, 23, 39, 47, 56

fascipes, Orthopodomyia, 18

faurani (300), Culex, 23, 34, 45, 51

ferox (81), Toxorhynchites, 8, 34, 43, 49, 51

finlayi (126), Wyeomyia, 11, 37, 47, 54

flabellata (147), Phoniomyia, 12, 37, 47, 51

flavicosta (240), Aedes, 19, 43, 44, 51

flavisetosus (162), Limatus, 13, 31, 48, 55

flavitibia (372), Corethrella, 28, 36, 45, 55

flochi (301), Culex, 23, 34, 45, 53

fluminensis (12), Anopheles, 3, 41, 50, 54

fluminensis (85), Toxorhynchites, 8, 40, 47,

52 |

fluviatile (94), Trichoprosopon, 9, 42, 44, 56

fluviatilis (250), Aedes, 20, 38, 44, 56

foliaceus (273), Culex, 21, 36, 45, 54

forattinii (274), Culex, 21, 32, 45, 56

forattinii (173), Sabethes, 14, 31, 45, 50

forceps (230), Psorophora, 18, 31, 47, 52

fulva (373), Corethrella, 28, 36, 45, 56

62 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

fulvithorax (258), Aedes, 20, 39, 49, 52

fulvus (239), Aedes, 19, 43, 49, 51

fuscatus (342), Culex, 26, 37, 48, 52

gairus (337), Culex, 26, 41, 50, 52

galvaoi (48), Anopheles, 5, 31, 46, 50

galvaoi (327), Culex, 25, 34, 45, 54

galvaoi (148), Phoniomyia, 13, 32, 45, 55

genumaculata (225), Psorophora, 18, 32, 47,

56 |

geometrica (213), Uranotaenia, 17, 42, 44, 55

geometricus (10), Anopheles, 3, 32, 49, 55

gilesi (69), Anopheles, 7, 40, 48, 53

glaucodaemon (182), Sabethes, 15, 34, 50, 54

goeldii (56), Anopheles, 6, 41, 50, 53

gordoni (283), Culex, 22, 34, 44, 51

guarani (52), Anopheles, 6, 42, 50, 53

guarujaensis (36), Anopheles, 4, 41, 48, 55

hedys (338), Culex, 26, 41, 50, 54

hermanoi (195), Coquillettidia, 16, 37, 45, 52

hildebrandi (353), Culex, 27, 34, 44, 50

horei (76), Toxorhynchites, 7, 36, 44, 50

hosautos, Wyeomyia, 10

howardi (127), Wyeomyia, 11, 37, 47, 51

humboldti (101), Trichoprosopon, 9, 36, 47,

humilis (302), Culex, 23, 42, 44, 56

hypocindyna (197), Coquillettidia, 16, 34, 50,

identicus, Sabethes, 15

imitator (339), Culex, 26, 42, 44, 56

imperfectus (32), Anopheles, 4, 32, 45, 56

inadmirabilis (303), Culex, 24, 34, 50, 56

incaudata (149), Phoniomyia, 13, 41, 50, 52

indecorabilis (304), Culex, 24, 42, 44, 53

indubitans (204), Mansonia, 17, 34, 50, 53

infuscata (374), Corethrella, 28, 36, 45, 55

inimitabilis, Culex, 26

innominatus (287), Culex, 22, 34, 44, 50

innovator (328), Culex, 25, 34, 44, 50

intermedius (13), Anopheles, 3, 40, 47, 52, 54

intermedius (343), Culex, 26, 37, 48, 52

intermedius (187), Sabethes, 15, 38, 44, 56

intricatus, Culex, 24

iridescens (363), Culex, 27, 39, 45, 55

iridescens (389), Lutzomiops, 29, 36, 45, 55

isabelae (306), Culex, 24, 34, 45, 54

jacobinae (241), Aedes, 19, 41, 50, 51

johnnyi (307), Culex, 24, 34, 45, 51

juquianus (390), Lutzomiops, 29, 36, 45, 55

juxtamansonia (196), Coquillettidia, 16, 31,

47, 53

kerri (308), Culex, 24, 34, 45, 51

kerri (128), Wyeomyia, 11, 33, 47, 52

knabi (129), Wyeomyia, 11, 37, 47, 54

konderi (60), Anopheles, 6, 35, 46, 48, 50

kummi (375), Corethrella, 28, 36, 44, 51

kummi (118), Wyeomyia, 10, 37, 47, 53

laneanus (68), Anopheles, 7, 32, 45, 55

lanei (49), Anopheles, 5, 35, 46, 55

lanei (344), Culex, 26, 32, 45, 55

lanei (231), Psorophora, 18, 42, 48, 52

lanei (174), Sabethes, 14, 31, 45, 50

lepidus (242), Aedes, 19, 31, 48, 53

leucocelaenus (253), Aedes, 20, 34, 44

leucomelas (253), Aedes, 20, 34, 38, 44, 55

leucophoebus (254), Aedes, 20, 35, 45, 50

leucostigma (120), Wyeomyia, 10, 39, 49, 56

leucotarsis (109), Wyeomyia, 10, 36, 46, 48,

52 |

lewisi (7), Anopheles, 2, 41, 50, 51

limai (19), Anopheles (A.), 3, 32, 45, 56

limai (30), Anopheles (N.), 4, 35, 37, 49, 56

limai (401), Dixella, 30, 41, 49, 53

limai (110), Wyeomyia, 10, 36, 47, 54

lineata, Psorophora, 18

lineatus (6), Anopheles, 2, 39, 47, 56

locuples (171), Sabethes, 14, 41, 49, 52

longfieldae (166), Sabethes, 14, 34, 44, 52

longipalpis (219), Orthopodomyia, 18, 40, 44,

longipalpis (95), Trichoprosopon, 9, 39, 47,

longipes, Trichoprosopon, 9

longirostris (150), Phoniomyia, 13, 42, 44, 52

lopesi (376), Corethrella, 28, 36, 45, 54

lopesi (151), Phoniomyia, 13, 32, 45, 55

luederwaldti (96), Trichoprosopon, 9, 36, 45,

es)

lugens (309), Culex, 24, 39, 47, 55

lunatum (103), Trichoprosopon, 9, 42, 44, 52

luteopleurus (359), Culex, 27, 42, 44, 53

luteoventralis (130), Wyeomyia, 11, 42, 44,

lutzi (22), Anopheles, 3, 32, 34, 47, 53

lutzi (391), Lutzomiops, 29, 36, 45, 55

lutzi (111), Wyeomyia, 10, 37, 47, 52

lutzianus (186), Sabethes, 15, 37, 47, 54

lutzii (66), Anopheles (K.), 7, 34, 40, 42, 43,

lutzii (50), Anopheles (N.), 5, 6, 32, 47, 52

lutzii (232), Psorophora, 19, 42, 44, 52

lutzii (175), Sabethes, 14, 42, 47, 50

lygrus (275), Culex, 22, 41, 50, 54

lynchi (198), Coquillettidia, 16, 42, 50, 53

Belkin et al: Topotypic Brazilian Mosquitoes 63

maculata (3), Chagasia, 2, 40, 48, 53

maculipes (14), Anopheles, 3, 42, 43, 56

magnificus (398), Edwardsops, 30, 36, 45, 53

manaosensis (296), Culex, 23, 34, 44, 51

manaosensis (392), Lutzomiops, 29, 36, 45,

marajoara (31), Anopheles, 4, 35, 46, 53

mariae (77), Toxorhynchites, 7, 31, 49, 51

mathesoni (214), Uranotaenia, 17, 36, 45, 55

-mathisi (248), Aedes, 20, 40, 49, 50

mattinglyi (119), Wyeomyia, 10, 36, 48, 51

mattogrossensis (15), Anopheles, 3, 39, 47, 52

mauesensis (276), Culex, 22, 36, 45, 51

medioalbipes (112), Wyeomyia, 10, 38, 44, 51

mediomaculatus (251), Aedes, 20, 42, 44, 53

mediopunctatus (17), Anopheles, 3, 42, 43, 56

megapus (326), Culex 25, 41, 50, 54

melanocephala, Wyeomyia, 11 |

melanoides (131), Wyeomyia, 11, 41, 50, 54

melanonymphe (188), Sabethes, 15, 34, 44

metcalfi (59), Anopheles, 6, 35, 37, 50, 54

microphyllus (345), Culex, 26, 41, 50, 54

minor (21), Anopheles, 3, 37, 47, 54

mojuensis (290), Culex, 23, 34, 45, 53

molestus (367), Culex, 28, 36, 49, 52

montemor (67), Anopheles, 7, 32, 45, 55

mystes (132), Wyeomyia, 11, 34, 50, 52

nataliae, Uranotaenia, 17

neglectus (346), Culex, 26, 38, 50, 55

neglectus (74), Toxorhynchites, 7, 39, 49, 56

negrensis (133), Wyeomyia, 11, 36, 44, 50

neivae (2), Chagasia, 2, 32, 48, 53

neivai (199), Coquillettidia, 16, 37, 45, 55

neivai (156), Phoniomyia, 13, 37, 46, 47, 54

neoapicalis (233), Psorophora, 19, 42, 44

niger (51), Anopheles, 6, 42, 43, 55

niger (388), Lutzomiops, 29, 36, 45, 52

nigrescens (310), Culex, 24, 42, 44, 56

nigrescens (393), Lutzomiops, 29, 36, 45, 54 —

nigricorpus (312), Culex, 24, 42, 44, 50

nigrimacula (313), Culex, 24, 37, 48, 52

nigritarsis (53), Anopheles, 6, 32, 48, 53

nimbus, Anopheles, 2

nitidus (170), Sabethes, 14, 42, 44, 53

noctivaga (215), Uranotaenia, 17, 40, 48, 52

noroestensis (54), Anopheles, 6, 35, 37, 45,

46, 56

nuneztovari, Anopheles, 6

oblita (113), Wyeomyia, 10, 38, 39, 44, 47,

obscurum (90), Trichoprosopon, 8, 36, 47, 54

ocellatus (314), Culex, 24, 42, 44, 56

oedipus (316), Culex, 24, 41, 50, 54

originator (360), Culex, 27, 36, 44, 51

oswaldi (256), Aedes, 20, 38, 39, 44, 55

oswaldoi (58), Anopheles, 6, 40, 49, 51, 54

oswaldoi (277), Culex, 22, 35, 50, 54

palaciosi (329), Culex, 25, 34, 45, 54

pallidiventer (91), Trichoprosopon, 8, 38, 44,

pallidoventer (152), Phoniomyia, 13, 42, 44,

pallidoventer (217), Uranotaenia, 18, 42, 44,

pallipes (366), Culex, 28, 41, 49, 52

palmata (153), Phonioinyia, 13, 37, 47, 52

paraensis (161), Limatus, 13, 42, 44, 53

paramaxi (278), Culex, 22, 33, 45, 53

parvus (61), Anopheles, 6, 31, 47, 53

paulistana (402), Dixella, 30, 37, 45, 56

paulistensis (42), Anopheles, 5, 32, 35, 37,

48, 56 |

pennai (243), Aedes, 19, 30, 37, 45, 55

personata (134), Wyeomyia, 11, 38, 44, 55

perterrens (235), Psorophora, 19, 43, 44, 49,

perventor (244), Aedes, 19, 31, 48, 54

peryassui (22), Anopheles, 3, 4, 34, 47

pessoai (40), Anopheles, 5, 35, 37, 46, 56

pessoai (205), Mansonia, 17, 30, 45, 53

phlogistus, Culex, 24 3

pilicauda (154), Phoniomyia, 13, 41, 50, 54

pilipes (223), Psorophora, 18, 39, 49, 52

pillosa (377), Corethrella, 28, 36, 45, 55

pintoi (114), Wyeomyia, 10, 37, 47, 54

plectoporpe (317), Culex, 24, 41, 50, 52

pleuristriatus (347), Culex 26, 42, 44, 56

posticatus (82), Toxorhynchites, 8, 39, 47, 54

pseudomaculipes (25), Anopheles, 4, 40, 47,

pseudotibiamaculatus (70), Anopheles, 7, 35,

46, 55

pseudotitillans (206), Mansonia, 17, 42, 44,

purpureus Peryassu (177), Sabethes, 14, 34,

40, 47, 53

purpureus Theobald (176), Sabethes, 14, 42,

AG, BOs

purpureus (72), Toxorhynchites, 7, 42, 44, 50

pusillus (78), Toxorhynchites, 7, 37, 47, 52

putumayensis (318), Culex, 24, 39, 50, 51

quasicyaneus (178), Sabethes, 15, 40, 48, 53

quasilongirostris (255), Phoniomyia, 13, 42,

44,52

64 Contrib. Amer. Ent. Inst., vol. 7, no. 5, 1971

quasilongirostris of Dyar, 1928, Phoniomyia,

quinquefasciatus, Culex, 22

rachoui (26), Anopheles, 4, 35, 45, 51

rachoui (319), Culex, 25, 34, 45, 53

ramosi (44), Anopheles, 5, 43, 49, 55

reducens (348), Culex, 26, 37, 48, 52

remipes (172), Sabethes, 14, 43, 50, 51

remipusculus (177), Sabethes, 14, 34, 47

renatoi (280), Culex, 22, 37, 45, 56

retrosus (341), Culex, 26, 37, 48, 52

reversum (104), Trichoprosopon, 9, 36, 47,

thyacophilus (245), Aedes, 19, 37, 48, 51

rockefelleri (18), Anopheles, 3, 40, 48, 52

rondoni (62), Anopheles 6, 7, 39, 47, 52

rooti (121), Wyeomyia (A.), 11, 33, 37, 47,

rooti (136), Wyeornyia (D.), 11, 12, 32, 33,

47, 52

rozeboomi (5), Chagasia, 2, 31, 49, 51

sabethea (115), Wyeomyia, 10, 36, 47, 54

sampaioi (221), Orthopodomyia, 18, 37, 47,

sawyeri (38), Anopheles, 5, 31, 50, 51

scapularis (246), Aedes, 19, 41, 48, 52

scintillans (224), Psorophora, 18, 43, 44, 53

scutipunctata (234), Psorophora, 19, 39, 47,

59 3

selvicola (378), Corethrella, 28, 36, 45, 55

serrata (116), Wyeomyia, 10, 38, 47, 56

serratimarge (320), Culex, 25, 41, 50, 54

serratus (247), Aedes, 20, 42, 44, 52

shannoni (27), Anopheles, 4, 32, 50, 53

shannoni (200), Coquillettidia, 16, 30, 36, 45,

shannoni (179), Sabethes, 15, 31, 45, 50

shannoni (138), Wyeomyia, 12, 37, 47, 54

shopei (349), Culex, 27, 35, 45, 53

silvai (321), Culex, 25, 34, 45, 54

simile (105), Trichoprosopon, 9, 36, 47,55 .

soaresi (92), Trichoprosopon, 9, 36, 47, 51

solstitialis (79), Toxorhynchites, 8, 38, 44, 56

soperi (364), Culex, 28, 30, 37, 45, 56

soperi (189), Sabethes, 15, 37, 47, 51

souzai (396), Sayomyia, 29, 36, 45, 53

spinosus (281), Culex, 22, 38, 44, 56

spixi (180), Sabethes, 15, 31, 45, 51

splendens (88), Trichoprosopon, 8, 39, 49, 50

stigmopteryx (4), Chagasia, 2, 39, 44, 56

striata (379), Corethrella, 28, 36, 45, 56

strodei (43), Anopheles, 5, 41, 50, 53

subcomplosa (139), Wyeomyia, 12, 32, 33, 48,

54952553656

tarsata (380), Corethrella, 28, 36, 44, 51

tarsata (140), Wyeomyia, 12, 37, 47, 51

terrens (255), Aedes, 20, 43, 44, 52

theobaldi (322), Culex, 25, 38, 44, 55

theobaldi (157), Phoniomyia, 13, 37, 46, 47,

theobaldi (106), Trichoprosopon, 10, 36, 47,

thomasi (7), Anopheles, 2, 41, 50

thomasi (324), Culex, 25, 34, 44, 51

tibialis (222), Psorophora, 18, 41, 49, 52

tibiamaculatus (28), Anopheles, 4, 39, 48, 53

titillans (207), Mansonia, 17, 43, 44, 53

torrentia (403), Dixella, 30, 36, 45, 55

townsendi (220), Orthopodomyia, 18, 37, 47,

townsendi (89), Trichoprosopon, 8, 42, 50, 53

toxorhynchus (259), Aedes, 20, 39, 49, 52

travassosi (381), Corethrella, 29, 36, 45, 52

triannulatus (63), Anopheles, 6, 7, 40, 48, 53

trichopus (102), Trichoprosopon, 9, 34, 50,

il eed

trichopygus (73), Toxorhynchites, 7, 43, 49,

7 eee

tridentatus (183), Sabethes, 15, 31, 45, 51

trilobulatus (325), Culex, 25, 34, 46, 56

tripartita (158), Phoniomyia, 13, 31, 49, 56

tripunctatus (252), Aedes, 20, 42, 44, 56

tropicalis (261), Haemagogus, 20, 30, 31, 48,

53 |

trychnus (334), Culex, 26, 41, 50, 54

undulata (141), Wyeomyia, 12, 33, 47, 52

unicolor (399), Edwardsops, 30, 36, 45, 53

varipes, Psorophora, 18

violaceus (83), Toxorhynchites, 8, 43, 49,51

virgultus (282), Culex, 22, 42, 44, 49, 52

vittata (382), Corethrella, 29, 36, 45, 55

walcotti (97), Trichoprosopon, 9, 36, 47, 51

whitmani (383), Corethrella, 29, 36, 45, 51

whitmani (190), Sabethes, 16, 37, 47, 51

wiedemanni (81), Toxorhynchites, 8, 34, 49

wilsoni (365), Culex, 28, 37, 48, 51

wilsoni (208), Mansonia, 17, 30, 46, 48, 56

worontzowi (350), Culex, 27, 41, 46, 56

wygodzinskyi (404), Dixella, 30, 36, 45, 55

No.

(Continued from inside front cover)

Parts of Volume 4, with prices

Mosquito studies (Diptera, Culicidae), 1968.

X. Peters, T. Michael. Dixinae originally described from North America. 7 pages.

XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes

originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay.

29 pages. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XII. Berlin, O. G. W. A revision of the Neotropical subgenus Howardina of Aedes.

Carlson, Robert W. and Fred B. Knight, 1969.

Biology, taxonomy, and evolution of four sympatric Agrilus beetles (Coleoptera:

Buprestidae). 104 pages, 47 figures. Price: $3.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XIII. Barr, Ralph A. and Sylvia Barr. Pupae of the genus Psorophora in America

north of Mexico. 19 pages, 11 figures.

XIV. Berlin, O. G. W. Redescription of Toxorhynchites (T.) gigantulus from the

Philippines. 7 pages, 2 figures.

XV. Zavortink, Thomas J. A new species of treehole breeding Anopheles from

the southwestern United States. 12 pages, 4 figures.

Price: $1.25, postpaid.

Matthews, Robert W., 1970. A revision of the genus Spathius in America north of Mexico

(Hymenoptera, Braconidae). 86 pages, 31 figures. Price: $3.50, postpaid.

postpaid.

Parts of Volume 5, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XVI. Zavortink, Thomas J. A new species of treehole breeding Aedes (Ochlerotatus)

from southern California. 6 pages, 2 figures.

XVII. Adames, Abdiel J. and Charles L. Hogue. Two new species of Deinocerites from

Costa Rica. 6 pages, 6 figures.

XVIII. Berlin, O. G. W. The subgenus Micraedes of Culex. 30 pages, 12 figures, 1 map.

Price: $2.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XIX. Zavortink, Thomas J. The treehole Anopheles of the New World. 35 pages, 12

figures. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XX. Schick, Robert X. The Terrens Group of Aedes (Finlaya). 158 pages, 60 figures.

Price: $5.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

V. Reinert, John F. Genus Aedes, subgenus Diceromyia Theobald in southeast Asia.

43 pages, 15 figures. Price: $1.50, postpaid.

Howden, Anne T. The Tanymecini of the West Indies (Coleoptera: Curculionidae). 74 pages,

76 figures. Price: $2.75, postpaid.

Kethley, John B., 1970. A revision of the family Syringophilidae (Prostigmata: Acarina).

76 pages, 38 figures. Price: $2.75, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

VI. Mattingly, P. F. The genus Heinzmannia Ludlow in southeast Asia. 104 pages,

51 figures. Price: $4.00, postpaid.

Parts of Volume 6, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXI. Belkin, John N., Sandra J. Heinemann, and William A. Page. The Culicidae of

Jamaica. 458 pages, 126 figures. Price: $15.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

Vil. Tyson, W. H. Genus Aedeomyia Theobald in southeast Asia. 27 pages, 6 figures.

Vill. Tyson, W. H. Genus Aedes, subgenus Mucidus Theobald in southeast Asia. 54 pages,

21 figures. Price: $3.50, postpaid.

Parts of Volume 7, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXII. Zavortink, Thomas J. A new subgenus and species of Aedes from Arizona.

11 pages, 2 figures.

XXIII. Schick, Robert X. Additions and corrections to the revision of the Aedes

terrens group. 27 pages, 8 figures. Price: $1.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1971.

XXIV. Adames, Abdiel J. A revision of the crabhole mosquitoes of the genus

Deinocerites. 154 pages, 58 figures. Price: $6.00, postpaid.

(Continued from inside back cover)

No. 3. Contributions to the mosquito fauna of southeast Asia, 1971.

IX. Zavortink, Thomas J. The genus Orthopodomyia Theobald in Southeast Asia.

37 pages, 12 figures.

X. Dobrotworsky, N. V. The genus Culiseta Felt in Southeast Asia. 24 pages, 8 figures.

XI. Sirivanakarn, Sunthorn. A proposed reclassification of Neoculex Dyar based

principally on the male terminalia. 24 pages, 3 figures. Price: $3.50, postpaid.

No. 4. Not yet published.

No. 5. Mosquito studies (Diptera, Culicidae), 1971.

XXV. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes originally

described from Brazil. 64 pages. Price: $2.50, postpaid.

4+L/

pet.

Contributions

of the

American Entomological Institute

Volume 7, Number 6, 1971

Ms, oy

MOSQUITO STUDIES (Diptera, Culicidae)

XXVI. Winter biology of Culex tarsalis in

Imperial Valley, California

Michael J. Nelson

oT Mer yo

SNM Naas

007 141971 }

Linn. wm, ore

CONTRIBUTIONS

of the

AMERICAN ENTOMOLOGICAL INSTITUTE

The Contributions are for larger papers on insects. Each paper is a separate number, with separate

pagination and index. Separate numbers aggregating about 500 pages constitute a volume. Issues appear

irregularly, as suitable manuscripts are available.

Copies are sold separately or in subscriptions to complete volumes. Complete volumes are $12.00.

The price of separate numbers varies. Subscribers are billed for each volume with its beginning number,

and receive the parts as issued. Orders for separate numbers that total less than $8.00 must be accom-

panied by payment. Address orders or correspondence to the American Entomological Institute, 5950

Warren Road, Ann Arbor, Michigan 48105, U.S.A.

Parts of Volume 1, with prices

No. 1. Dasch, Clement E., 1964. The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae).

75 pages, 69 figures. Price: $2.25, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1965.

I, Belkin, John N., etal. A project for a systematic study of the mosquitoes of Middle America.

II. Belkin, John N., et al. Methods for the collection, rearing and preservation of mosquitoes.

78 pages, 4figures. Price: $2.25, postpaid.

No. 2a. Same as no. 2, but in Spanish. Price: $2.25, postpaid.

No. 3. Matthews, Robert W., 1965. The biology of Heriades carinata Cresson. (Hymenoptera,

Megachilidae). 33 pages, 23 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1965.

Ill. Ramalingam, Shivaji, and John N. Belkin. Two new Aedes from Tonga and Samoa.

IV. Belkin, John N. The mosquitoes of the Robinson-Peabody Museum of Salem expedition to

the southwest Pacific, 1956.

34 pages, 3 figures. Price: $1.00, postpaid.

No. 5. Mosquito studies (Diptera, Culicidae), 1965.

V. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann,

Mosquitoes originally described from Middle America. 95 pages. Price: $2.75 postpaid.

No. 6. Mosquito studies (Diptera, Culicidae), 1966.

VI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann.

Mosquitoes originally described from North America. 39 pages. Price: $1.00, postpaid.

No. 7. DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae (Homopotera,

Cicadellidae). A synopsis of the genus Ponana.

86 pages, 257 figures. Price: $2.50, postpaid.

No. 8. Contributions to the mosquito fauna of southeast Asia, 1967.

I. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald in Thailand.

55 pages, 20 figures. Price: $1.75, postpaid.

Parts of Volume 2, with prices

No. 1. Contributions to the mosquito fauna of southeast Asia, 1967.

Il. Bram, Ralph A. The genus Culex in Thailand (Diptera: Culicidae).

296 pages, 104 figures. Price: $9.00, postpaid.

No. 2. Schuh, Toby, 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region.

35 pages, 30 figures. Price: $1.25, postpaid.

No. 8. Snider, Richard J., 1967. The chaetotaxy of North American Lepidocyrtus s. str. (Collembola:

Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid. e

No. 4. Contributions to the mosquito fauna of southeast Asia, 1968.

III. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald of southeast

Asia. 76 pages, 26figures. Price: $2.25, postpaid.

No. 5. Contributions to the mosquito fauna of southeast Asia, 1968.

IV. Knight, Kenneth L. Species of the subgroup Chrysolineatus of group D, genus Aedes,

subgenus Finlaya Theobald. 45 pages, 12 figures. Price: $1.50, postpaid.

Parts of Volume 3, with prices

No. 1. Mosquito studies (Diptera, Culicidae), 1968.

Vil. Belkin, John N. The Culicidae of New Zealand.

182 pages, 30 figures. Price: $5.50, postpaid.

No. 2. Mosquito studies (Diptera, Culicidae), 1968.

VIll. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia.

221 pages, 41 figures. Price: $6.75, postpaid.

No. 3. Fleming, Richard C., 1968. Head musculature of sphinx moths (Lepidoptera: Sphingidae).

32 pages, 37 figures. Price: $1.00, postpaid.

No. 4. Mosquito studies (Diptera, Culicidae), 1968.

IX. Belkin, John N. The type specimens of New World mosquitoes in European museums,

69 pages. Price: $2.00, postpaid.

(Continued on back cover)

Contributions

of the

American Entomological Institute

Volume 7, Number 6, 1971

—_—

MOSQUITO STUDIES (Diptera, Culicidae)

XXVI. Winter biology of Culex tarsalis in

Imperial Valley, California

Michael J. Nelson

CONTENTS

ABSTRACT. .

INTRODUCTION

BACKGROUND .

Bionomics :

Vector Relationships

Overwintering of Arboviruses .

METHODOLOGY AND PRELIMINARY S STUDIES

Study Area .

Sampling Procedure.

Collecting methods. .. .

Light traps with CO, attractant .

CO, traps Seta g

Shelters

Larval sollccHons:

Sources of error . ‘ |

Error due to sampling procedure

Bias due to behavior of mosquitoes

Shelter preference

Dispersal . F

Physiological Age-rading

Criteria

Dissections .

Sources of error .

Gonotrophic ‘mau alion:

Autogeny .

Analysis of Data. .

RESULTS AND DISCUSSION

Weather

Population Bea

Mobility .

Females

Males

Immatures

Physiological Pacancters”

Fat body ian ea

Insemination

Blood feeding and ovarian a development.

Patty

CONCLUSIONS . .

REFERENCES CITED

FIGURES

OOM OUYUNAMNBRWWNHNK

MOSQUITO STUDIES (Diptera, Culicidae)

XXVI. WINTER BIOLOGY OF CULEX TARSALIS

IN IMPERIAL VALLEY, CALIFORNIA’

Michael J. Nelson”

ABSTRACT

Culex tarsalis Coquillett, 1896 is the most important vector of the viruses of

western equine encephalomyelitis and St. Louis encephalitis in the western United

States during the summer, but the overwintering mechanism for these 2 viruses is un-

known. In most areas, tarsalis cannot carry virus through the winter because blood-

fed females do not survive the coldest months. Overwintering females develop a large

fat body and experience a developmental diapause. They usually do not take their

first blood meal until the late winter or early spring. However, the mild winters in

the Imperial Valley of southern California are more conducive to mosquito activity.

From September 1967 until April 1970, a population of tarsalis was studied at

the Wister Unit of the Imperial Waterfowl Management Area, Imperial County, Cali-

fornia, to determine to what degree feeding and other activities were maintained

throughout the year. Each month, adult mosquitoes were collected in light traps

with CO, attractants, CO, bait can traps, and resting shelters. Monthly searches

for breeding sites were made.

The following characteristics of the females were recorded: external condition,

insemination, stage of ovarian development, presence or absence of muscle tissue

remnants, meconium, vertebrate blood, fat body, coiled ovarian tracheoles, and fol-

licular relics. The degree of error in the interpretation of these age-grading criteria

due to autogeny was similar to that of comparable studies of the winter biology

of tarsalis in the Central Valley of California.

There were 2 depressions of the population density each year: a major one in

the winter and a lesser in the summer. In contrast to studies to the north, at Wister

during every winter month females were attracted to CO,, males were found in

shelters, and pupae and larvae of all instars were collected. Fat body developed only

slightly and was largely exhausted before the coldest days of winter. The percentages

‘Contribution from project “Mosquitoes of Middle America” supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478. Based on PhD dissertation submitted to the University

of California, Los Angeles.

* Department of Zoology, University of California, Los Angeles, California 90024. Present

address: WHO-Aedes Research Unit, c/o Ministry of Public Health, Devavesm Palace, Bangkok,

Thailand.

Z Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

of blood-engorged, gravid, and parous females decreased in the autumn, but recovered

in December.

In the Imperial Valley of California, winter diapause of tarsalis is very weakly

expressed. The duration of diapause is different at the various latitudes within the

range of this species. A combination of short photoperiod and low temperature ap-

parently initiates diapause at all latitudes. The relatively high fall and winter temper-

atures in the Imperial Valley are probably largely responsible for the weak expression

and short duration of diapause there.

Potentially, tarsalis is a good candidate for overwintering arboviruses in the Im-

perial Valley. However, other workers have thus far failed to isolate the viruses from

this species or from vertebrates in December and January at Wister. Without direct

evidence of the viruses in the winter, it is not possible to determine the population

density of mosquitoes and the level of blood feeding necessary to maintain the cycle

of transmission.

INTRODUCTION

Culex tarsalis Coquillett, 1896 is the most important vector of western equine

encephalomyelitis virus (WEE) and St. Louis encephalitis virus (SLE) in the west-

ern United States. Although the mosquito-avian transmission cycle of these viruses

has been well documented for the summer months, the overwintering mechanism

is unknown. In most areas, tarsalis cannot overwinter virus because blood-fed fe-

males do not survive the coldest months. However, the mild winters in the Imperial

Valley of southern California are more conducive to mosquito activity.

In August 1967, following a report in June of 2 horse cases of WEE in Imperial

County, Dr. Telford H. Work and I made a preliminary mosquito survey of the

area. The Wister Unit of the Imperial Wildlife Management Area (p. 7) appeared

to be a potentially favorable area for virus activity, as large populations of both

mosquitoes and birds were present. Monthly mosquito collections and virus iso-

lation attempts were initiated in September 1967. It soon became evident that,

contrary to the behavior of tarsalis in other study areas, the Wister population

remained relatively active throughout the winter. We speculated that if blood feed-

ing continued at high level throughout the year, a mosquito-avian cycle could con-

tinue without interruption, and in the spring migrating birds might carry the virus

northward to areas from which it disappeared each winter. Therefore, I initiated

a 33-month study to determine to what degree tarsalis continued to feed and to

maintain other activities throughout the year. Seasonal fluctuations in the follow-

ing aspects of its biology were » vestigated and compared to similar studies in

other areas:

I. Population density II. Physiological parameters

A. Mobility A. Fat body development

B. Females B. Insemination

C. Males C. Blood feeding and ovarian eres aien

D. Immatures D. Oviposition

I wish to express my unending gratitude to Dr. John N. Belkin for his invaluable

assistance and guidance during this study. I thank Dr. A. Ralph Barr, Dr. Austin J.

MacInnis, Dr. Marietta Voge, and Dr. Donald Landenberger for their helpful ad-

vice in the development and writing of the dissertation, Dr. William C. Reeves

for his critical reading of the manuscript and valuable suggestions, and Dr. Telford

H. Work who first stimulated my interest in the mosquitoes of the Imperial Valley

Nelson: Winter Biology of Culex tarsalis 3

of California. I am greatly indebted to the staff of the Imperial Waterfowl Manage-

ment Area, especially C. Ray Knight, Richard Weaver, and Carl Miller, without

whose cooperation this project would not have been possible. I wish also to ex-

tend my sincerest appreciation to Claire Price who typed the preliminary and final

copies of the manuscript, to Margaret Kowalczyk who did such excellent work

in inking the graphs, and Caryle L. Stallard for editorial assistance and the prepara-

tion of the text copy for lithoprinting. This investigation was supported in part

by U.S. Public Health Service grants T1-AI-132 and T0O1-AI-00070. Some of the

field work was supported by National Science Foundation grant GB 8075.

BACKGROUND

BIONOMICS

Culex tarsalis occurs in the western, central and southern United States, in west-

ern Canada north to Northwest Territories (Carpenter and LaCasse, 1955:296) and

in Mexico south to the state of Chiapas (Vargas, 1956:27). Jenkins cites collec-

tions up to 2700 meters in Utah (1950:914).

Normally, a raft of 200-300 eggs is deposited on the surface of the breeding

water and hatching takes place after 2 or 3 days (Brennan and Harwood, 1953:156).

The immature stages are found in a variety of habitats, most commonly in still

ground pools in hot, irrigated valleys. The breeding water may range from fresh

to very foul and usually contains much organic matter (Carpenter and LaCasse,

1955:296; Jenkins, 1950:914; Sjogren, 1968). In the field, larvae survive freezing

temperatures (Washino and Bellamy, 1963) and thermal waters at 39° C (Jenkins,

1950:914). The larval stage, consisting of four stadia, lasts only 10 days and the

pupal stage 3 days at 24-26° C (Brennan and Harwood, 1953:155-156).

In the summer, females are gonoactive. During the first day or two after emer-

gence they rest, imbibe plant juices and mate. Then a host is sought for a blood

meal. Although preferred hosts are homeothermic vertebrates, especially birds (Tem-

pelis et al., 1965), feeding occurs readily on reptiles and amphibians (Gebhardt

et al., 1966; Henderson and Senior, 1961) and, in the laboratory, even on cater-

pillars (Harris et al., 1969). Blood digestion takes about 3 days at 24-27° C, but

requires 3 weeks or longer at 10-13° C (Bellamy and Reeves, 1963:320; Rush,

Kennedy and Eklund, 1963a:261).

Activities such as flying, mating, host-seeking and feeding occur largely during

the twilight hours. During the day, adults rest in protected shelters but will be-

come active and bite if disturbed. Females are capable of dispersing over relatively

great distances. Dow, Reeves and Bellamy (1965) showed that marked adults dis-

persed up to 15 km into a larvicided area, independently of wind direction. Bailey

et al. (1965:111) recovered a marked specimen over 24 km downwind from its

point of release 2 nights previously. They estimated dispersal distances at 32-40 km.

Longevity is a matter of weeks or months. Bellamy and Reeves (1963:319) re-

ported that some females held at outdoor winter temperatures in nan Joaquin

Valley, California, survived more than 8 months.

In early fall, the female population is composed of 2 distinct physiological types

(Blackmore and Dow, 1962:292; Nelson, 1964:251). The gonoactive group con-

tinues to seek hosts, takes blood meals, and oviposit. It is present in ever diminish-

ing numbers, and by late fall almost no individuals of this category are to be

4 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

found. The females destined to overwinter emerge, mate, feed on plant juices and

convert the carbohydrate mainly to lipid in the progressively larger fat body (Har-

wood and Takata, 1965). These females do not take a blood meal and their ovaries

do not develop past the resting stage. They seek protected shelters and are rela-

tively inactive during the winter. Males do not develop a large fat body and do

not survive the winter.

Terms such as hibernation, diapause, reproductive diapause, and facultative dia-

pause have been used to describe the winter condition of tarsalis females. Arrest

of development in mosquitoes may occur in the egg, larval or adult stage, de-

pending on the species (Clements, 1963:220-237). When environment unfavorable

for development, such as low temperature, acts directly on an insect and produces

torpor only for the duration of the stimulus, it is usually called ‘“‘quiescence”’ (Lees,

1955:3). “Diapause” is less dependent on direct influence of unfavorable environ-

ment. When development is interrupted in every generation, regardless of external

conditions, the term “obligatory diapause”’ is used. Initiation of diapause may be

by indirect effect of the environment, such as decreasing photoperiod, so that

some generations do not diapause but others do. This is known as “facultative

diapause” (Lees, 1955:5), and Bellamy and Reeves (1963:321) use this term for

tarsalis. Harwood and Halfhill (1964) and Harwood and Takata (1965) have dem-

onstrated the importance of photoperiod in initiating diapause in tarsalis, but low

temperature reinforces the effect of shortened day length.

“Hibernation” is often used for the winter reproductive diapause of adult mos-

quitoes (e.g. Bennington, Sooter and Baer, 1958:299). It is very different from

the total arrest of all developmental processes often seen in diapausing embryos

or immatures of many insects, where a reactivation period is usually required be-

fore cessation of diapause, such as exposure for several weeks to low tempera-

tures, followed by a return to temperature favorable for development. C. tarsalis

females are moderately active in the winter, unless the temperature is much be-

low freezing (Rush, 1962:180; Bellamy and Reeves, 1963:321) and will feed when

transferred to “‘room’”’ temperature (Rush, 1962:179).

After 2-6 months, depending on the area and the severity of the winter, the

overwintered, inseminated, nulliparous females emerge from their shelters, imbibe

plant juices, seek a vertebrate host, and take their first blood meal. Early in the

season, many females may be attracted to hosts, but the rate of biting is low until

the fat body is largely exhausted (Rush and Tempelis, 1967:309). Because of the

low ambient temperature, blood digestion and ovarian development are much slow-

er during the first gonotrophic cycle of the overwintered females than later in

the season (Rush, Kennedy and Eklund, 1963a:261).

VECTOR RELATIONSHIPS

C. tarsalis is the most important vector of western equine encephalomyelitis

(WEE) and St. Louis encephalitis (SLE) in the western United States. Although

WEE is usually not fatal to man, the epidemic of 1952 in the Central Valley of

California resulted in many human deaths (Longshore et al., 1956:84).

WEE was first isolated from tarsalis by Hammon, Reeves, Brookman et al. (1941),

and numerous subsequent studies have elucidated its epidemiology. The primary

cycle involves tarsalis as the vector and wild birds, especially passerines, and do-

Nelson: Winter Biology of Culex tarsalis 5

mestic fowl as reservoir hosts (Hammon, Reeves and Sather, 1951). Infected mam-

mals, including horses and humans, apparently do not circulate virus in high enough

titer to infect mosquitoes, and therefore they are a dead end in the transmission

cycle,

After a female mosquito feeds on an infected host, WEE virus invades the mid-

gut cells and replicates. Subsequently it invades almost every other tissue of the

body, especially the salivary glands. Although the virus is found in the ovaries

of tarsalis (Thomas, 1963:159) and has been isolated from males of Culiseta mel-

anura (Stamm et al., 1962:74), transovarian transmission is considered unlikely

(Reeves, 1961:66; Thomas, 1963:164). The extrinsic incubation period in tarsalis

can be completed within as short a time as 4 days and a female can transmit the

virus for the duration of its life (Barnett, 1956; Bellamy et al., 1967; Thomas,

1963). There have been no indications of pathology in mosquitoes infected with

arboviruses (arthropod-borne viruses), except for Semliki Forest virus which causes

cytopathology of salivary glands in Aedes aegypti (Mims et al., 1966; Lam and

Marshall, 1968).

OVERWINTERING OF ARBOVIRUSES

Each winter, many arboviruses, including WEE and SLE, disappear from their

vectors and reservoirs, and for many years workers have attempted to determine

how the viruses survive the cold months (Reeves, 1958; 1961; 1965).

Possibly, resident birds with chronic latent infections overwinter WEE virus, or

migrating birds with short or chronic viremia may reintroduce the virus each spring.

Infections usually last a few days (Hammon, Reeves and Sather, 1951:360; Kissling,

Chamberlain et al., 1957:49), but Reeves, Hutson et al. (1958) recovered WEE

virus from blood and other tissues of several passerine species from 1-10 months

after experimental infection. Lord and Calisher (1970) and Stamm and Newman

(1963) report southward transport of WEE during fall migrations, but Kissling,

Stamm et al. (1957:44) found lower incidence of WEE antibodies in spring mi-

grants than in permanent and winter residents.

Bats have recurrent viremia of SLE virus several weeks after infection (Sulkin,

Allen and Sims, 1966:409). Transplacental transmission of virus occurs (Sulkin,

Sims and Allen, 1964) and virus and antibodies have been isolated from bats vir-

tually every month of the year (Allen et al., 1970).

WEE has been isolated from 3 genera of snakes, Thamnophis (garter snakes),

Pituophis (gopher snakes) and Coluber (racers), and from the frog Rana pipiens

(Gebhardt, Stanton et al., 1964; Burton et al., 1966). In the laboratory, infected

tarsalis females transmit the virus to garter snakes (Gebhardt, Stanton and de St.

Jeor, 1966), and Thomas and Eklund (1960) showed that snakes inoculated with

WEE in the fall could overwinter the virus and transmit it to biting mosquitoes

in the spring. Viremia sometimes recurs more than 5 months after initial infection

(Burton et al., 1966:1031; Gebhardt and Stanton, 1966:30). Since the virus has

been found in the offspring of infected snakes (Gebhardt, Stanton et al., 1964:

173), and both snakes and frogs can be infected orally (Spalatin et al., 1964), many

new transmission pathways are feasible: Snakes or frogs may be bitten by infected

mosquitoes, they may eat infected mosquitoes, snakes may eat infected frogs, and

so on.

The mosquito Culiseta inornata (Williston, 1893), a winter species in Imperial

6 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Valley (Chew and Gunstream, 1970:558), is a laboratory vector of WEE (Hammon

and Reeves, 1943:433) and harbors the virus in the field (Hammon, Reeves, Benner

and Brookman, 1945). However, Washino et al. (1962:270) failed to isolate virus

from 4900 females of this species collected mostly in the fall, winter and spring

in San Joaquin Valley, California.

In northern Colorado, where winters are severe, 1 winter isolation of western

equine encephalomyelitis from tarsalis was made from a pool of females collected

in a mine on December 30, 1953 (Blackmore and Winn, 1956). However, based

on the long and well-defined hibernation of the population in that area, Bennington,

Sooter and Baer (1958:304) concluded that tarsalis was an unlikely candidate to

overwinter the. virus. In central Washington, Rush, Brennan and Eklund (1958:

292) failed to isolate WEE virus from overwintering females collected from late

September to early April, although many isolations were made in the summer.

In eastern Oregon, virus was not found in the cold months, and the earliest iso-

lates from tarsalis were in July in one study (Rush, Kennedy and Eklund, 1963b:

286) and not until August in another (Rush and Tempelis, 1967:313). In 13 years

of attempts in central North Dakota, the earliest WEE isolation was in June, and

in western Idaho, July was the earliest month (Rush, Kennedy and Eklund, 1963a:

260). It is the conclusion of the above workers that tarsalis is not a winter vector

in the northwestern United States.

Although in San Joaquin Valley, California, diapause is less pronounced, iso-

lates of WEE virus have not been made from November 15 to January 15, the

period of minimal blood feeding (Reeves, Bellamy and Scrivani, 1958:82-83). Both

WEE virus (Bellamy et al., 1967) and SLE virus (Bellamy et al., 1968) have been

experimentally overwintered in tarsalis. However, Bellamy and Reeves (1963:322)

and Burdick and Kardos (1963:534) concluded that tarsalis is not an important

overwintering vector of virus because of its winter biology. Nelson (1964:252)

speculated that during mild winters this mosquito may occasionally maintain the

virus cycle, but that it is probably not the principal overwintering agent.

METHODOLOGY AND PRELIMINARY STUDIES

STUDY AREA

The Imperial Valley is in the southern extreme of California, south of the Salton

Sea (fig. 1). Various other studies of tarsalis have been made in the Coachella

Valley, which is northwest of the Salton Sea, and the Central Valley which is

divided into the Sacramento Valley in the north and the San Joaquin Valley in

the south.

The Imperial Valley is an irrigated desert of some 2200 km? in Imperial County.

Water for irrigation of such crops as alfalfa, grains, flax and sugar beets comes

primarily from the Colorado River via the All-American Canal and is distributed

by a large network of canals. The drains flow into either the Alamo River or New

River, both of which empty into the Salton Sea. The Sea is actually an inland

lake over 50 km in length, 18 km in width, and approximately 70 m below sea

level. It was formed in 1905 when the Colorado River flooded and filled the dry

sink. Since water leaves the Salton Sea only by evaporation, the salinity has in-

creased to approximately that of sea water (deStanley, 1966).

The Imperial Waterfowl Management Area was established by the State of Cali-

Nelson: Winter Biology of Culex tarsalis i

fornia Department of Fish and Game primarily to prevent crop depredation on

surrounding farms by migratory waterfowl. It also provides sanctuary for an abun-

dance of wildlife, although hunting is permitted during certain seasons. The Wister

Unit of the Area (fig. 2) is on the east shore of the Salton Sea, about 11 km

northwest of Niland on State Highway 111. It is about 2 x 6 km, divided into

sections by dirt roads, dikes, and canals for irrigation of the marshes. Most irri-

gation is done during the fall and winter to provide wintering habitat for migra-

tory waterfowl from the north. The 3 main crops, barley, wild millet and alkali

bulrush, are harvested each year by the ducks and geese. Cattails are plentiful

in the ponds and tamarisk occurs along the borders and in the drier areas with

mesquite.

The most common wintering waterfowl are snow goose, Canada goose, pintail

duck, green-winged teal and American widgeon. Other common birds are Brewer’s

blackbird, red-winged blackbird, yellow-headed blackbird, starling, house finch,

mourning dove, ground dove, pheasant, Gambel’s quail, nighthawk, killdeer, black-

necked stilt, avocet and other small shore birds. Common mammals are Audubon

cottontail rabbit, black-tailed hare, striped skunk and raccoon. Gopher snakes and

rattlesnakes are seen occasionally, frogs are abundant, and several freshwater fish

inhabit the ponds, notably Gambusia affinis, an efficient predator of mosquito

larvae.

Culex tarsalis is the most common mosquito at Wister throughout the year.

Culiseta inornata is abundant in the winter. Culex pipiens quinquefasciatus Say,

1823, Psorophora confinnis (Lynch Arribalzaga, 1891), Aedes vexans (Meigen,

1830), and Anopheles franciscanus McCracken, 1904 are common at other times

of the year. Culex erythrothorax Dyar, 1907 and Aedes dorsalis (Meigen, 1830)

are less common.

On the map of Wister (fig. 2), circled numerals mark the sites where the various

kinds of collections were made. Sites 1-5, 7, 9 and 10 were on the border of

duck ponds that were flooded from about August to March. Sites 6 and 11 were

about 0.5 km from the ponds and sites 12 and 13 about 1 km from the ponds.

All sites were within a few meters of canals. Sites 7-9 and 11-13 were barley

fields where waterfowl fed in the winter.

SAMPLING PROCEDURE

The Wister study was divided into 2 phases: a preliminary phase and a defini-

tive phase. The purpose of the preliminary phase, the 11 lunar months from Sep-

tember 27, 1967 until July 22, 1968, was to determine monthly fluctuations in

population density, to gather preliminary data on overwintering activity, and to

collect large numbers of mosquitoes for virus isolation attempts done by Dr. Telford

H. Work and coworkers at the Division of Infectious and Tropical Diseases, School

of Public Health, University of California, Los Angeles. Seven light traps with

CO, attractant and 7 CO, bait can traps were deployed at sites 3-5, 9-11 and

13 (fig. 2) for 3 or 4 consecutive days during the dark phase of the moon each

month. Collection dates are given with the population data in tables 5 and 6.

On November 30, 1968, use of 3 more light traps and CO, traps was initiated

at sites 2, 6 and 7. Data from these sites were not included with those from the

other 7 sites for estimation of population density for the first 11 months. Rou-

tine daytime resting collections from a culvert (site 6) were also initiated in Novem-

a5 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

ber. Sporadic collections from other culverts and from wooden bridges were made

throughout the first 11 months. Occasional larval collections were made.

The definitive phase of the study included the 21 lunar months from August

31, 1968 until April 12, 1970. The emphasis during this period was on the phys-

iological parameters of the population. Again, collections were made each month

for 3 days during the dark phase of the moon. Light trap collections were dis-

continued. CO, traps were deployed at sites 2, 6 and 7, and routine shelter col-

lections were made from a culvert (site 6) and from under 4 wooden bridges (sites

1, 2, 5, 7). Routine larval collections were made.

COLLECTING METHODS |

Light traps with CO, attractant

The CDC (Center for Disease Control, Atlanta, Georgia) miniature light trap

(Sudia and Chamberlain, 1962) is a small fan and light assembly mounted in a

plastic cylinder. A catching net receives the live insects and an aluminum shade

protects them from sun and rain. The trap operates on a 6-volt battery. Photo-

tactic insects are attracted to the light, caught by the suction of the fan and de-

livered into the catching bag. A block of about 1 kg of dry ice was wrapped in

2 sheets of newspaper, tied with string, and hung next to each light trap at about

1.8 m above the ground (Newhouse, et al., 1966). All traps were deployed before

sunset and retrieved shortly after sunrise. Usually, more mosquitoes are attracted

to light traps if there are no competing light sources. Although mosquitoes are

generally more active during a full moon, more of them are attracted to light

traps when there is little or no moonlight (Bidlingmayer, 1967:206). For this

reason, I scheduled each collecting trip of 3 or 4 consecutive days at Wister to

include the day of the new moon for that month.

The attraction of insects to light is not well understood. One view is that in-

sects utilize a near source of light for a mechanism of orientation which is adapted

for distant sources such as the moon (Wigglesworth, 1965:285). While the angle

between the line of sight to the moon and the longitudinal axis of the insect

remains the same, the insect will fly in a constant compass direction. However,

when an insect orients in this manner to a near light source, ““The angle of inci-

dence of light can be kept constant only if the insect continually turns towards

the source. It will thus move along a logarithmic spiral ending at the light itself...”

(Wigglesworth, 1965:285). Another view is expressed by Verheijen (1960). He pro-

poses that animals use light cues of low intensity for orientation at night. A high

intensity artificial light source interferes with normal photic orientation, resulting

in drifts of the animal toward the light source. Attraction to light varies with

species, sex, age and other factors (Southwood, 1966:174-178, 200-201). In my

study, light traps were used with CO, attractant, which further complicates inter-

pretation of physiological data from mosquitoes collected by this method. For

these reasons, use of light traps was discontinued after the preliminary phase of

the study. Physiological measurements were made only from females collected in

CO, traps and shelters.

CO, traps

The CO, traps (Bellamy and Reeves, 1952) were made from cylindrical “‘lard

cans’ (36 cm x 30 cm diam). A circular hole (23 cm diam) was cut in the lid

and bottom of each can and replaced by an inwardly pointing, hardware cloth

Nelson: Winter Biology of Culex tarsalis 2

cone. A 2 cm circular opening was left in the apex of each cone. The cans were

painted dark red inside and out, and a |-liter capacity, bottomless, polyethylene

cup was fastened to the top of each can on the inside with a strip of sheet metal.

The dry ice, prepared in the same manner as for the light traps, was inserted in

the cup. Mosquitoes enter the traps through the hole in either cone and are un-

able to find their way out again. Each trap was suspended from a tree or bush

at about 1.5 m above the ground.

Apparently, carbon dioxide activates hee -seeking females which then fly up-

wind to locate the source (Clements, 1963:276). Reeves (1953) found that 3 spec-

ies of mosquitoes were attracted to CO, released from large “‘stable’’ traps at rates

comparable to those of their preferred hosts. C. tarsalis, which feeds on a variety

of vertebrates, was attracted to CO, released at rates similar to those of chicken,

man and horse (or cow). More females were attracted at the higher rate of CO,

release. Although females of tarsalis usually do not bite in the daytime, many

attacked the collector when he entered the traps, and mosquitoes were observed

probing at the rubber tubing where the gas was released. Of over 8000 adults

collected, only 22 were males. In my study only 3 males and 1 engorged female

were collected in 444 trap-nights in which 6112 nonengorged females were col-

lected. Therefore, I rejected the possibility that the mosquitoes might be attracted

to the traps as daytime resting shelters and considered that the mosquitoes col-

lected in CO, traps were seeking hosts. (Although 58 gravid females, each with

a very few developing eggs, were collected in the CO, traps, I believe that they

were also searching for a blood source, p. 18.)

Shelters

A mechanical aspirator (Husbands, 1958) was used to collect mosquitoes from

daytime resting shelters. This device was made from a battery-powered vacuum

clothes brush that I fitted with a collecting tube (38 cm x 1.27 cm diam) and

interchangeable clear plastic cartridges (10 cm x 2.5 cm diam) that held the mos-

quitoes from each collection. Collections were made at about midday from a ce-

ment culvert and 4 wooden bridges. The culvert (fig. 3) consistently contained

the largest numbers of mosquitoes. It was a tubular structure (ca. 15 m x 1 m

diam) under State Highway 111. It had running or standing water 1-15 cm deep

at all times. Mosquitoes were usually distributed about evenly throughout the mid-

dle 2/3 of its length. The walls were smooth, and nearly every mosquito could

be collected.

The 4 wooden bridges (sites 1, 2, 5, 7) crossed canals at 0.8 km intervals on

a dirt road running north to south (fig. 4). Each bridge was supported by wooden

beams about 1.5 m above the surface of the slowly moving water in the canal.

Mosquitoes rested on the horizontal undersurface of the bridges, on the vertical

side of the beams, and on the sloping earth banks. During the first 11 months

of the study, all mosquitoes that could be found were collected. However, many

parts of the bridges were inaccessible, and even with a flashlight it was difficult

to see mosquitoes against the darkly colored, rough wooden and earth surfaces.

In August 1968, five 3 gal. (11.4 liter), cylindrical, dark red ice cream cartons

were stapled to the southern, interior vertical surface of each bridge between the

beams (fig. 5). Collections thereafter were made only from these containers. The

red cartons were also placed in various other shaded sites at Wister, but almost

no mosquitoes were attracted to them.

10 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Larval collections

Each month, a systematic search for breeding sites was made. A 1-pint (0.473

liter) sauce pan attached to a wooden pole served as the sampling tool. If a breed-

ing site was too shallow for the conventional dipper, a pipette was used to re-

move all larvae and the mean number of larvae per 0.1 m” of water surface was

determined. At site 8 (fig. 8), a sewage oxidation pond, routine collections were

made from September, 1969 until March,1970. One dipper of water per 0.1 m?

of water surface was taken from an area of 6 m’ of the pond (fig. 9).

SOURCES OF ERROR

There are at least 2 potential sources of error in the data gathered by the meth-

ods described above: The first arises from the collecting procedure; the second is

introduced by the behavior of the mosquitoes.

Error due to the sampling procedure

The sampling procedure may have been inadequate to describe accurately the

Wister population: Collections were made during a very short period each month,

samples were not random, and in some months very few mosquitoes were collected.

The monthly collections of 3 or 4 consecutive days were assumed to be repre-

sentative of the population for the entire lunar month. This probably was not the

case when some characteristic of the population was changing rapidly. Temporary

aberrant weather surely made some collecting periods very atypical for the month.

Biweekly or weekly collections would have been better, and daily collections would

have been best, but impractical. In the graphic representations of the various pa-

rameters of the population, a line connects the values for each pair of consecutive

monthly collecting periods. In this way, seasonal trends can be seen more easily

and data from 2 or more collecting methods can be compared on the same graph.

However, it must be remembered that each point is an average for 3 or 4 days,

not an entire month. 3

Since samples were taken at the same sites every month, they were not random.

The frequency distribution of numbers of mosquitoes per trap per night (trap-

night) was not normal. The monthly population means were sometimes influenced

by very large or very small numbers of mosquitoes captured at | or 2 sites. Stan-

dard errors are given (tables 5, 6) to indicate the variability in the numbers of

mosquitoes captured, but the standard error cannot be used for statistical tests

of differences among the means.

In some months, only a few females were examined for a particular attribute

(e.g. fat body development in CO, traps and shelters). When sample size (N) was

very small ((10), the point on the graph representing the percentage of females

with the attribute is bracketed. For all points sample size is given above the graph.

Bias due to the behavior of the mosquitoes

The behavior of tarsalis may bias the estimation of physiological parameters

in 2 ways: Females may prefer more secluded shelters in the winter, or most of

them may disperse to the foothills.

Shelter preference

Overwintering females of tarsalis have been found in a variety of shelters. In

northern Utah (Dow et al., 1956), northern Colorado (Blackmore and Winn, 1956;

Nelson: Winter Biology of Culex tarsalis 11

Mail and McHugh, 1961) and western Nebraska (Keener, 1952), they are common-

ly found in mines and cellars during the coldest months, but some workers are

of the opinion that these do not constitute the successful overwintering popula-

tion. Dow et al. (1956) found that females disappeared from mines in late winter.

The temperature in the mines was about 10° C and the mosquitoes remained ac-

tive, probably exhausting their energy reserves prematurely. Bennington, Blackmore

and Sooter (1958:298) collected 2 overwintering females from rodent burrows

where the temperature was below 10° C and noted a correspondence of spring

emergence with soil temperature inversion. Mail and McHugh (1961) brought over-

wintering females into the laboratory and found that survival was longest at high

humidity and at a temperature of freezing or slightly below. In southern Alberta,

Shemanchuk (1965) found occasional wintering females in large animal burrows

where the temperature was considerably higher than the ambient temperature but

still dropped below freezing nearly every day. He, too, noted a correspondence

of spring emergence and soil temperature inversion. In the summer, Harwood and

Halfhill (1960) found females and males in vegetation, rock fissures, and animal

burrows, but winter females were found almost exclusively in rock piles and talus

slopes in central Washington (Harwood, 1962; Rush et al., 1958) and in eastern

Oregon (Rush, 1962; Rush et al., 1963b). Harwood (1962) thought that the colder

and more stable northfacing talus slopes were most favorable for overwintering

adults by reducing activity and energy utilization.

Some workers in Central Valley, California, believe that tarsalis prefers “natu-

ral’ winter shelters to “artificial” ones. Probably, a more useful distinction would

be based on the degree of protection from wind and precipitation, and on regimes

of temperature, humidity and light intensity. The man-made structures in question

are usually more exposed. Mortenson (1953) collected overwintering female tarsalis

from such natural sites as tree stumps, tree roots, a tree hole, hollow logs, natural

wood piles, brush piles, animal burrows, a woodrat’s nest, and a rock ledge in

San Joaquin Valley, California. He stated that this species was less abundant in

‘artificial’? shelters. Loomis and Green (1955) continued Mortenson’s study and

concluded that tarsalis females preferred ‘‘natural’? resting shelters to such arti-

ficial sites as chicken houses, bridges, and cellers, but their conclusion was based

on the percentage of tarsalis of all mosquitoes of various species collected rather

than on actual numbers, so that preferences of the other species may have biased

the results. Ryckman and Arakawa (1952) found tarsalis adults in woodrats’ nests

in late November and early December in Riverside County, but did not find them

in all other habitats searched. Kliewer et al. (1969) in Fresno County, California,

observed a decrease in the number of tarsalis in culverts in winter after an earlier

increase, and thought this might be due to a sudden avoidance of these resting

sites. They noticed an apparent preference for piles of rocks. Bailey (1965:109)

states that many overwintering females are found in man-made structures, but that

they prefer natural shelters in the Sacramento Valley.

At Wister, there may well be a winter preference of the females for sites other

than cement culverts and wooden bridges. In fact, this may be the explanation

for an apparent decrease in the population at these sites in the fall (fig. 15). How-

ever, there is no evidence that the winter biology of a hypothetical, inaccessible,

subterranean population of females would be different than that of the females

in the more open shelters. The winter studies of tarsalis in the San Joaquin Valley

of Bellamy and Reeves (1963), Burdick and Kardos (1963), Kliewer et al. (1969)

and Nelson (1964) were all based on collections from such man-made shelters as

12 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

culverts, bridges, sheds and wooden boxes, and a definite period of winter dia-

pause was noted. Since the Wister study was based on collections from the same

kinds of shelters, the data can be compared to those from the San Joaquin Valley.

Dispersal

Some workers have indicated that in the fall, tarsalis disperses from the valleys

to pass the winter in the surrounding foothills. In the Sacramento Valley, Cali-

fornia, Bailey et al. (1965:109) concluded that there was a definite fall move-

ment into the foothills, especially to the hills downwind from the valley. This

dispersal occurred at the same time that the rice fields, the major breeding source,

were drained. Abell (1959) studied mosquito populations in the foothills of Fresno

County, California, at an intermittent stream that offered plentiful breeding sites

from late fall through spring. Twice as many adults were attracted to light traps

from mid-October to mid-November as in July. As breeding did not occur in the

foothill area in the fall, the adults probably had dispersed there. In a 5-year study

in an adjacent canyon, Kliewer et al. (1969:16-17) did not find farsalis adults

in any numbers in culverts until late fall. They were abundant throughout the

winter. In the valley below, light trap catches were highest in the summer. Since

teneral (recently emerged) adults were absent throughout the spring, summer and

early fall in the hills, the population probably did not originate there. Indications

were that in the autumn the mosquitoes dispersed from the valley to the foot-

hills where they overwintered. In January, they began to feed and returned to

the valley without breeding significantly in the foothills.

In the San Joaquin Valley, both the valley and foothill populations of tarsalis

experience a well defined winter diapause. Therefore, there is no reason to believe

that the females remaining at Wister in the winter may be physiologically different

from those that may have dispersed to the foothills.

PHYSIOLOGICAL AGE-GRADING

CRITERIA

Females were dissected and the following attributes related to their winter bi-

ology and physiological age were recorded: external condition, insemination, mus-

cle remnants, meconium, fat body development, presence of blood, ovarian de-

velopment, ovarian tracheation and follicular relics.

As a mosquito ages in the field, the external scales and hairs on the body,

legs and wings are rubbed off. I divided the degree of external wear into 3 cate-

gories, based on the characters used in taxonomic keys to identify tarsalis females:

(1) Recently emerged: Inverted dark-scaled “‘V’’s on venter of each abdominal seg-

ment complete; white tarsal bands complete (Carpenter and LaCasse, 1955:269).

(2) Intermediate: Inverted ‘‘V’’s unclear; white tarsal bands complete. (3) Rubbed:

Inverted “‘V’’s nonexistent; white tarsal bands unclear; white scales present on in-

ner margin of antennal torus, distinguishing tarsalis from Culex peus Speiser, 1904

and other species of Culex (Chaniotis and IItis, 1960).

A female mates once, soon after emergence from the pupa, and the sperm stored

in the spermathecae are used to fertilize all the egg batches she produces. Usually,

only recently emerged females are noninseminated. Diapausing females are nearly

all inseminated.

Nelson: Winter Biology of Culex tarsalis 13

A meconium, a mass of greenish-brown semisolid substance in the midgut, oc-

curs only in recently emerged adults, as do certain abdominal muscle tissue rem-

nants, but the time required for the disappearance of these 2 physiological age

indicators varies considerably with temperature (Rosay, 1961:528).

The fat body of mosquitoes is not a single organ, but a loose meshwork of

lobes dispersed throughout the abdomen. It functions as a food storage organ and

as an excretory organ where uric acid accumulates. In mosquito larvae, it contains

lipid, glycogen and protein, but protein is never deposited in the adult fat body

(Clements, 1963:49-51). The glycogen is used mainly for activities such as flying.

The lipids appear to be more important for hibernation (Buxton, 1935; Schaefer

and Washino, 1969). In diapausing tarsalis females, the fat body becomes large

and compact in winter and is largely exhausted by spring. Bullock et al. (1959:

184-185) recognized 4 stages of fat body development in Culex tritaeniorhynchus

Giles, 1901. Their classification was adapted to tarsalis by Burdick and Kardos

(1963:530). I modified the latter classification for my study: Class 0. — No fat

cells seen. Class 1. — Fat cells around spermathecae only. Class 2. — Traces of

fat in the abdomen, anterior to the ovaries. Class 3. — Abdomen filled with fat

body, but not distended. Class 4. — Abdomen distended by fat body. Classes 3

and 4 of my classification are approximately the same as Classes 2 and 3, respec-

tively, of Burdick and Kardos (1963:530).

Blood digestion and ovarian development occur simultaneously. The adjective

“engorged”’ is used to describe females with any amount of blood in the midgut.

When the blood has been completely digested, the eggs are usually in Stage IV

(see below), and the female is “‘gravid’”’ until the eggs are deposited. Females with

neither blood nor developing eggs are ‘“‘empty”’.

The reproductive system of tarsalis females consists of 2 ovaries and 2 lateral

oviducts that join to form a common oviduct. The 2 ovaries contain a variable

number of ovarioles (ca. 300). Each ovariole consists of a growth zone anteriorly

(germarium) and usually 2 follicles posteriorly. A follicle contains an oocyte and

7 nurse cells. Only 1 egg at a time develops in each ovariole, and the eggs of

most ovarioles develop in synchrony after a blood meal. The fully developed eggs

pass through the lateral oviduct, are fertilized from the 3 spermathecae attached

to the common oviduct, and are deposited in a raft on the water surface.

Christophers (1911) divided the period of ovarian development into 5 phases

(Stages I-V). Mer (1936) added Stage N. The following is a brief summary of

the classification as I have adapted it to tarsalis: Stage N. — The 8 cells of the

spherical follicle are undifferentiated. Stage I. — The oocyte becomes clearly dif-

ferentiated from the 7 nurse cells. Stage II. — Yolk granules appear around the

oocyte nucleus. Stage III. — Yolk granules obscure the oocyte nucleus and begin

to fill the follicle; the follicle becomes broadly oval. Stage IV. — The oocyte fills

nearly the entire follicle and begins to elongate. Stage V. — The egg becomes

fully elongate; the micropyle cup becomes distinct. Stage N is seen only in nulli-

parous females. Stage I is generally thought to be the resting stage for culicine

mosquitoes, but Rosay (1969:610) found that either Stage I or Stage II could

be the resting stage for some culicine species before their first blood meal, and

some anautogenous farsalis females developed Stage II eggs before a subsequent

blood meal.

The tightly coiled ‘‘skeins’ of the ovarian tracheoles of nulliparous (i.e. no his-

tory of oviposition) female mosquitoes are irreversibly stretched during develop-

ment of the ova in the first gonotrophic cycle (Detinova, 1945; 1962:48-51). After

14 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Oviposition, the ovary returns to approximately its original size, but the uncoiled

tracheoles become visible if the ovary is allowed to dry on a microscope slide.

This criterion of parity has been evaluated with farsalis females of known history

and applied successfully to field populations, but there is generally a portion of

the ovaries that either cannot be read or is intermediate between the normal nulli-

parous and parous conditions (Blackmore and Dow, 1962; Burdick and Kardos,

1963; Kardos and Bellamy, 1961; Nelson, 1964, 1966).

In a like manner, the stomach tracheation has been used to distinguish blood-

fed from non-blood-fed females (Detinova, 1962:51-53). Midgut tracheae of nulli-

parous females form a series of tight bands with coiled tracheoles. After diges-

tion of a blood meal, the tracheae are stretched and the tracheoles are uncoiled.

Plant juices do not stretch the stomach tracheae because they first enter the crop

and only gradually pass to the stomach. Detinova (1962:52) reports that this cri-

terion is reliable for distinguishing parous from nulliparous females of Culex. pi-

piens, but I found that it was inapplicable to tarsalis females from a laboratory

colony.

In many species of mosquitoes and other haematophagous Diptera, after each

Oviposition a “follicular relic’? remains, which is seen as a dilatation on the pedi-

cel of the ovariole (Detinova, 1962:25-29; 1968). Polovodova (1949) was the first

to use this as a means of determining parity in mosquitoes (with Anopheles), but

the method was not generally known to the non-Russian speaking world until a

1958 review by Gillies. The Polovodova technique proved to be superior to the

ovarian tracheation method to distinguish parous from nulliparous female tarsalis

in a laboratory colony and in the field (Nelson, 1966), but Nelson (1964:243)

was unable to determine the number of ovipositions with this technique. Rosay

(1969:608) commonly found fewer dilatations than ovipositions in tarsalis females

of known history, and she reported dilatations in specimens that had never ovi-

posited.

DISSECTIONS

In the laboratory, field-collected mosquitoes were immobilized by chloroform,

sorted to species and sex, and counted. Each tarsalis female was placed in a glass

“embedding button” (5 mm deep, 20 mm inside diam) with 2 drops of 0.675%

NaCl. The buttons were stacked to retain moisture. Identification of each female

was verified under 20X of a Bausch and Lomb stereoscopic dissecting microscope,

and the external condition was recorded. The mosquito was then oriented ventral

side uppermost and head to the left, and held by the thorax with a pair of fine

forceps. A minuten pin dissecting needle was inserted between the cerci, and by

slight downward pressure the 3 hard, brown, oval spermathecae and their ducts

and surrounding tissue were withdrawn. In most cases, active sperm could be seen

without crushing the spermathecae. In doubtful cases the spermathecae were trans-

ferred to a slide with a drop of saline solution and examined at 100X with a com-

pound microscope.

The points of a second pair of fine forceps were then inserted into the lateral

portions of the membrane between the 6th and 7th abdominal segments, and drawn

to the right, freeing the gut and ovaries in most cases. The remaining portion

of the abdomen was teased open and the degree of fat body development and

the presence or absence of muscle remnants were recorded. The midgut was checked

for the presence of the meconium and for the presence and degree of digestion

of blood.

Nelson: Winter Biology of Culex tarsalis 15

After the ovaries were separated, 1 was placed on a microscope slide marked

with a grid to facilitate counting of developing ova. With minuten pins, the ovar-

ian sheath was removed, and the ovarioles separated. A record was made of Chris-

tophers’ stage of ovarian development and the presence or absence of dilatations

on the pedicel. Two rows of 5 circles were marked on a microscope slide with a

grease pencil, If the oocytes were not obscured by yolk (Stages I and II), the second

ovary was placed in a drop of distilled water in one of the circles on the slide.

When the ovaries had dried, the presence or absence of skeins could be determined

at a magnification of 100X.

SOURCES OF ERROR

Generally, for each blood meal taken by a female mosquito a complement of

eggs develops, and eggs will not develop without a blood meal. A gravid or parous

female is therefore considered to have a history of blood feeding, and a nulli-

parous female cannot have taken any blood. However, females of some popula-

tions of mosquitoes do not always develop eggs after a blood meal (gonotrophic

dissociation, gonotrophic discordance) and others develop eggs without blood (auto-

geny).

Gonotrophic dissociation

““Gonotrophic dissociation” (Swellengrebel, 1929:1378) occurs in several spec-

ies of Anopheles when fat body development rather than egg maturation follows

a blood meal in overwintering females (Guelmino, 1951). In the same genus, some

species are known to require more than 1 blood meal to initiate egg development.

Rao (1947:44) called this phenomenon “gonotrophic discordance’. Females that

have blood-fed but have not developed eggs are “‘pre-gravids’’ (Gillies, 1954:59).

Eldridge (1968) has reported that low temperature stimulates gonotrophic dissoc-

iation in Culex pipiens and suggests that it may occur in f¢arsalis as well, but it

has not been observed in several studies (e.g. Bellamy and Reeves, 1963:322).

Autogeny

The adjective “‘autogene”’ was coined by Roubaud (1929) to describe females

of a population of Culex pipiens that produced a small raft of eggs without feed-

ing. ““Autogeny”’ is currently used for the phenomenon of the development of

the ovarioles beyond the resting stage without a blood meal (Spielman, 1957:404).

The source for ovarian development comes from reserves accumulated during the

larval stage with the result that only 1 complement of eggs is produced auto-

genously. Autogeny in a colony of tarsalis was first reported by Bellamy and Kardos

(1958) and by Chao (1958) in a subcolony of the same San Joaquin Valley strain.

Moore (1963) found high levels of autogeny in field populations in Sacramento

Valley, California.

If the percentage of gravid or parous females is to be used as an estimate of

blood feeding, this estimate will be too high if the population is autogenous. The

complications in the interpretation of data on gonoactivity and parity in farsalis

due to autogeny have been discussed by various workers (Burdick and Kardos,

1963:528; Kardos and Bellamy, 1961:450; Kliewer et al., 1969:17; Nelson, 1966:

11). In order to determine the degree of error produced by autogeny, I attempted

to estimate its extent in the Wister populations. The following experiments and

observations were of a preliminary nature. Only those aspects of autogeny were

studied that related to physiological age-grading.

16 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Autogeny is a somewhat elusive phenomenon to investigate because of the ap-

parent interplay among hereditary and environmental factors. It is genetically de-

termined (Spielman, 1957), but its expression in tarsalis depends on larval nutri-

tion, photoperiod and temperature (Chaniotis, 1960; Harwood, 1967; Kardos, 1959).

Therefore, these 3 factors were kept constant for all laboratory colonies.

Several autogenous colonies of tarsalis were established from immatures and

adults collected from Wister and vicinity. When workers first attempted to colo-

nize tarsalis, they were successful only after subjecting the mosquitoes to a sim-

ulated twilight in a large cage (Brennan and Harwood, 1953). I was able to colo-

nize the mosquitoes from Wister in a 1 ft (0.028 m°) cage without simulated

twilight. A photoperiod of 14 hr. light and 10 hr. dark was maintained. Temper-

ature in the colony room fluctuated between 24° C and 27° C and relative hu-

midity varied from 53% to 58%. To increase humidity in the cages, I covered

them with plastic bags. Moist raisins were provided as a carbohydrate source. Lar-

vae, reared in 2-liter white enamel pans, were fed either Fleischmann’s yeast or

laboratory chow (Purina Pigeon Checkers) but this diet was not standardized by

weight. The water was aerated to prevent the formation of surface scum.

Three separate criteria were used for detecting autogeny: (1) development of

eggs without a blood meal, (2) oviposition without a blood meal, (3) occurrence

of Stage III eggs in nonblooded females. Percentage of females that develop eggs

without a blood meal was the best index of autogeny. Females with Stage III-V

eggs 5-10 days after emergence from pupae were considered autogenous. The sec-

ond index, percentage of females ovipositing without a blood meal, was used as

Table 1. — Autogeny in Culex tarsalis collected as larvae and pupae at Wister and vicinity

and reared in the laboratory. 1969.

NON-BLOOD-FED BLOOD-FED

Index of Autogeny Mean No. Eggs per ? Mean No. Eggs per °

. E z Bak 2a

3% ae Ge 6 Bas eS See a

— e 2 oF 2 & 2 a re 2

ES a 2) a B25 Z, a 5 iz = 5 7,

May 14 fF, 24 436 (162) 3

Jun Ji P 83 11

F, oe) 55

F, 902!) 28453 a. 48

Jul 8 Pp S$. < 9057 57 14

Sep 11 : 47 a7 119 10

eee 10 ee 10

Oct 11 ; 17 30. f49) 5

ee 716 49 7 31

nee 9 33 = (66) 3

8.11,15- - a3. 74 23 96 96 ie: See 4 pee mee oe ee

Nov 11 : 0 12

8 - ay) 42 96 18

Nelson: Winter Biology of Culex tarsalis 17

a measure of autogeny when females were not dissected. This index tended to be

lower than the first, because some females developed eggs but did not oviposit.

Also, the number of eggs oviposited tended to be lower, because some of the

eggs were occasionally retained in the ovaries. The third index of autogeny, per-

centage of Stage III females without blood in the midgut, was used on field-col-

lected females. In the laboratory, females with anautogenously developed ovaries

always had blood in the midgut until after Stage III. Therefore, nonblooded fe-

males from the field with Stage III ovaries were classified as autogenous. This

index was lower than the first 2 because many females that oviposit autogenously

subsequently blood feed and develop eggs anautogenously.

In June 1969, I found several egg rafts in a colony of non-blood-fed mosquitoes

that had been reared from larvae collected at site 12. This colony was maintained

for 6 generations without a blood meal before I discontinued it 5 months later.

(Each new generation was transferred to a separate cage.) The colony from site

14 was allowed to feed during the parent generation, but thereafter it reproduced

autogenously for 6 generations (6 months) before it was discontinued. The colony

from site 8 died after 12 generations (10 months) without a blood meal.

Tables 1 and 2 show the preliminary data from experiments and field observa-

tions of autogeny: Table 1 for females that were collected as larvae or pupae in the

field and table 2 for adult collections. For colonies, the generation (P, F,, F,,

etc.) is designated. For each collection, the percentage of autogenous females is

shown beneath the index used for its determination. Data based on (10 observa-

Table 2. — Autogeny in Culex tarsalis collected as adults in CO, traps and shelters at Wister.

1969.

NON-BLOOD-FED BLOOD-FED

Index of Autogeny Mean No. Eggs per? Mean No. Eggs per &

3 § Choma aaa: aoe

ae = = eae a Qu a,

Es eee Bs By eke Ban saan S a2 8 See,

CO, trap

Jun ae ig 20 i

F, 13 76

12 228 2 pe

Jul - 20 61 22 11

Sep - 20 29 QT) 4

Oct - 22 41

Shelter

All* - 30 155 58 10s. (419) 9

Jan 5 . 138 1]

(1968)

* Females collected from sites 1,2,5-7. September 1967-April 1970.

18 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

tions are bracketed. Two site numbers are given that are not shown in the map

of Wister (fig. 2): Site 14 was a grassy roadside ditch (ca. 60 cm x 300 cm x 12

cm deep), ca. 8 km south of Wister on State Highway 111. Site 15 was a foul,

grassy ditch (ca. 1 m x 2 m x 0.5 m deep), ca. 40 km south of Wister on State

Highway 86 at the Keystone Avenue intersection.

Consistently, fewer eggs per female were developed autogenously than anauto-

genously. From a combined collection from sites 8, 11 and 15 in October, 16

of 39 females were allowed to feed on a pigeon, and all females were isolated

in 9-dram (16 g) plastic vials. The bottom 1 cm of each vial was filled with water,

and a moist raisin was placed on the cheesecloth top. Eighty-three percent of the

non-blood-fed individuals developed eggs autogenously, and 74% of them ovipos-

ited. The mean number of eggs that developed for 19 females was 96 as opposed

to 212 for the blood-fed individuals. The difference between the means was sig-

nificant (p = (0.001). However, the range overlapped: As many as 136 eggs devel-

oped autogenously in 1 female and as few as 118 developed anautogenously. In

other colonies, the number of eggs oviposited autogenously ranged up to 186 (site

14, F,), and 1 blood-fed female developed only 25 eggs (site 11, F;). Females

that took small blood meals usually developed fewer eggs. In the Sacramento Val-

ley, Moore (1963:240) reported that 1 female developed 220 eggs autogenously.

Mean number of autogenously developing eggs was highest in May at 116 and

declined to less than 35 in October. In June, 92% of the females collected as

pupae were autogenous. By October only about 20% were autogenous. Burdick

and Kardos (1963:533), using intermediate ovarian tracheation as the criterion,

concluded that autogeny decreased from September to October in the San Joaquin

Valley.

In August and September 1968, 4 gravid females were collected in CO, traps

at Wister. In 1969, about 20% of the females caught in CO, traps from June to

October were gravid (table 2). With 1 exception, no engorged females were col-

lected in these traps. None of the females with Stage III ovaries had any trace

of blood in the midgut. Therefore, all of the gravid females in CO, traps were

probably developing eggs autogenously. The average number of developing eggs

per female was 22. This is much less than the number of eggs normally produced

by autogenous females. The non-blood-fed females of 2 filial generations developed

approximately the normal number of eggs for autogenous females. It may be

that the field individuals had not stored sufficient reserves during the larval stage

to complete the development of a normal number of eggs autogenously. Ovarian

development began, but when reserves were exhausted an additional source of pro-

tein was sought in the form of a blood meal. An alternative explanation is that

these were females that had taken a very small blood meal and had begun ovarian

development. By Stage III, the blood meal was completely digested, and they were

attracted to CO, in search of a second blood meal. In either case, the gravid fe-

males in CO, traps did not need a blood meal to complete development of their

eggs and to oviposit. In 11 such females, an average of 20 eggs was deposited,

of which most hatched.

Two criteria for distinguishing parous from nulliparous females are less accu-

rate when the population is autogenous: ovarian tracheation and follicular relics.

The degree of stretch of the tracheal skeins is determined by the number of eggs

in the ovaries. The skeins of autogenous-parous females are intermediate between

those of nulliparous and blooded-parous females, because autogenous females de-

velop fewer eggs. Burdick and Kardos (1963:531) examined a large series of labo-

Nelson: Winter Biology of Culex tarsalis 19

ratory-reared tarsalis females (with histories of ovarian activity unknown to them

until after the experiment) in an attempt to separate them into nulliparous, auto-

genous-parous, and blooded-parous categories on the basis of ovarian tracheation.

By assigning all doubtful specimens to the intermediate (autogenous) category, no

nulliparous specimens were incorrectly identified as blooded-parous and only 4%

of the blooded-parous specimens were judged to be nulliparous. However, so many

females in these 2 categories were incorrectly judged to be autogenous-parous, that

only 39% of the nulliparous specimens and 68% of the blooded-parous specimens

were correctly identified. Eighty percent of the autogenous-parous females were

correctly identified. The other 20% were approximately evenly distributed between

the other 2 categories, but of all females identified as autogenous-parous, less than

a third were correctly identified.

Follicular relics occur in both autogenous-parous and blooded-parous females.

Usually more ovarioles of blooded-parous females will have relics because more

eggs are developed, but this criterion cannot be used to distinguish them from

autogenous-parous females because of the overlap in the range of the number of

eggs developed by these 2 groups. Figures 10 and 11 show the frequency distribu-

tion of the number of developing eggs per female from collections at Wister. Based

on my laboratory observations (table 1), I expected the following 3 peaks in egg

frequencies, with a minimum of overlap: retained eggs (ca. 1-10), autogenously

developing eggs (ca. 50-100), and anautogenously developing eggs (ca. 200-250),

However, there was no clear division into these 3 groups: 116 of the 315 females

examined had from 1 to 7 Stage V eggs; 70 of these had 1 egg. These were prob-

ably all females with retained eggs, but the Stage V eggs of females with more

than 7 may also have been retained. The rest of the distribution was continuous

and did not separate into an autogenous and an anautogenous group. Therefore,

the frequency distribution of follicular dilatations is not useful to separate auto-

genous-parous from blood-fed-parous females. However, lack of dilatations is a good

criterion for identifying nulliparous individuals (Nelson, 1966).

In summary, at Wister autogenous-gravid or parous females could not be dis-

tinguished from blood-fed-gravid or parous females with a high level of accuracy.

However, if monthly levels of autogeny in Imperial Valley are similar to those

in Central Valley, which probably is the case, based on my limited data and on

Central Valley studies, then the bias due to autogeny should be the same in both

areas, and the physiological age-grading data can be compared.

ANALYSIS OF DATA

All of the data from each of the 2658 dissections (date, dissection number,

collecting method, site and all the age-grading results) were coded and entered

onto standard IBM punch cards. The following programs were written in Fortran

IV for a Watfor compiler (Blatt, 1968) and were executed on the model IBM 360/91

of the Campus Computing Network of the University of California, Los Angeles:

(1) a print-out of all or any portion of the dissection data in easily readable, non-

coded form, (2) a 2-way classification table of a variable number of categories

to show the frequency of females for each combination of categories (e.g. if the

2 variables were external wear, ‘“‘ext’’, and fat body development, “‘fat’’, the table

showed frequency of females with the combination of categories ext = 1, fat = Q;

ext = 1, fat = 1; ext = 2, fat = 1; etc. for all possible combinations), (3) percent-

20 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

age of females in each of the 6 stages of ovarian development, tabulated by col-

lection method and by month, (4) percentage of females inseminated each month,

tabulated by Christopher’s stage and by collecting method, (5) histogram of the

frequency distribution of the number of developing eggs per female for all months.

Other computations were executed by an Olivetti Underwood Programma 101

desk computer. A prewritten program (Williams, 1968:33) was used to calculate

the monthly mean number of males and females per collecting method and the

standard error of the mean. A t-test (Williams, 1968:188) was employed to de-

tect the degree of difference between the mean number of eggs developed by blood-

fed vs. non-blood-fed females. Short programs were written for computing per-

centages and confidence intervals.

RESULTS AND DISCUSSION

WEATHER

A standard U.S. Weather Bureau Station was established at Wister in May 1967

but was discontinued in May 1968. The closest other station with complete weather

data was operated by the Imperial Irrigation District, about 50 km south of Wister.

Differences in temperature were minimal between the 2 stations for the 13 months

in question. The monthly mean lows averaged less than 1° C lower at Wister,

and the mean highs less than 0.5° C lower. Recorded rainfall was also nearly the

same. Therefore weather data for my entire study were taken from the Imperial

station.

Winters are mild and summers very hot at Imperial, California (fig. 12, table

3). The coldest months were December and January, but temperatures only rarely

Table 3. — Winter and summer temperatures (°C) at Imperial, California.

1967 1968 1969 1970

WINTER is ie ea aes ee

Coldest month : mean 11:7 le oe | 12.8

mean minimum 5.0 3.3 5.6

Coldest 2 weeks: mean minimum 3.9 Supe 2.8

Coldest day : minimum 0.6 -2.8 -1.7

SUMMER

Hottest month : mean 34.4 32.8 35.6

mean maximum 41.7 40.6 42.8

Hottest day : maximum 45.0 45.6 47.2

dropped below 0° C: not at all in winter 1967-1968, 4 days in December 1968

and 6 days in January 1970. The lowest temperature during the study was —2.8°

C for 1 night in December 1968. The mean low temperature for the coldest month

was about 5.3° C for 2 winters and 3.3° C for the winter 1968-1969. Summer

temperature reached 47.2° C in August 1970. Mean maximum temperature for

the hottest month (July or August) exceeded 40° C all 3 summers. Rain occurred

sporadically and infrequently, usually in the winter except for a major storm in

July 1968. Average annual rainfall for all years recorded by the Imperial station

Nelson: Winter Biology of Culex tarsalis Ps

was 72.7 mm. Relative humidity, recorded daily at 7:30 a.m. PST at Wister from

May 1967 to May 1968, ranged from 9% to 98%. Monthly mean relative humidity

was approximately 70% in the winter and 50% the rest of the year.

In most areas outside of California, where the winter biology of tarsalis has

been studied, winters are severe. In southern Alberta, Canada, Shemanchuk (1965:

461) reported an extreme of more than 35° C below zero, and the air temperature

was well below freezing nearly every day from November through March. In a

western Nebraska study, ambient temperature reached a low of —30.6° C (Keener,

1952:209). In North Dakota, temperatures below —17° C persist for 2 weeks or

more, but in central Washington temperatures below —17° C last for only short

periods (Anderson and Harwood, 1966:6). Rush, Brennan and Eklund (1958:291-

292) reported below freezing temperatures all during January.

In the San Joaquin Valley, California, winter temperatures are milder, but still

considerably colder than the irrigated desert of Imperial County, only 2 degrees

of latitude to the south. In Kern County, Bellamy and Reeves (1963:318) reported

average minimum temperatures of well below 5° C for many 2-week periods dur-

ing the winter, and there were biweekly lows below —1° C both winters of Nelson’s

(1964:249) study. Washino and Bellamy (1963:162) reported that temperature fell

below freezing and reported a minimum of —7.8° C for 2 days in January. In

Fresno County, Kliewer (1969) showed minimum temperatures below freezing for

nearly every week during a 2-4 month period for each of 5 years.

POPULATION DENSITY

The numbers of tarsalis adults collected by the 3 methods are listed in table

4. Light traps with CO, attractant, used only during the first 11 collecting per-

Table 4. — Number of adult Culex tarsalis collected by 3 methods at Wister, California.

September 1967 — April 1970.

Number of Mean

Number Collecting Number of Number per

Collected Periods Collections Collection

Light trap with CO, $5235 11 280 119

CO, trap 6115 33 444 14

Shelter 5171 30 366 14

Total 44521

iods (lunar months) of the study, attracted by far the greatest number of adults

per trap per night (trap-night), and no trap failed to capture at least a few adults,

except in occasional cases of mechanical failure. CO, traps and shelters yielded

fewer adults per collection, or sometimes none.

MOBILITY

If adults are not active every day, or if the population is very small, progres-

: Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

sively fewer mosquitoes will be collected at the same site on consecutive days.

At Wister, no effect of the 3 sampling procedures on the population was observed.

Collecting from a particular site one day tended neither to decrease nor increase

the number of mosquitoes collected the next day. Of 134 such comparisons with

light traps, 56% were decreases and 44% were increases. (Instances of no change

from one day to the next were not counted.) For CO, trap collections, 46% of

211 comparisons were decreases, and of 314 comparisons for shelter collections,

44% were decreases. For all 3 methods, decreases and increases for consecutive

collections were about equal, even during the winter months when the population

density was low. It may be inferred that the population of adults was very large

in comparison to the numbers removed by the 3 sampling techniques, and adults

were very mobile all year.

In the northwestern United States and Canada, overwintering tarsalis females

are nearly completely immobilized by the cold (Rush et al., 1958:291); Shemanchuk,

1965:459). Rush (1962:179-180) collected wintering tarsalis females in the field

and subjected them to various temperatures in the laboratory. Although there was

some survival at temperatures as low as —10° C for 24 hours, at —3.3° C only

very slow walking was observed, and true flight was not possible until 2.2° C.

Strong flight of 9 meters or more was not achieved until 4.4° C. In the San Joa-

quin Valley, California, where winter temperatures are less severe, overwintering

females are immobilized only during brief cold periods, and shelters become re-

populated a week after all mosquitoes are collected (Bellamy and Reeves, 1963).

In the Imperial Valley, shelters were repopulated daily, and afternoon tempera-

ture was never low enough to reduce adult flying ability noticeably.

FEMALES

The best data on the winter fluctuations of the female population density were

obtained during the preliminary phase of the project when many samples were

taken every month from September 1967 through July 1968 (fig. 13, table 5).

The light traps and CO, traps showed nearly identical fluctuations, which was

to be expected since CO, attractant was employed with both. Fewest females

per trap-night were found during the late November-early December collecting per-

iod, the only time that the number fell below 10 and was less than the number

of females of Culiseta inornata caught in the same traps. Average monthly mini-

mum temperatures were lowest in December and January during the winter 1967-

1968 (fig. 12) and the coldest 2-week period was January 1-15, but progressively

more females were obtained by all 3 methods after the collecting period of Novem-

ber 30-December 2. Light traps were not used for the remainder of the study and

fewer CO, traps were employed. Supplemental shelter collections were made from

3 wooden bridges, but usually only a few adults were found. During the next

2 winters, the fewest females were found after the coldest months. In the winter

1968-1969, the minimum temperature was in December (fig. 12) but the culvert

showed the lowest population in February and the CO, traps in March (fig. 13,

table 6). The trend in 1969-1970 was not clear since collections were discontinued

in April. During the coldest months of both winters (December 1968, January

1970), the numbers in the culvert shelter increased while those in CO, traps de-

creased from the previous month.

For both summers, there was an apparent depression in population density dur-

ing the hottest month, but collection data were incomplete for the first summer,

and during both summers there were temporary aberrant weather conditions. In

Winter Biology of Culex tarsal

Nelson

a = (UROL oY} JO IOIIO Plepue]s 94} JO 9}¥UTIso peseIqun) *9 x

; : : : 2 € "7 OTT 91 v7 L'6 17-61 Inf

: ; 2 ‘ : L rare vy ET ETT 6°6€ 7Z-OC UNE

€ Et EE CE Lol IZ 6°S O'vZ IZ V'v7 V'TEl L7-S7 ABW

€ SZ 0°6 LS O'6T IZ 6°C L'8 IZ 6'v1 6°€6 67-LTZ dy

“d : O'9T SOI S61 IZ OP OSI IZ SEI 9°96 L7-ST EW

€ 8°S ETI 6b LOI 07 v7 18 IZ ET 9°OL Z WeW-67 994

€ 6°0 Lz Oe O'ST 8Z 70 ct 87 SY 761 O€-L7 URL B96I

€ Bp et (a3 ETT iz a) vl IZ aS SEZ O€-87 99

€ 3 O'1Z SI ie IZ €0 €°0 IZ SE L'6 Z 99-0€ AON

- , : : ; LZ 8" L3 8Z Tp €€0E S-7 AON

; ; : ; ; 07 €°8 8°87 IZ 9°ES 6'09r 67-LTZ deg LOGI

N *9 x *9 x N *9 x N ao x

2 5 é

Vaan) sdey °Q) [OO ypia

sdeq} 7Y8r]

"8961 9UNC—L9O6[ ISNSNY “(9 OTIS)

I9}]9YS WaATNO eB pue ‘sden %QJ ‘7Q_D YIM sdeq} YB] UL paydeT[Oo sywsun] XaIND 1[Npe Jo souepunge A[YyUOW| “Aysuep uoNe[ndog — °¢ aqQuL,

Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

9 ; 0) 9 : 0 : 0 ‘6 : 0 ; 07 CI-II Adv

6 50 a 6 : 0) : lO € ; L'0 ; €0 8-9 Ie]

6 a Or 6 E 1'0 v0 cl € 9°0 07 Le Le 8-9 qo

6 : 10 6 S'0 61 v0 ET € ae €Ol Lo L's G-¢ UeL OL6I

6 90 Wl 6 70 9'0 a) 60 € SZ OE E0 Ly 8-¢ 90

6 [eE vl 6 10 Z'0 Z'0 L'0 € SE] LZ O71 OSI OI-L AON

6 L€ Vel 6 70 Ul S'0 Cl 4 v7 Lu 13 8°61 ET-O1T 90

8 Or vil 6 60 9] 60 9°7 € 601 L’Sv vy O'7L PI-IT das

6 L'0 g'] 6 : 0) pl 70 € : 0 Cl oa 9I-pl snyv

6 797 9°€9 6 : ) S'0 LA € €0 E9 OTT €°69 EI-TT ie

6 0°61 es 8 8°0 ST a gs € Ol OL LO L’v9 OI-pl ung

9 E€ (exe) 6 v0 6'0 60 ve € vl Lv EOI LV 17-81 At

9 alk xs 6 : ) v'0 9°0 € eT C7 | L’8 17-61 idy

9 90 8°0 6 70 70 70 9°0 € €0 L'0 ST L’€ 6I-LI JeW

9 8°0 ST 6 £0 70 €0 8°0 € £0 I ET ET LI-ST 994

6 Gl Ly 6 E0 8°0 ie (G46 € g's Le 79 ESI O7-SI URE 696

9 0) Cl 6 ST ES Ll Ee € [Se E101 9°¢ €°99 vI-TT 99d

€ cI o's Ol 91 (os 6'0 67 € TIP S'6L S°8 Or 97-77 AON

6 ETI Tp cl 07 L’9 6'0 v7 € 0'OL Gal O81 OCP tT AON

6 OTT g°S9 al ET SY 60 vs € 0'6¢ EPOI O'rZ £891 P7-7T dog

9 6'CE 8°SOl 8 Tl Ove rag Cv € Se E61 p77 ey9l Z das-Te 3nV 896

».4 xX ».4

N “9, x N “o x “9, x N “9 x a x

sdeyy “QD sosplig WAND

‘OL61 [IdW—896I 1oquieides ‘sraxpays pue sden %Q) Ut poyoeT]oo symsuny xaIND y[Npe Jo souepunge ATyUOW “Aysuop uoHe[ndog — °9 a1qeL

Nelson: Winter Biology of Culex tarsalis 25

the summer of 1968, light trap collections were greatest in mid-May and progres-

sively diminished until mid-July, but this collecting method was discontinued after

that month. Fewer females were caught in CO, traps in June than May, but this

collection method was not employed in July. By late August, CO, traps attracted

4 times as many as the previous May, and 25 times the June catch. In early July,

unusually heavy rains in the nearby Chocolate Mountains caused the canals at Wister

to overflow, and a large portion of the area was inundated. This may have lowered

the population by flooding the adult resting sites and washing larvae into the Salton

Sea. The subsequent increase in population may be attributed to the increase in

available breeding ponds as the flood waters receded. In the summer of 1969,

a considerable drop in numbers in shelters and CO, traps was evident in mid-

August. Shelter collections a month before and a month after were high. Shortly

before the collecting period in August, there had been a wind storm which may

have dispersed the mosquitoes, but high winds during other months had not no-

ticeably reduced the population in shelters. (However, light trap and bait can catches

were reduced on windy days.)

The annual population curve is bimodal. Mosquitoes were most abundant in

the spring and fall, somewhat less abundant for a short time in the summer and

least abundant in winter. Work, Vanis and Wallace (1969) continued the CO, baited —

light trap collections at Wister after I stopped them in July 1968, and they too

observed spring and fall population peaks in 1968 and 1969.

Since rainfall in the Imperial Valley is insufficient at almost any time of the

year to provide breeding sites, breeding is largely dependent on irrigation prac-

tices. Wister is primarily a management area for migratory waterfowl, and the duck

ponds are flooded from about August until March. Other areas of Imperial Valley

are irrigated in the spring and summer, but the pattern of seasonal abundance

is about the same. Magy (1955) maintained light traps in Imperial Valley at sites

ranging from about 40 to 65 km south of Wister. In 1954, tarsalis females were

most abundant in October, with a smaller population peak during April and June.

Larval collections showed the same bimodal pattern. Chew and Gunstream (1970:

560) found peak populations of tarsalis in light traps in spring and fall in the

Coachella Valley, at a site about 50 km northwest of Wister. In a survey of several

other areas of southern California, including Imperial Valley, only 1 peak was evi-

dent, in the late spring. In all the areas studied, this species was present in light

traps every month of the year, and it was the only species besides Culiseta inornata

that occurred through the winter months in any numbers.

C. tarsalis demonstrates latitudinal differences in seasonality in various parts of

its range from western Canada to southern Mexico. Similar differences are seen

in several other species, notably Culiseta inornata, which is a winter species in

Imperial Valley (Chew and Gunstream, 1970:558), but is most active in the spring

and autumn in the northern United States and in the summer in Canada (Horsfall,

1955:351; Shemanchuk, 1959:908). Probably the difference in the time of opti-

mal temperature is one of the most important factors influencing this seasonality.

In southern Alberta, Canada, light trap catches of tarsalis were highest in July,

but by September they dropped to zero (Shemanchuk, 1959:908). In North Dakota,

tarsalis is inactive sometimes for more than 6 months, but the period of inactivity

in central Washington is usually less (Anderson and Harwood, 1966:6). In Wash-

ington, the number of adults in light traps were low in May and peaked in July

(Hammon, Reeves, Brookman and Gjullin, 1942:281), and by late September vir-

tually all surviving females had migrated to their winter resting sites (Rush, Brennan

26 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

and Eklund, 1958:289). Emergence of the overwintered population began the first

half of March (Rush, Brennan and Eklund, 1958:291; Harwood, 1962:29). In north-

ern Colorado, hibernating females were found in mines at the end of August when

the numbers of females attracted to CO, traps had declined (Blackmore and Dow,

1962:293). The emergence of overwintered females in that state varied from late

March to mid-April (Bennington, Blackmore and Sooter, 1958:298). In western

Nebraska, hibernating females disappeared from a cellar at the end of April, but

no females were caught in light traps until June (Keener, 1952:208).

In the San Joaquin Valley, California, populations also show a midsummer peak

and a winter low, but fall populations are depressed later and spring populations

increase sooner. Collections in light traps (Hayes et al., 1958:221; Reeves, 1970:

3), CO, traps (Hayes et al., 1958:221), and shelters (Hayes et al., 1958:221;

Reeves and Hammon, 1962:176-177) were highest in late July and August. Oc-

casional adults were found throughout the winter in light traps (Hayes et al., 1958:

221), but no mosquitoes were attracted to CO, traps during December and Jan-

uary and sometimes not until March or late April (Hayes et al., 1958:221; Nelson,

1964:247). }

Jenkins (1950:913) noted that most collection records of tarsalis east of the

Mississippi River are in late fall. Snow and Pickard (1956:147-148) found peak

larval abundance in September in the Tennessee Valley, an area where tarsalis is

an uncommon species.

Eads (1965) states that on the Lower Rio Grande Valley of Texas, tarsalis

is a winter mosquito. He presents light trap data for 2 years in which no adults

were collected from June to September, although this was the period of greatest

abundance for many other mosquito species. On close inspection of his data, I

believe that a very definite bimodal pattern is evident for both years. Adults are

most abundant in November, common in December and January, and then increase

again in February of one year and March of the next. The Lower Rio Grande

Valley is semitropical, and the winters are even milder than in Imperial Valley.

The average January temperature was 15.6° C, as opposed to 12.2° C at Imperial,

but the minimum occasionally dropped below freezing. It is difficult to explain

the summer disappearance of tarsalis from the Lower Rio Grande, since the aver-

age July temperature was 28.9° C, compared to 34.4° C in Imperial Valley, and

other species of mosquitoes were abundant. Most of the 760 mm of rainfall oc-

curred in the fall and spring, and standing water was usually more abundant in

the winter months, but potential breeding sites were plentiful in the summer due

to agricultural irrigation.

Figure 14 is a diagrammatic summary of population fluctuations in the major

areas that have been discussed above. C. tarsalis is a summer mosquito to the

north, a fall and spring species in the Imperial Valley, and a winter mosquito

in southern Texas (with a mild midwinter depression). The series cannot be ex-

plained entirely by differences in temperature regimes, since the Lower Rio Grande

Valley summer temperatures are not excessive.

MALES

Although males collected in shelters at Wister were less abundant in winter than

during the fall, they never disappeared (fig. 15, tables 5, 6). During the late fall

and early winter of both years, males were distinctly more numerous than females,

but during the spring and summer months the converse was true.

Throughout most of the range of tarsalis, males do not survive the winter, al-

Nelson: Winter Biology of Culex tarsalis 27

though Harwood (1962:28-29) did collect 1 male in an emergence trap placed

over a talus slope from March 11 to April 29, 1961. Because this collection, and

others of male Anopheles freeborni Aitken, 1939, were made from a north-facing

slope, Harwood (1962:30) speculated that the colder, more stable site was most

favorable to hibernation of adults by conserving their energy reserves. Usually,

the first males seen in the spring are brightly colored members of the first spring

brood, a few weeks after females emerge from their wintering sites (Rush and

Tempelis, 1967:309). In southern Alberta, Shemanchuk (1965:459) found females

in mammal burrows from September to January and April to June, but males

were present in September only. Bennington, Sooter and Baer (1958:300) collected

females under a bridge in northern Colorado in mid-April, but males did not ap-

pear until late May. Both sexes had virtually disappeared by the end of October.

Keener (1952:208) collected females from cellars in western Nebraska from mid-

January until the end of April, but no males were found. In Fresno county, Cali-

fornia, males were absent from culverts in January, February and March (Kliewer

et al., 1969:17), and in Kern County they are absent from shelters for a month

or longer, and do not reappear in numbers until late March or April (Bellamy

and Reeves, 1963:317; Burdick and Kardos, 1963:533; Nelson, 1964:248).

At Wister, the presence of tarsalis males throughout the year is the clearest

evidence of a basic difference in winter biology between that population and those

mentioned above. Since blooded and gravid females and larvae are also present all

winter, there is no distinct “new generation” of adults indicated by the first ap-

pearance of males in the spring. Continuous breeding produces a continuous popu-

lation of males.

The apparent increase in percentage of males in shelters for both winters at

Wister may be due to a change in behavior of the females. Perhaps the majority

of the females disperse from Wister during the cooler months (p. 12) or seek more

secluded resting sites such as animal burrows (p. 10). Bellamy and Reeves (1963:

317,318,321) found a secondary peak of abundance of males in late autumn in

Kern County. They thought that there may be an increased tendency for males

to congregate in shelters at that season or survival may be better at the cooler

autumn temperatures.

IMMATURES

Larvae and pupae were found every month of the year at Wister. Although

much of the area was flooded to provide habitat for migratory water fowl, larvae

were seldom found in the duck ponds (fig. 6). Gambusia affinis (mosquito fish),

an efficient predator of mosquito larvae (Hoy and Reed, 1970), was very abun-

dant in the ponds. The best tarsalis breeding sites were shallow, grassy ditches

(fiz. 7) with recent accumulation of seepage from the ponds or the canals. Spo-

radic breeding occurred at or near almost every site enumerated on fig. 2, espec-

ially sites 1, 6 and 12. Unfortunately, these sites were usually dry after a few

weeks, and therefore breeding levels over time could not be determined for any

one seepage ditch.

Table 7 shows fall and winter collections from site 8, the large sewage oxida-

tion pond which was the only breeding place that contained water all year. One

dip was made for each 0.1 m’ of surface area collected, therefore mean number

of larvae per dip can also be interpreted as mean number per 0.1 m’™ of water

surface. Lesser numbers of larvae were collected in the winter months, but no

trend is readily apparent from the limited data. Frequent changes in the level of

28 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Table 7. — Immatures. Monthly collections of larvae and pupae of Culex tarsalis in the sewage

oxidation pond (site 8) at Wister, California.

Date of Mean number Number

collection of immatures of dips

per dip

1969 Sep 13 0.8* 60

Oct 10 50.0* 60

Nov 9 = 4 60

Dec 7 9.5 63

1970 Jan 5 Of 20

Feb 7 0.2 20

Mar 7 0.8 20

: Estimate.

* Larvae were plentiful in 3 seepage ditches in January.

the pond corresponded to decreases in numbers collected, even from one day to

the next. On January 5, 1970, although plentiful larvae of both Culiseta inornata

and Culex pipiens quinquefasciatus were found in the oxidation pond, only 1 larva

of tarsalis was collected in 20 dips. However, 3 other temporary breeding sites

in the area had larger number of tarsalis larvae that month.

In the northwestern United States, tarsalis larvae are not found in the winter

(Rush, Kennedy and Eklund, 1963a) because females are gonoinactive. In Kern

County, California, immature stages do not disappear completely in the winter,

but they are apparently much less abundant than in Imperial County. Brookman

(1950:85) and Washino and Bellamy (1963) found at least a few immature stages

of tarsalis throughout the winter months, but as the latter authors state, “during

the cold months an exhaustive search was necessary to find any larvae.” In Janu-

ary, only third and fourth instar larvae and pupae were found, and in February,

in 34 potential breeding sites, 7 larvae were found, all of which were first and

and second instar. By March, all larval instars were found, but no pupae. Probably,

the January population represented the last of the developing immatures from adults

that had been gonoactive in the fall, and the February population of early instars

was from females that had overwintered and subsequently became gonoactive. In

my study at Wister, immatures were more numerous, and pupae and all larval

instars were found every winter month.

On January 6, 1970, at 7:00 a.m., I noticed that Wister site 1, a small grassy

seepage ditch approximately 0.5 m x 2 m x 10 cm deep, was frozen over. Am-

bient minimum temperature for the day was —1° C. Under about 2 or 3 cm of

ice, 174 tarsalis and 207 Culiseta inornata pupae and larvae of all instars were

collected by pipette. In the laboratory the collection was divided equally between

2 plastic half-liter cups. The first cup was held at 25.6° C and the other was

refrigerated. After 16 hours at O° C, 5 tarsalis and 15 inornata larvae were im-

mobilized in the ice on the top two-thirds of the cup. The frozen and liquid

portions were separated and incubated at 25.6° C with the control. Almost no

mortality was observed for any of the 3 groups, and both species continued de-

velopment to the adult stage.

Washino and Bellamy (1963) found larvae under ice on 2 occasions in Kern

Nelson: Winter Biology of Culex tarsalis 29

County. Although development is slower and mortality probably higher at winter

temperatures, adults emerged from penned pupae in the field.

Immatures of tarsalis can also tolerate relatively high temperatures. In June 1968,

active larvae were found in a seepage ditch (site 11) about 5 cm deep where the

water temperature was 35° C. Jenkins (1950:914) reports that larvae have been

found in thermal waters at temperatures up to 39.0° C.

PHYSIOLOGICAL PARAMETERS

FAT BODY DEVELOPMENT

The percentage of females at Wister with Class 3 fat body (abdomen filled with

fat but not distended) was highest in November of both years (fig. 16, table 8).

Almost no females had abdomens that were noticeably distended by fat (Class 4).

No females collected in March, April or May 1969 had more than a trace of fat

body in the abdomen (Class 2). The pattern was nearly identical in the CO, traps

(although the data are less reliable because fewer females were examined). Fe-

males with Class 3 fat body were attracted to CO, in the same proportion that

they were found in shelters each month. Diapausing females should not be at-

tracted to CO, because they do not feed; therefore the maximum amount of fat

body developed by females at Wister was not characteristic of diapause.

In other areas where the winter biology of tarsalis has been investigated, fat

body is strongly developed. Where winters are coldest, tarsalis begins to store energy

reserves in early fall. By late fall, the abdomen is distended by a compact, lobular

fat body (Bennington, Sooter and Baer, 1958:301; Blackmore and Dow, 1962:

293; Harwood and Halfhill, 1964; Rush, Brennan and Eklund, 1958:289). Ap-

parently in the San Joaquin Valley, California, fat body develops less than in Wash-

ington (Harwood and Halfhill, 1964:597,598). Burdick and Kardos (1963:533) re-

ported that the largest percentage of females with abdomens filled or distended

with fat was in late December (91%) in Kern County. At Wister, the highest level

was in November of both years: 59% in 1968 and 38% in 1969.

The differences in degree and duration of fat body development may be due

to a direct effect of temperature or to latitudinal differences in response to photo-

period. Although decreasing day length seems to be the important factor for initi-

ation of fat body development in farsalis, its effect is reinforced by low tempera-

ture (Harwood and Halfhill, 1964; Harwood and Takata, 1965). Not all popu-

lations of the same species of mosquito show the same response to photoperiod,

even under a controlled temperature regime. In a laboratory strain of tarsalis from

Bakersfield, California, Harwood and Halfhill (1964:597) never observed fat body

as large or as compact as that found in hibernating females in central Washington.

This may reflect a difference in the natural populations of the 2 areas or it may

be due to changes in the Bakersfield colony after many years under laboratory

conditions. California and Washington strains of Anopheles freeborni showed lat-

itudinal differences in fat body response to photoperiod (Depner and Harwood,

1966). The Wister population of tarsalis shows no obvious difference in time of

initiation of fat body development from the other populations that have been

studied, but, probably due to the warmer fall and winter temperatures, the fat

body never develops to the same extent, and it is more rapidly metabolized as

the females continue to be active. |

Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

VE € L6 te 6 16 8-9 994

0 0 S-€ uel OL6I

Ol O€ OL IT 6 16 8-¢ 99d

8 (79) (8¢) cr SE 09 z OI-L AON

Iv OI 06 ZS 9 76 €1-O1 90

SZ 91 8 ) bl 9] v8 pI-TT des

II ST 78 | 6 (O01) 9I-vI 3ny

€S 6 16 ZE ZI 88 pI-TT ine

1a IT L8 I I 78 Ol 78 9 OI-pI ung

89 Iv LS I 69 SE ZS €] 17-81 ARW

LZ 6S LE v SZ SP ZS 17-61 Idyv

4 (O01) L (67) (IL) 6I-LI JUN

9 (LI) (¢¢) (0S) ZI 8 EE 8s LI-ST 99

L (pI) (98) i (67) (TL) O7-8I Url 6961

L i) = oa (ep) 101 ZI 88 pI-IT 99d

€ (19) (€¢) 67 6S Iv 97-b7 AON

L8 6p Iv 0 6 ZE a7 9¢ 1-7 AON

L9 ve vS 0 ria $6 SI CL I €] PT-7T des

LL OT 99 QT € SE ¢ S6 7 das-T¢ 8nV 8961

N € Z I 0 N € Z I a)

(%) ssojp Apog wy (%) sso Apog 10.4

SALVG

SdVUL °OD SUALTAHS

‘sde1y %Qd pur siojjeys Wo] sypsuny xajnD gTeuaj ul JUoWdojaAep Apog 184 — °g BGR,

Nelson: Winter Biology of Culex tarsalis 31

INSEMINATION

C. tarsalis females destined to overwinter usually mate first (Bellamy and Reeves,

1963:319; Burdick and Kardos, 1963:533; Kliewer et al., 1969:15) but insemina-

tion is not necessary for the fat body to develop (Harwood and Halfhill, 1964:

598). Since females mate soon after emergence from the pupal stage at all sea-

sons, the high percentage of females inseminated in the winter is probably due

to a dearth of immatures at that time that would give rise to noninseminated

females.

At Wister, I examined the spermathecae of too few females during some months

to form a definite conclusion about the insemination rate in the winter (table 9).

Table 9. — Insemination. Monthly percentage of inseminated female Culex tarsalis.

SHELTER CO, TRAP

Percent Percent

inseminated N inseminated N

1968 Aug 31-Sep 2 99 92 100 BS

Sep 22-24 86 104 96 57

Nov 2-4 62 26 94 78

Nov 24-26 68 44 100 10

Dec 11-14 84 hey (86) 7

1969 Jan 18-20 98 57 100 13

Feb 15-17 (100) 4 (100) 1

Mar 17-19 -- 0 - 0

Apr 19-21 100 19 96 26

May 18-21 -~- 0 -- 0

Jun 14-16 “~ 0 “ 0

Jul 11-13 97 32 98 43

Aug 14-16 (100) 7 (100) 4

Sep 11-14 78 63 96 23

Oct 10-13 82 50 95 38

Nov 7-10 52 40 (100) 9

Dec 5-8 46 11 90 10

1970 Jan 3-5 “ 0 oo 0

Feb 6-8 97 33 100 34

Mar 6-8 (100) 1 100 10

In early September 1968, 91 of 92 females examined from shelters were insemi-

nated. The percentage of inseminated females then declined for 2 months to 62%

in early November, and then steadily increased to 98% in January 1969. The fol-

lowing winter, there was again a suggestion of a fall decline of insemination rate

and subsequent increase by February. Nearly all females captured in CO, traps

were inseminated.

BLOOD FEEDING AND OVARIAN DEVELOPMENT

Engorged or gravid females were found all year (fig. 17). The percentage of

32 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

females with blood or developed ovaries was lowest in the late fall, at the same

time that percentage of females with fat body was greatest (fig. 16). Before the

coldest weeks of winter, fat body development decreased and blood feeding and

ovarian development increased. There was another apparent depression of blood

feeding and ovarian development in the summer. The data for 1968 are incom-

plete: the percentage of females engorged or gravid decreased in May and was

higher in early September, but shelter collections were not made in June or July.

In summer 1969, the lowest level was in July, the month before the apparent

depression of the female population (fig. 13). Although August was the hottest

month in 1969 (fig. 12), blood feeding increased that month. The level of blood

feeding may be influenced by the density of birds at Wister, which is much high-

er in the winter than in the summer.

Where winters are severe, engorged females are not found for many months.

In Utah (Blackmore and Dow, 1962:293) and in Colorado (Bennington, Sooter

and Baer, 1958:301), no engorged females are found by October. In the spring,

the first females do not bite bait animals until March in Oregon (Rush, Kennedy

and Eklund, 1963a:260). ,

In the San Joaquin Valley, numerous studies have documented the winter de-

pression in blood feeding (Kliewer et al., 1969:15; Nelson, 1964:248; Reeves, Bel-

lamy and Scrivani, 1958:84). Bellamy and Reeves (1963:316,321) found a nearly

complete cessation of blood feeding and ovarian development for a period of al-

most 3 months every year for 4 years, but did not find any period when feeding

ceased completely. At Wister, the reduction in blood feeding in the fall was not

as extreme as reported for the San Joaquin Valley. In 1967, engorged and gravid

females accounted for 23% of the female population during the lowest month

of blood feeding (fig. 17), 16% in 1968, and 9% in 1969. Resumption of blood

feeding was detected in late December the first two years and in early January

of the third. In Kern County, a gradual increase in the percentage of females en-

gorged is usually observed in late January and only rarely as early as late Decem-

ber (Bellamy and Reeves, 1963:316; Nelson, 1964:248).

In the Lower Rio Grande Valley, Texas, where tarsalis is essentially a winter

species, Eads (1965) collected many engorged or gravid females in light traps from

October to January. I have calculated the proportion of engorged or gravid fe-

males from the numerical data that he presented: 54% of the females were en-

gorged or gravid in October, but only 19% in November and 34% in December.

By January, the level had increased to 63%. No data was given for February or

March. These fluctuations in blood feeding occur about a month behind similar

fluctuations in population level (p. 26) and, like the population fluctuations, sug-

gest a bimodal curve of activity with a depression in early winter.

The blood meals of 43 females that I collected in September 1969 in the cul-

vert (site 6) were analyzed by precipitin test (Tempelis and Lofy, 1963) by Dr.

Tempelis at the University of California, Berkeley (table 10). Of particular inter-

est are the 3 snake bloods. As previously mentioned (p. 5), snakes are currently

under study as overwintering candidates of WEE virus. Only rarely are field-col-

lected tarsalis females found engorged with reptilian blood. Of over 10,000 en-

gorged females tested, Tempelis et al. (1965:174) detected 3 reptilian bloods in

Kern County, and Tempelis and Washino (1967:316) found 1 reptilian blood meal

in more than 900 tested in the Sacramento Valley.

Nelson: Winter Biology of Culex tarsalis 33

Table 10. — Hosts of Culex tarsalis as determined by precipitating antisera to blood meals of

females collected in shelters. September 1969.

Bird

Columbiformes 1

Negative* lips

Mammal ;

Rabbit 4

Man 1

Reptile

Snake 3

Negative lay

*Class Aves, order not determined.

PARITY

Parous females were found every month of the year at Wister (fig. 18). In shel-

ters, the percentage of parous females decreased in the fall of 1967 and 1969,

but by early winter it had increased. This corresponded to the simultaneous in-

crease in blood feeding and ovarian development (fig. 17) and to the decrease

in fat body development (fig. 16).

Even at the more northern latitudes, parous females do not disappear completely.

In northern Colorado, 2 of 597 females collected in a mine in early December

were parous (Blackmore and Dow, 1962:293-294). In eastern Colorado, Rush and

Tempelis (1967:311) found parous females in mid-April, before they found evi-

dence of the new spring generation of adults. In the San Joaquin Valley, at least

a few parous females are found in shelters throughout the winter (Burdick and

Kardos, 1963:532; Kliewer et al., 1969:17). Bellamy and Reeves (1963:319,320)

demonstrated experimentally that parous females could survive the winter in Kern

County, but survival was inferior to that of females that had not been allowed

to take a blood meal.

Although parity of shelter-collected females decreases in the fall in some areas,

it increases in females attracted to CO,. In northern Colorado, progressively fewer

females were caught in CO, traps in the late summer, but the percentage of those

females that were parous reached 100% by late September. The parity of empty

females in shelters dropped to zero during the same period (Blackmore and Dow,

1962:292). Nelson (1964:246) reported the same phenomenon in the San Joaquin

Valley later in the year. By the end of October, nearly all females collected in

CO, traps were parous, but parity in empty shelter-collected females had decreased.

He suggested that the fall population of tarsalis females is divided into two groups:

(1) Females that emerge from pupae in the summer take blood meals and oviposit

repeatedly for the duration of their lives. They usually do not survive the winter.

These are the host-seeking females with a high level of parity found in CO, traps

in dwindling numbers in the fall. (2) Females that emerge from pupae in the fall

experience diapause. They do not blood feed until after they have overwintered

(January in Kern County). Hence, few females found wintering in shelters are parous.

At Wister, too few females from CO, traps were examined to detect a diver-

gence in level of parity between females from CO, traps and those from shelters.

34 Contrib. Amer .Ent. Inst., vol. 7, no. 6, 1971

The indication is that there is no divergence. Fluctuations in parity level were

similar for the 2 methods from June to October 1969. In mid-October, 56% of

25 CO, trap females and 54% of 26 shelter-collected females were parous. In Kern

County, from the first to the second half of October 1952, parity decreased from

30% to 17% in shelters and increased from 90% to 93% in CO, trap collections

(Nelson, 1964:247).

CONCLUSIONS

The Wister population in Imperial Valley, California, remains active all year.

Winter diapause is so weakly expressed and of such short duration as to be nearly

nonexistent. In the fall, blood feeding and parity are somewhat reduced momen-

tarily and a fat body is developed. However, these indications of diapause begin

to disappear even before the coldest days of winter. The amount of fat body de-

veloped is less than typical for diapause, and females with a fat body are attracted

to CO,. Males, pupae and larvae of all instars are found throughout the winter.

The duration of diapause is different at various latitudes. In Canada and the

northwest United States, it may last 6 months. In California it is shorter. Benning-

ton, Sooter and Baer (1958:303) state that in California “true hibernation’’ does

not occur in tarsalis but studies by workers in the San Joaquin Valley have shown

almost complete cessation of activity for nearly 3 months. Females develop a fat

body and cease feeding almost completely; parous females are rare; males disap-

pear and larvae are very scarce. In the Lower Rio Grande Valley, Texas, there

is no winter diapause.

In the Imperial Valley, it does not seem that diapause would be necessary for

winter survival of tarsalis. Although blood feeding decreases in late fall, it resumes

before the coldest weeks of winter. Therefore, the temperature during this period

does not appear to inactivate the mosquitoes directly. Progressively shortened day

length in the fall may initiate diapause in populations of farsalis at all latitudes,

and low temperature probably reinforces the response. The temperature in the

fall and winter in the various areas studied is probably largely responsible for the

differences observed in degree of expression and duration of diapause. Where win-

ters are less severe, the females are more active, and energy reserves are depleted

more rapidly. The stimulus for breaking the diapause may be simply the complete

or partial exhaustion of the fat body, or as Bellamy and Reeves (1963:321) sug-

gest, accumulation of a critical amount of heat. In either case, the period of dia-

pause would be shorter where fall and winter temperatures are higher.

Potentially, tarsalis in the Imperial Valley is a good candidate for overwintering

arboviruses. However, without direct evidence of the viruses in the winter, it is

not possible to determine the population density and the level of blood feeding

necessary to maintain the cycle of transmission. Work, Vanis and Wallace (1969)

made 12 isolations of WEE, 10 of SLE and 21 of Turlock virus from farsalis

in 1968 and 1969 at Wister, but none of these isolates was from December or

January. Just as in Kern County, California, at Wister no virus has been found

when the level of blood feeding is lowest. Further investigations of the viruses

and their vectors and reservoir hosts will be necessary for a better understanding

of their seasonal relationship.

Nelson: Winter Biology of Culex tarsalis 35

REFERENCES CITED

Abell, D.L.

1959. Observations on mosquito populations of an intermittent foothill stream

in California. Ecology 40:186-193.

Allen, R., S.K. Taylor and S.E. Sulkin

1970. Studies of arthopod-borne virus infections in Chiroptera. VIII. Evidence

of natural St. Louis encephalitis virus infection in bats. Amer. J. Trop. Med.

Hyg. 19:851-859.

Anderson, A.W. and R.F. Harwood

1966. Cold tolerance in adult female Culex tarsalis (Coquillett). Mosquito News

26: 1-7.

Bailey, S.F., D.A. Eliason and B.L. Hoffmann

1965. Flight and dispersal of the mosquito Culex tarsalis Coquillett in the Sacra-

mento Valley of California. Hilgardia 37:73-113.

Barnett, H.C.

1956. The transmission of western equine encephalitis virus by the mosquito

Culex tarsalis Coq. Amer. J. Trop. Med. Hyg. 5:86-98.

Bellamy, R.E. and E.H. Kardos

1958. A strain of Culex tarsalis Coq. reproducing without blood meals. Mosquito

News 18:132-134.

Bellamy, R.E. and W.C. Reeves

1952. A portable mosquito bait trap. Mosquito News 12:256-258.

1963. The winter biology of Culex tarsalis (Diptera: Culicidae) in Kern County,

California. Entomol. Soc. Amer., Ann. 56:314-323.

Bellamy, R.E., W.C. Reeves and R.P. Scrivani ,

1967. Experimental cyclic transmission of western equine encephalitis virus in

chickens and Culex tarsalis through a year. Amer. J. Epidemiol. 85:282-296.

1968. Experimental cyclic transmission of St. Louis encephalitis virus in chickens

_and Culex mosquitoes through a year. Amer. J. Epidemiol. 87:484-495.

Bennington, E.E., J.S. Blackmore and C.A. Sooter

1958. Soil temperature and emergence of Culex tarsalis from hibernation. Mos-

quito News 18:297-298.

Bennington, E.E., C.A. Sooter and H. Baer

1958. The diapause in adult female Culex tarsalis Coquillett (Diptera: Culicidae).

Mosquito News 18:299-304.

Bidlingmayer, W.L.

1967. A comparison of trapping methods for adult mosquitoes: species response

and environmental influence. J. Med. Entomol. 4:200-220.

Blackmore, J.S. and R.P. Dow

1962. Nulliparity in summer and fall populations of Culex tarsalis Coq. Mosquito

News 22:291-294.

Blackmore, J.S. and J.F. Winn

1956. A winter isolation of western equine encephalitis virus from hibernating

Culex tarsalis Coq. Soc. Exp. Biol. Med., Proc. 91:146-148.

Blatt, J.M.

1968. Introduction to Fortran IV programming: using the Watfor compiler. Pacif-

ic Palisades, Goodyear Pub. Co. 313 p.

Brennan, J.M. and R.F. Harwood

1953. A preliminary report on the laboratory colonization of the mosquito,

Culex tarsalis Coquillett. Mosquito News 13:153-157.

36 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Brookman, B.

1950. The bionomics of Culex tarsalis Coquillett in irrigated areas of a lower

Sonoran environment. Doctoral dissertation. Univ. Calif., Berkeley. 192 p.

Bullock, H.R., W.P. Murdock, H.W. Fowler and H.R. Brazzel

1959. Notes on the overwintering of Culex tritaeniorhynchus Giles in Japan.

Mosquito News 19:184-188.

Burdick, D.J. and E.H. Kardos

1963. The age structure of fall, winter and spring populations of Culex tarsalis

in Kern County, California. Entomol. Soc. Amer., Ann. 56:527-535.

Burton, A.N., J. McLintock and J.G. Rempel

1966. Western equine encephalitis virus in Saskatchewan garter snakes and leop-

ard frogs. Science 154:1029-1031.

Buxton, P.A.

1935. Changes in the composition of adult Culex pipiens during hibernation.

Parasitology 27:263-265.

Carpenter, S.J. and W.J. LaCasse aoe

1955. Mosquitoes of North America (north of Mexico). Berkeley, Univ. Calif.

Press. 360 p.

Chaniotis, B.N.

1960. Autogeny in a colony of Culex tarsalis as affected by the level of pro-

tein in the larval diet. Master’s Thesis. Univ. Calif., Davis. 39 p.

Chaniotis, B.N. and W. Iltis

1960. Two additional characters identifying Culex tarsalis Coq. Mosquito News

20:319.

Chec, J.

1958. An autogenous strain of Culex tarsalis Coq. Mosquito News 18:134-136.

Chew, R.M. and S.E. Gunstream

1970. Geographical and seasonal distribution of mosquito species of southern

California. Mosquito News 30:55 1-562.

Christophers, S.R.

1911. The development of the egg follicle in anophelines. Paludism 2:73-88.

Clements, A.N.

1963. The physiology of mosquitoes. N.Y., Pergamon Press. 393 p.

Depner, K.R. and R.F. Harwood

1966. Photoperiodic responses of two latitudinally diverse groups of Anopheles

freeborni (Diptera: Culicidae). Entomol. Soc. Amer., Ann. 59:7-11.

DeStanley, M.

1966. The Salton Sea: yesterday and today. Los Angeles, Triumph Press. 128 p.

Detinova, T.S.

1945. [Determination of the physiological age of the females of Anopheles by

the changes in the tracheal system of the ovaries.] (In Russian.) Med. Para-

zitol. Parazit. Bolez. 14:45-49. English abstract in Appl. Entomol., Rev. (B)

34:180-181.

1962. Age-grouping methods in Diptera of medical importance with special ref-

erence to some vectors of malaria. World Health Organ., Monogr. Ser. 47.

2hGep:

1968. Age structure of insect populations of medical importance. Annu. Rev.

Entomol. 13:427-450.

Dow, R.P., G.A. Mail and C.S. Richards

1956. Observations on the overwintering of Culex tarsalis in northern Utah. Utah

Mosquito Abatement Ass., Annu. Meeting, 9th, Proc.:12-15.

Nelson: Winter Biology of Culex tarsalis 37

Dow, R.P., W.C. Reeves and R.E. Bellamy

1965. Dispersal of female Culex tarsalis into a larvicided area. Amer. J. Trop.

Med. Hyg. 14:656-670.

Eads, R.B.

1965. Biological notes on Culex tarsalis in the Lower Rio Grande Valley of Texas.

Mosquito News 25:61-63.

Eldridge, B.F. )

1968. The effect of temperature and photoperiod on blood feeding and ovarian

development in mosquitoes of the Culex pipiens complex. Amer. J. Trop.

Med. Hyg. 17:133-140.

Gebhardt, L.P. and G.J. Stanton

1966. The role of poikilothermic hosts as virus reservoirs. Jn: Symposium on

Arbovirus Diseases, Animal Vectors and Reservoirs. Pacific Science Congr.,

Lith, Proc. 20:30-34.

Gebhardt, L.P., G.J. Stanton, W.H. Hill and G.C. Collett

1964. Natural overwintering hosts of the virus of western equine encephalitis.

New England J. Med. 271:172-177.

Gebhardt, L.P., G.J. Stanton and S. de St. Jeor

1966. Transmission of WEE (western equine encephalitis) virus to snakes by in-

fected Culex tarsalis mosquitoes. Soc. Exp. Biol. Med., Proc. 123:233-235.

Gillies, M.T.

1954. The recognition of age groups within populations of Anopheles gambiae

by the pre-gravid rate and the sporozoite rate. Ann. Trop. Med. Parasit. 48:

58-74.

1958. A review of some recent Russian publications on the technique of age

determination in Anopheles. Trop. Dis. Bull. 55:713-721.

Guelmino, D.J.

1951. The physiology of Anopheles maculipennis during hibernation. An attempt

to interpret the phenomenon of gonotrophic dissociation. Trop. Med. Para-

sitol., Ann. 45:161-168.

Hammon, W. McD. and W.C. Reeves

1943. Laboratory transmission of western equine encephalomyelitis virus by mos-

quitoes of the genera Culex and Culiseta. J. Exp. Med. 78:425-434.

Hammon, W. McD., W.C. Reeves, S.R. Benner and B. Brookman

1945. Human encephalitis in the Yakima Valley, Washington, 1942, with forty-

nine virus isolations (western equine and St. Louis types) from mosquitoes.

J. Amer. Med. Ass. 128:1133-1139.

Hammon, W. McD., W.C. Reeves, B. Brookman and C.M. Gjullin

1942. Mosquitoes and encephalitis in Yakima Valley, Washington. V. Summary

of the case against Culex tarsalis Coquillett as a vector of St. Louis and western

equine viruses. J. Infect. Dis. 70:278-283.

Hammon, W. McD., W.C. Reeves, B. Brookman, E.M. Izumi and C.M. Gjullin

1941. Isolation of the viruses of western equine and St. Louis encephalitis from

Culex tarsalis mosquitoes. Science 94:328-330.

Hammon, W. McD., W.C. Reeves and G.E. Sather

1951. Western equine and St. Louis encephalitis viruses in the blood of experi-

mentally infected wild birds and epidemiological implications of findings. J.

Immunol. 67:357-367.

Harris, P., D.F. Riordan and D. Cooke

1969. Mosquitoes feeding on insect larvae. Science 164:184-185.

38 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Harwood, R.F.

1962. Trapping overwintering adults of the mosquitoes Culex tarsalis and Anoph-

eles freeborni. Mosquito News 22:26-31.

1967. The relationship between photoperiod and autogeny in Culex tarsalis. En-

tomol. Exp. Appl. 9:327-331.

Harwood, R.F. and J.E. Halfhill

1960. Mammalian burrows and vegetation as summer resting sites of the mos-

quitoes Culex tarsalis and Anopheles freeborni. Mosquito News 20:174-178.

1964. The effect of photoperiod on fat body and ovarian development of Culex

tarsalis (Diptera: Culicidae). Entomol. Soc. Amer., Ann. 57:596-600.

Harwood, R.F. and N. Takata

1965. Effect of photoperiod and temperature on fatty acid composition of the

mosquito Culex tarsalis. J. Insect Physiol. 11:711-716.

Hayes, R.O., R.E. Bellamy, W.C. Reeves and M.J. Willis

1958. Comparison of four sampling methods for measurement of Culex tarsalis

adult populations. Mosquito News 18:218-227.

Henderson, B.E. and L. Senior

1961. Attack rate of Culex tarsalis on reptiles, amphibians, and small mammals.

Mosquito News 21:29-32.

Horsfall, W.R.

1955. Mosquitoes. Their bionomics and relation to disease. New York, Ronald

Press. "723 0.

Hoy, J.B. and D.E. Reed

1970. Biological control of Culex tarsalis in a California rice field. Mosquito

News 30:222-230.

Husbands, R.C.

1958. An improved mechanical aspirator. Calif. Vector Views 5:72-73.

Jenkins, D.W.

1950. Bionomics of Culex tarsalis in relation to western equine encephalomy-

elitis. Amer. J. Trop. Med. 30:909-916.

Kardos, E.H.

1959. The effect of larval nutritional level on development of autogeny in a

colony of Culex tarsalis Coq. Calif. Mosquito Contr. Ass., Proc. 27:71-72.

Kardos, E.H. and R.E. Bellamy

1961. Distinguishing nulliparous from parous female Culex tarsalis by examina-

tion of the ovarian tracheation. Entomol. Soc. Amer., Ann. 54:448-451.

Keener, G.G., Jr.

1952. Observations on overwintering of Culex tarsalis Coq. (Diptera: Culicidae)

in western Nebraska. Mosquito News 12:205-209.

Kissling, R.E., R.W. Chamberlain, W.D. Sudia and D.D. Stamm

1957. Western equine encephalitis in wild birds. Amer. J. Hyg. 66:48-55.

Kissling, R.E., D.D. Stamm, R.W. Chamberlain and W.D. Sudia

1957. Birds as winter hosts for eastern and western equine encephalomyelitis

viruses. Amer. J. Hyg. 66:42-47.

Kliewer, J.W., T. Miura and H.C. Chapman

1969. Seasonal occurrence and physiology of Culex tarsalis in foothills of Fresno

County, California. Entomol. Soc. Amer., Ann. 62:13-18.

Lam, K.S.K. and I.D. Marshall

1968. Dual infections of Aedes aegypti with arboviruses. II. Salivary-gland dam-

age by Semliki Forest virus in relation to dual infections. Amer. J. Trop. Med.

Hyg. 17:637-644.

Nelson: Winter Biology of Culex tarsalis 39

Lees, A.D.

1955. The physiology of diapause in arthropods. Cambridge, Univ. Press. 151 p.

Longshore, W.A., Jr., ILM. Stevens, A.C. Hollister, A.G Gittelsohn and E.H. Lennette

1956. Epidemiological observations on acute infectious encephalitis in California,

with special: reference to the 1952 outbreak. Amer. J. Hyg. 63:69-86.

Loomis, E.C. and D.H. Green

1955. Resting habits of adult Culex tarsalis Coquillett in San Joaquin County,

California, November, 1953 through November, 1954. A preliminary report.

Calif. Mosquito Contr. Ass., Proc. 23:125-127.

Lord, R.D. and C.H. Calisher

1970. Further evidence of southward transport of arboviruses by migratory birds.

Amer. J. Epidemiol. 92:73-78.

Magy, H.I.

1955. A mosquito survey of Imperial Valley, Imperial County-December, 1953-

January, 1955. Calif. State Dep. Public Health, Bur. Vector Contr. June 9,

1955. M-41 (Mimeographed). 9 p.

Mail, G.A. and R.A. McHugh

1961. Relation of temperature and humidity to winter survival of Culex pipiens

and Culex tarsalis. Mosquito News 21:252-254.

Mer, G.G.

1936. Experimental study of the development of the ovary in Anopheles elutus

Edw. (Dipt., Culic.). Bull. Entomol. Res. 27:351-359.

Mims, C.A., M.F. Day and I.D. Marshall

1966. Cytopathic effects of Semliki Forest virus in the mosquito Aedes aegypti.

Amer. J. Trop. Med. Hyg. 15:775-784.

Moore, C.G.

1963. Seasonal variation in autogeny in Culex tarsalis Coq. in northern California.

Mosquito News 23:238-241.

Mortenson, E.W.

1953. Observations on the overwintering habits of Culex tarsalis Coquillett in

nature.. Mosquito Contr. Ass., Proc. 21:59-60.

Nelson, R.L.

1964. Parity in winter populations of Culex tarsalis Coquillett in Kern County,

California. Amer. J. Hyg. 80:242-253.

1966. A comparison of two techniques for distinguishing parous from nulliparous

Culex tarsalis Coquillett. Mosquito News 26:11-13.

Newhouse, V.F., R.W. Chamberlain, J.G. Johnston, Jr. and W.D. Sudia

1966. Use of dry ice to increase mosquito catches of the CDC miniature light

trap. Mosquito News 26:30-35.

Polovodova, V.P.

1949. [The determination of the physiological age of female Anopheles by the

number of gonotrophic cycles completed.] (In Russian, not seen.) Med. Para-

zitol. Parazit. Bolez. 18:352.

Rao, V.V.

1947. On gonotrophic discordance among certain Indian Anopheles. Indian J.

Malariol. 1:43-50.

Reeves, W.C.

1953. Quantitative field studies on a carbon dioxide chemotropism of mosquitoes.

Amer. J. Trop. Med. Hyg. 2:235-331.

1958. Problems of overwintering and natural maintenance of mosquito-borne

viruses. Int. Congr. Trop. Med. Malaria, 6th, Proc. 5:48-57,

40 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

1961. Overwintering of arthropod-borne viruses. Progr. Med. Virol. 3:59-78.

1965. Ecology of mosquitoes in relation to arboviruses. Annu. Rev. Entomol.

10:25-46.

1970. Evolving concepts of encephalitis prevention in California. Calif. Mos-

quito Contr. Ass., Proc. 37:3-6.

Reeves, W.C., R.E. Bellamy and R.P. Scrivani

1958. Relationships of mosquito vectors to winter survival of encephalitis viruses.

I. Under natural conditions. Amer. J. Hyg. 57:78-89.

Reeves, W.C, and W. McD. Hammon

1962. Epidemiology of the arthropod-borne viral encephalitides in Kern County.

Univ. Calif. Pub. Public Health 4: 1-257.

Reeves, W.C., G.A. Hutson, R.E. Bellamy and R.P. Scrivani

1958. Chronic latent infections of birds with western encephalomyelitis virus.

Soc. Exp. Biol. Med., Proc. 97:733-736.

Rosay, B. |

1961. Anatomical indicators for assessing the age of mosquitoes: the teneral

adult (Diptera: Culicidae). Entomol. Soc. Amer., Ann. 54:526-529.

1969. Anatomical indicators for assessing age of mosquitoes: changes in ovarian

follicles. Entomol. Soc. Amer., Ann. 62:605-611.

Roubaud, E.

1929. Cycle autogene d’attente et generations hivernales suractives inapparentes

chez le moustique commun Culex pipiens L. Acad. Sci. Paris, Compt. Rend.

188:735-738.

Rush, W.A.

1962. Observations on an overwintering population of Culex tarsalis with notes

on other species. Mosquito News 22:176-181.

Rush, W.A., J.M. Brennan and C.M. Eklund

1958. A natural hibernation site of the mosquito Culex tarsalis Coquillett in the

Columbia River basin, Washington. Mosquito News 18:288-293.

Rush, W.A., R.C. Kennedy and C.M. Eklund

1963a. Evidence against maintenance of western equine encephalomyelitis virus

by Culex tarsalis during spring in northwestern United States. Amer. J. Hyg.

77:258-264. .

1963b. Evidence against winter carryover of western equine encephalomyelitis

virus by Culex tarsalis. Mosquito News 23:285-286.

Rush, W.A. and C.H. Tempelis

1967. Biology of Culex tarsalis during the spring season in Oregon in relation to

western encephalitis virus. Mosquito News 27:307-315.

Ryckman, R.E. and K.Y. Arakawa

1952. Additional collections of mosquitoes from woodrats’ nests. Pan-Pacific

Entomol. 28:105-106.

Schaefer, C.H. and R.K. Washino

1969. Changes in the composition of lipids and fatty acids in adult Culex tarsalis

and Anopheles freeborni during the overwintering period. J. Insect Physiol.

15:395-402.

Shemanchuk, J.A.

1959. Mosquitoes (Diptera: Culicidae) in irrigated areas of southern Alberta and

their seasonal changes in abundance and distribution. Can. J. Zool. 37:899-

912.

1965. On the hibernation of Culex tarsalis Coquillett, Culiseta inornata Williston,

and Anopheles earlei Vargas (Diptera: Culicidae) in Alberta. Mosquito News

25:456-462.

Nelson: Winter Biology of Culex tarsalis 41

Sjogren, R.D.

1968. Notes on Culex tarsalis breeding in sewage. Calif. Vector Views 15:42-43.

Snow, W.E. and E. Pickard

1956. Seasonal history of Culex tarsalis and associated species in larval habitats

of the Tennessee Valley region. Mosquito News 16:143-148.

Southwood, T.R.E.

1966. Ecological methods, with particular reference to the study of insect popu-

lations. London, Methuen and Co. 391 p.

Spalatin, J., R. Connell, A.N. Burton and B. J. Gollop

1964. Western equine encephalitis in Saskatchewan reptiles and amphibians, 1961-

1963. Can. J. Comp. Med. Vet. Sci. 28:131-142.

Spielman, A.

1957. The inheritance of autogeny in the Culex pipiens complex of mosquitoes.

Amer. J. Hyg. 65:404-425.

Stamm, D.D., R.W. Chamberlain and W.D. Sudia

1962. Arbovirus studies in South Alabama, 1957-1958. Amer. J. Hyg. 76: 61-81.

Stamm, D.D. and R.J. Newman

1963. Evidence of southward transport of arboviruses from the U.S. by migrating

birds. An. Microbiol. 11(A):123-133.

Sudia, W.D. and R.W. Chamberlain

1962. Battery-operated light trap, an improved model. Mosquito News 22:126-

129,

Sulkin, S.E., R. Allen and R. Sims

1966. Studies of arthropod-borne virus infections in Chiroptera. HI. Influence

of environmental temperature on experimental infection with Japanese B and

St. Louis encephalitis viruses. Amer. J. Trop. Med. Hyg. 15:406-417.

Sulkin, S.E., R. Sims and R. Allen

1964. Studies of arthropod-borne virus infections in Chiroptera. II. Experiments

with Japanese B and St. Louis encephalitis viruses in the gravid bat. Evidence

of transplacental transmission. Amer. J. Trop. Med. Hyg. 13:475-481.

Swellengrebel, N.H.

1929. La dissociation des fonctions sexuelles et nutritives (dissociation gono-

trophique) d’Anopheles maculipennis comme cause du paludisme dans les Pay-

Bas et ses rapports avec “‘l’infection domiciliare”. Inst. Pasteur Paris, Ann.

43:1370-1389.

Tempelis, C.H. and M.F. Lofy

1963. A modified precipitin method for identification of mosquito blood meals.

Amer. J. Trop. Med. Hyg. 12:825-831.

Tempelis, C.H., W.C. Reeves, R.E. Bellamy and M.F. Lofy

1965. A three year study of the feeding habits of Culex tarsalis in Kern County,

California. Amer. J. Trop. Med. Hyg. 14:170-177.

Tempelis, C.H. and R.K. Washino

1967. Host-feeding patterns of Culex tarsalis in the Sacramento Valley, California,

with notes on other species. J. Med. Entomol. 4:315-318.

Thomas, L.A.

1963. Distribution of the virus of western equine encephalomyelitis in the mos-

quito vector, Culex tarsalis. Amer. J. Hyg. 78:150-165.

Thomas, L.A. and C.M. Eklund

1960. Overwintering of western equine encephalomyelitis in experimentally in-

fected garter snakes and transmission to mosquitoes. Soc. Exp. Biol., Proc.

105:52-55.

42 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Vargas, L.

1956. Especies y distribucion de mosquitos mexicanos no anofelinos. Inst. Sa-

lubridad y Enfermedades Trop., Rev. 16:20-36.

Verheijen, F.J.

1960. The mechanisms of the trapping effect of artificial light sources upon

animals. Arch. Neerland. Zool. 13:1-107.

Washino, R.K. and R.E. Bellamy )

1963. Winter observations on larval populations of Culex tarsalis Coquillett in

Kern County, California. Mosquito News 23:162-163.

Washino, R.K., R.L. Nelson, W.C. Reeves, R.P. Scrivani and C.H. Tempelis

1962. Studies on Culiseta inornata as a possible vector of encephalitis virus in

California. Mosquito News 22:268-274.

Wigglesworth, V.B. |

1965. The principles of insect physiology. Ed. 6. London, Methuen Co. 741 p.

Williams, J.B.

1968. Statistical analysis. Olivetti Underwood Corp., Systems Groups, Marketing

Services Division. 324 p. ig

Work, T.H., R. Vanis and H.G. Wallace

1969. Arboviruses in southern California. Div. Infect. Trop. Dis., Sch. Public

Health, Univ. Calif., Los Angeles. (Unpublished manuscript.) 8 p.

Publication dates for Volumes 5 and 6

Contributions of American Entomological Institute

Volume 5

No. 1 January 28, 1970

No. 2 January 28, 1970

No. 3 February 27, 1970

No. 4 April 10, 1970

No. 5 May 5, 1970

No. 6 June 2, 1970

No. 7 August 17, 1970

Volume 6

No. 1 August 3, 1970

No. 2 August 17, 1970

Nelson: Winter Biology of Culex tarsalis

IOATY

OpeIO[O)

“ATV A [elioduly oy} pue erusosiyed jo sdew— | onstJy

SID] IUOTLY

Qe ae 00S

SO =

sI9}OWIO[LY Py >

OOT 0

oN Re ap tr, Se x

2 u eS in eas Se

jewodwy —¥ ec: ROHR A

uinbeor ues

‘

Age A [elodwy \

IO1STM oh Bag UO}eS \

U SAAR ORS. \

S \

eee: x | Aayye Rr

oJusurTeOeS

Aaqe A ,

EIJayOROD

‘Suen

= eax

— ~ eo am

aod Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

WG, she, *

ew Ma, a

PL eS

Vb,

“. Saas Ma, os, ee We, la,

i) SLES Ee "ate. Me te, aw.

Ue, ag, tll cence ole, at Ma, cY?

fo, alle, Mea, “alt, . lee, i Sia

Me an tg. a 4

Me Ny Me, ashlee,

le Ma Ne, a

Wile, hae MG la, SMe, Ale,

4, NY (9) le, Al

ae ; aie, atl A

r | | | |

kilometers

Figure 2.—The Wister Unit of the Imperial Waterfowl Management Area, Imperial

County, California. Circled numerals indicate sites where routine collections of

Culex tarsalis were made.

Nelson: Winter Biology of Culex tarsalis 45

Figure 3.—A diurnal shelter for Culex tarsalis adults: a cement culvert (site 6)

under California State Highway 111 at Wister.

46 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Figure 4.—A diurnal shelter for Culex tarsalis adults: a wooden bridge (site 5). The 2

m white rod is marked at 0.5 m intervals.

Figure 5.—Red ice cream cartons under a wooden bridge (site 5) from which Culex

tarsalis adults were collected each month with an aspirator.

Nelson: Winter Biology of Culex tarsalis 47

Figure 6.—A duck pond at Wister.

Figure 7.—A typical breeding site of Culex tarsalis at Wister: a shallow, grassy depression

(site 12) with a recent accumulation of seepage from an irrigation canal.

48 Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

ii eit

Figure 8.—The sewage oxidation pond at Wister. The arrow indicates the collecting area

(site 8) shown in fig. 9.

& ie

Figure 9.—Site 8 at the edge of the sewage oxidation pond where immatures of Culex

tarsalis were collected each month. The wooden stake projects approximately 0.25 m

above the water surface.

Nelson: Winter Biology of Culex tarsalis 49

124

Number of

females

Number of eggs per female

0 100 200 300

Figure 10.—Frequency distribution of the number of developing eggs (Stages III-

V) per female Culex tarsalis collected at Wister. September 1967-April 1970.

Number of

females

Number of eggs per female

0 10 20 30

Figure 11.—Frequency distribution of the number of developing eggs per female

for those individuals with 30 eggs or less.

1971

vol. 7, no. 6,

Contrib. Amer. Ent. Inst.,

OL6l

V Wo ds Pit Ne OS Vv

a — ]yeyures

6961

"BIUIOST]e ‘TeLodwy]

‘JOLIISIC] UOTIVSLLI] [eLIodwy sy} Aq pops1Oosl uoT}eyId1901d A[YJUOU! [e}0} pu SoIN}eIOdUIO} SUSI} XO pue URW A[YJUOW—'Z[ INST

896I L996!

Poe Ae WOW ed Ta NOUS Ys ie Wy Ae ee ae

oe ee

ig a. Fs

—— UINUITUIW «~UPOUT

WNUITXPUl UPD —

—— wInwIxeul

‘A[[ROTWUIeSOT poyojd suvow orjowuryjue AjyUOW “(LZ ‘S ‘Z “T SeqIs) SIo}OYS Ospliq

py pue “(9 o}IS) IoqJOYS yOAIND ke ‘sdey %QD ‘7QD YIM sdeqy 1YSI] Ul poyo][Oo soyeuley sippsup] XajnD Jo sioquiInN—‘¢E[ sINsTy

OL6I 6961 8961 EPG!

Vo ae eg I VN a ee Os YUE UF UNV A eee oe

= v

is N

= / \

C) °

Ce / \

o) / \

Be Fes = :

. \

eo / :

= /

aa / \

s ‘ / \

= \ / Ps :

Kn. Vv = :

= x \ OQ. u 4 oe oe * \

¢ \ : A ? . / \

az ¥ O--O x \

D 4

7 y

sspllg t-——--G

SAID CO - = <2 = 0

dei “G5 >> =

tO) + usr] x——_—_-x

Ol ™

as |

.@?)

(ore)

00S

5Z Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

Canada and

northwestern U.S.

7 Se FN

Imperial Valley ese. J eaeeene os ete ae sa Sear Be NORE

(Aypothetical Pe Rs

intermediate)

Lower Rio Grande

Valley, Texas

(Hypothetical extreme)

Figure 14.—Diagrammatic representation of the seasonal fluctuations in abun-

dance of Culex tarsalis collected in light traps at various latitudes.

Nelson: Winter Biology of Culex tarsalis

‘(L-S ‘Z ‘| SOUS) S19][9US UI Ppo}d9]]09 s7ppSunJ XajND Jo UoTjeTNdod 94} UT soTeUI Jo o8e}USdIEd ATYJUOW—"ST einst4

OL6

W ad

{Pa <N eo

6961

Pee oh

SOT

8961

soyeul

yU90I9 g

O01

s(soTeuley pue seTeut)

uotye—ndod [e100],

18S

SSE

c99

ve6

609

Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

“APO 1¥J € SSPTD YIM S1/DSUD] XajNZD s{eUId] JO osejUadIOd ATYJUOW *Apog je4—‘9][ oINsIy

OL6l 6961 8961

gpa NN Oe Ne i a Ne ee Nee ore

VPN

v \

® \ | 2

dU, CO) ee-= ace: =

PINS a

TOT

O€

TLT

€

Apoq }e}

YyIM

JUN9IOg

O09

OO!

- 193]94S

°N

- dery, “oD

Nelson: Winter Biology of Culex tarsalis

"(L-S ‘Z ‘] SalIs) SIO1[9US UI Pa1oaT[OO pIAIS IO pasIoZua ‘AjyduIa s1Msun] XajnD s[eUlej Jo osejuadIed ATYJUOW—"L] ons

oad

posiosus — =

Aydwao —

696l

896l

e¥)

O€

ee)

L961

jud010g

Contrib. Amer. Ent. Inst., vol. 7, no. 6, 1971

OL61

W a

‘sdel} “OD pure SIO}[OYS UL PdJOOT[OO ‘SaTeWay S1psun] xajnD Aydwis snosed Jo a8ejUs0I0d ATYJUO| *AiLWIeg—"Q] sNst4q

10S: #1»

6961 8961 L961

PE oN OS ts 2 WN

snoied

jUDd0IOg

OOl

oD AUN Aw oO NN Ni rs Sk es a i SSO SS

Se eu) We Gis ee -den ¢Q5

No.

(Continued from inside front cover)

Parts of Volume 4, with prices

Mosquito studies (Diptera, Culicidae), 1968.

X. Peters, T. Michael. Dixinae originally described from North America. 17 pages.

XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes

originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay.

29 pages. Price: $1.25, postpaid. a

Mosquito studies (Diptera, ‘Culicidae), 1969.

XII. Berlin, O. G. W. A revision of the Neotropical subgenus Howardina of Aedes.

Carlson, Robert W. and Fred B. Knight, 1969.

Biology, taxonomy, and evolution of four sympatric Agrilus beetles (Coleoptera:

Buprestidae). 104 pages, 47 figures. Price: $3.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1969.

XIII. Barr, Ralph A. and Sylvia Barr. Pupae of the genus Psorophora in America

north of Mexico. 19 pages, 11 figures.

XIV. Berlin, O. G. W. Redescription of Toxorhynchites (T. ) gigantulus from the

Philippines. 7 pages, 2 figures.

XV. Zavortink, Thomas J. A new species of treehole breeding Anopheles from

the southwestern United States. 12 pages, 4 figures.

Price: $1.25, postpaid.

Matthews, Robert W., 1970. A revision of the genus Spathius in America north of Mexico

(Hymenoptera, Braconidae). 86 pages, 31 figures. Price: $3.50, postpaid.

postpaid.

Parts of Volume 5, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XVI. Zavortink, Thomas J. A new species of treehole breeding Aedes ee

from southern California. 6 pages, 2 figures.

XVII. Adames, Abdiel J. and Charles L. Hogue. Two new species of Deinocerites from

Costa Rica. 6 pages, 6 figures.

XVID. Berlin, O. G. W. The subgenus Micraedes of Culex, 30 pages, 12 figures, 1 map.

Price: $2.00, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XIX. Zavortink, Thomas J. The treehole Anopheles of the New World. 35 pages, 12

figures. Price: $1.25, postpaid.

Mosquito studies (Diptera, Culicidae), 1970.

XX. Schick, Robert X. The Terrens Group of Aedes (Finlaya). 158 pages, 60 figures.

Price: $5.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

V. Reinert, John F. Genus Aedes, subgenus Diceromyia Theobald in southeast Asia.

43 pages, 15 figures. Price: $1.50, postpaid.

‘Howden, Anne T. The Tanymecini of the West Indies (Coleoptera: Curculionidae). 74 pages,

76 figures. Price: $2.75, postpaid.

Kethley, John B., 1970. A revision of the family Syringophilidae (Prostigmata: Acarina).

76 pages, 38 figures. Price: $2.75, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

VI. Mattingly, P. F. The genus Heizmannia Ludlow in southeast Asia. 104 pages,

dl figures. Price: $4.00, postpaid.

Parts of Volume 6, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXI. Belkin, John N., Sandra J. Heinemann, and William A. Page. The Culicidae of

Jamaica. 458 pages, 126 figures. Price: $15.00, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1970.

Vil. Tyson, W. H. Genus Aedeomyia Theobald in southeast Asia. 27 pages, 6 figures.

Vill. Tyson, W. H. Genus Aedes, subgenus Mucidus Theobald in southeast Asia. 54 pages,

21 figures. Price: $3.50, postpaid.

Parts of Volume 7, with prices

Mosquito studies (Diptera, Culicidae), 1970.

XXII. Zavortink, Thomas J. A new subgenus and species of Aedes from Arizona.

11 pages, 2 figures.

XXII. Schick, Robert X. Additions and corrections to the revision of the Aedes

terrens group. 27 pages, 8 figures. Price: $1.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1971.

XXIV. Adames, Abdiel J. A revision of the crabhole mosquitoes of the genus

Deinocerites. 154 pages, 58 figures. Price: $6.00, postpaid.

(Continued from inside back cover)

Contributions to the mosquito fauna of southeast Asia, 1971.

IX. Zavortink, Thomas J. The genus Orthopodomyia Theobald in Southeast Asia.

37 pages, 12 figures.

X. Dobrotworsky, N. V. The genus Culiseta Felt in Southeast Asia. 24 pages, 8 figures.

XI. Sirivanakarn, Sunthorn. A proposed reclassification of Neoculex Dyar based

principally on the male terminalia. 24 pages, 3 figures. Price: $3.50, postpaid.

Contributions to the mosquito fauna of southeast Asia, 1971.

XII. Mattingly, Peter F. Illustrated keys to the genera of mosquitoes (Diptera, Culicidae).

84 pages, 34 plates. Price: $3.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1971.

XXV. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes

originally described from Brazil. 64 pages. Price: $2.50, postpaid.

Mosquito studies (Diptera, Culicidae), 1971.

XXVI. Nelson, Michael J. Winter biology of Culex tarsalis in Imperial Valley,

California. 56 pages. Price: $2.50, postpaid.

Noe

+ 3

apie

‘

~ 1

—_

«)

re io -

*Y -

ry iy ih

a The

i Aras ri if

Nbaeey ,

; i 4 i uh

4) Ep

aa)

Ana

ith

‘ft

dit

LIBRARIES

TITUTION NOILALILSNI

INSTITUTI

417 LIBRARIES SMITHSONIAN

SMITHSONIAN INSTITUTION NOLLALILS

VINOSHLINS S31UVUGIT

NOILNLILSNI

NVINOSHLIWS

IBRARIES

NOILNLILSN

: Li B RARIES SMITHSONIAN

NSTITUTION

TITUTION NOILLALILSNI

NVINOSHLIWS SZluyvuait

HSONIAN

SMIT

iuvygit SMITHSONIAN”

NOILNLILS

NVINOSHLIWS

a \

SS(UVUGIT LIBRARIES

INSTITUTION

“LIBRARIES SMITHSONIAN

SMITHSONIAN

VINOSHLIWS

TITUTION |

NVINOSHLIWS

rAelEeS

NALILSNI

NOILNLILSNI

NVINOSHLIAS S3INVUEIT LIBRARIES

INSTITUTION NOILOLILSNI

ES SMITHSONIA

39)

1 LIBRARI

INSTITUTION

NVINOSHLINS S3INVUGIT LIBRARIES

NVINOSHLINS S31uVvudI

OILALILSNI

Saluvyudi

RAR |

SMITH

NOILAL

LLSNI

1 LIBRARI

JiUVUEl

NOSHLIWS

uVudII

Nv!

LIBRARIES

NOILLMLILSNI

NOILALILSNI

INSTITUTION

SN Me

WA NY RAS

“ ASEAN :

SMITHSONIAN

NOILNLILSNI

4 NABRARIES

1_LIBRARIES

Saluvugl

NYINOSHLIWS

LIBRARIES

ALILSNI

NVINO

TITUTION NOILOLILSNI

iNS

SMITHSONIAN

LIBRARIES

NVINOSHLINS S31uVvUa

ON NOILNLILSNI

MITHSONIAN

: QO

0 =}

. ote

of.

SMITHS

“nish

NVINOSHLINS S31YV

SMITHSONIAN INSTITI

SIJ(UVUGIT LIBRARIES

NVINOSHLIANS S31YY

VINOSHLIWS

INSTITL

NOLLOLILSNI

S31uyV

INSTITUTION

SMITHSONIA

Saiyy

IT LIBRARIES

SMITHSONIAN

INSTITUT

sziuvug

NVINOSHLIWS S3IUYY

NVINOSHLIWS

SMITHSONIAN INSTITI

ARIES

Ill

3 9088 00905 6607

|| LIB

Tee

jen ie z é 5 ae aah : s 2 ; = ; pag Sieh 3 See nw vikGh as ig3G oe ah taki

PAO RE NG z 4 . : i , S ee

; : : : : : PGS als 3 bby ai

se ah he

Rees

; aipenlbadgaay-aip

high son ete A f ; Coss eWiaky ute f

Mili Hs : i ats SNe: 5 GRUB Nay mishap aed

Sn Sb Bb ig ee pane

; i . / DE Wa fee peisly de

Ne Pe GE Opes ¥ es a Ease tes) Pe ee aah

ia be ; i z Been Piha ti ek a bo np x p ape 4 Hak ap Ai a eyed wif patients “ine, Pajjeala aervesd ibees

i es sa Bra Ss chp ee Php ipa AGAR AS the Gidget

Graphs wakes) say

en aries eh ee hie Sal ea

PES A

if est Hb ap 3 “ys Z 2 PRUE MAE Se Te Aiba Ae

RY Den wi had a gia Be. EY % . Bo tes) sie ots f

BUN Rian aha: : i :

: ye , tgliontroh'y ¢ SWAG EWES? 5

pid. PND pr : i $ $ 2 ne Se this Svat TEDW ae Aiebie ol ote digi f

a ‘ eine fee ay

ie ey ey ee

Ui ie hi Sat

RSP arg ica,

qs e poked is Fok Ast sah ss ier ney

US WB

faire

Pit 8

PN NER 4 - Z f . P Z y f

ee Dak iy es ae a fe ; ; ; ; Se ta i ey ; Hh a : : PSG AB EN STE EIEN AH ; ie eka ae be ee sit} ify ¢ Z ; raed yeh Z SOSH HA hee Aah tsb

i i } i ‘i i fA " “ GEA g YE i q : wads a FO 7h sian aig 4m ect:

ee oh i Bed ‘fh 4 ¥ 3 i wisn

i 2 gh: 4 f Sar Bey RGpeh Zatti ap yaa Liyang

Hi Spy Ay Beh As a eS AaOHE (ap high a Teh is

, sp is y ‘ , se Z al Palas M i 5 F y

Laing ah , # ; 5 Ma : i bh ed ‘ ‘ ics See at fall 2 i AE Op a nib aLes simi aag 2 aS Eig tv sbeebs SGD ip io oe 4

i : 4 F ¢ ¢ » g i 3 “ } AeA 2y

2 A fy i ye 1B 23 if Metra MENON

Labrie 4

Bek pey ang apie

PMG ep Heat #4 : : : é Ania iP Geka sy 2

re ad yt eae ,

Oe aR aE ‘4 BY SiN aie SPadwety

™ 4 GUE BAYA’ 3 BP IDE RB EY Ze Sy Lege 4,

LEST A Ay Sh 3 ie oT pethpenniogs

et bathynits OSL ADVE TBS De AS AP NPN Ee ay

a cp a i 5 Hiss cea eat rue ie Baga

Fae ire

Pe Ose ups be ee Os usd. ; tes ite ay

ine ih CLAUS RPG ee i Z sky 4 $f LE aes Haat 2 abe ie ie ode Sipe se asviehe,

LN SPA SNE NEG ga RA? é ‘ ue ee 8 blige

ti Siabs is) 5 Sub MA RAG : PRB AN A Sh ABS } Sih 4) ; en mi dea % "a : fie! hae Les ee ey 2h int i ‘i ape sito nce aed. j uterine j PEO bah ie Can fae

pomemess ue elig i ; MS v Y RORY fe i aie fy Ah: . 5 pitage 5 He Coat: , asda He Ese Z vale ; A OE Ti a ane ING Chess Piel Bata ‘ % “i oR bee és bale. et £

se yibeae

ysis fakin Big foie

VPA A :

ME dari Eri peo Intl degnch a vase Sib

Hpbi aga ee ph Oe ba fisifccib? ch iape eu,

sot MEPS NY. i pes sig Batons

Ag aig ws aN j i is bays ¥ VA, 4 nyt ah és “ ip : e Pali “s ; i ‘ : i - ii : § 8 ts BAS ne BA be! wife rs “hte ways pai vhs pens

4} g

Persie aise

ite

Lite Hayes ag ath Wi Hin xipsls

Ate S\hunilealy whee f i ?

ef i ae barat phe! BAIA aD Wale ey sara: ipieals ie ee ere

; ed isd SHO HAY: Tait Beles ae 4 h Fefoeinet pis

GME RPAE EOS shy. hs a

Fagan fips i

pagans. ; z Sat cakes : be ie A F 2 SAA EH ye tai : EN iA 4 A : 3 iteonipecls

HX ae Bales ihe intot he 5

mia Gate Saas i : BSS Bade AS 2 : Fuiinceed § é G his fe j : 3 { sas 1 545) Binys NERNEY Lips

sales gh : ; id, SS SEPP aS ee SEAL AER ab BV aid TARE 9 LENE AE ah Ay “ins 4 f fei fs i Sites ica eid ; PAYDAY I IAGS 5 os ets oe hepsi

; ; } IEAM a MAA SY i sh Alte! He ; 24s CIS ee fom) op oa Vera fees iy wena as ep: ; a ip: i

. i ies, eRe fo BODO, ib EO) 0 Pee Stan Fae nif “ Mp) ee sill la

pai eee nete Ge ip Ug Ge stieriiy Ay asi af a aeons ‘ SPE Ath Ph AINE Sth sa ay ACIP RAI en ei Kk'y

Bee eT nis pet Pape iret AT Se rn

el abt) poy gee