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Contributions ,/

of the

American Entomological Institute

Volume 17, 1979-1981

Zavortink, Thomas J. Mosquito studies (Diptera,

Culicidae) XXXV. The new sabethine genus

Johnbelkinia and a preliminary reclassification of the

composite genus Trichoprosopon. 61 pages. October 8,

1979.

Berlin & Belkin. Mosquito studies (Diptera, Culicidae)

XXXVI. Subgenera Aedinus, Tinolestes, and

Anoedioporpa of Culex. 104 pages. Nov. 13, 1980.

Huang, Yiau-Min and James C. Hitchcock. Medical

Entomology Studies-XII. A revision of the Aedes

scutellaris Group of Tonga (Diptera: Culicidae).

107 pages. June 10, 1980.

. Harrison, Bruce A. Medical Entomology Studies - XIII.

The Myzomyia series of Anopheles (Cellia) in Thailand,

with emphasis on intra-interspecific variations (Diptera:

Culicidae). 195 pages. November 13, 1980.

Mattingly, Peter. Medical Entomology Studies - XIV.

The subgenera Rachionotomyia, Tricholeptomyia, and

Tripteroides (Mabinii Group) of genus Tripteroides in

the Oriental Region (Diptera:-Culicidae). 147 pages.

May 13, 1981. Vs dab a Deas

\ AUG 27 1981

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Contributions

of the

American Entomological Institute

Volume 17, Number 1, 1979

eae

MOSQUITO STUDIES (Diptera, Culicidae)

XXXV. The new sabethine genus Johnbelkinia and a preliminary reclassification

of the composite genus 7richoprosopon. By Thomas J. Zavortink

MOSQUITO STUDIES (Diptera, Culicidae)

XXXV. THE NEW SABETHINE GENUS JOHNBELKINIA

AND A PRELIMINARY RECLASSIFICATION OF THE COMPOSITE

GENUS TRICHOPROSOPON'

By

Thomas J. Zavortink?

CONTENTS

INTRODUCTION .. .

MATERIAL AND METHODS .

SYSTEMATICS

Taxonomic History .

Taxonomic Characters .

Proposed Classification

BIONOMICS .. .

KEYS TO GENERA AND SUBGENERA

TAXONOMIC TREATMENT .

Genus Johnbelkinia .

Keys to Species

1. Johnbelkinia ulopus

2. Johnbelkinia longipes .

3. Johnbelkinia leucopus

REFERENCES CITED

PIGURES este cscc

CONSPECTUS OF PRELIMINARY RECLASSIFICATION OF COMPOSITE

GENUS TRICHOPROSOPON . . LORE Wc cr > he amie Mak biak aout te

INDEX TO SCIENTIFIC NAMES .

Contribution in part 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 Com-

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

*Department of Biology, University of San Francisco, San Francisco, CA 94117.

2 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

INTRODUCTION

In their study of American sabethine mosquitoes, Lane and Cerqueira (1942)

placed all the New World species of the tribe Sabethini without obvious specializa-

tions in structure or coloration of the adult stage into the single genus Trichoproso-

pon. Within this genus they recognized seven subgenera almost exclusively on the

basis of features of the adults. The delimitation and internal classification of the ge-

nus proposed by Lane and Cerqueira have been followed in large part by all subse-

quent authors treating species of the group, and they are followed with only minor

alterations in the recent world catalog of mosquitoes by Knight and Stone (1977:

312-316). As the larvae and pupae of a greater number of species of Trichoprosopon

have been discovered and examined carefully, it has become evident that these stages

provide a wealth of excellent taxonomic characters, and equally obvious that the ex-

isting classification for the genus is inadequate and artificial. Since Trichoprosopon

appeared to be in need of restudy, and since it is the most generalized group of

American Sabethini, it seemed appropriate that it be revised before the more special-

ized or derived groups in the tribe, and thus I undertook a study of the genus.

As a result of my study of Trichoprosopon, I believe the species presently includ-

ed in the genus belong to four separate phyletic lines. Since it appears that some of

these lines may be more closely related to mosquitoes presently placed in other gen-

era than they are to each other, I choose at this time to recognize each of the four

lines as a distinct genus. The present paper is the first in a series treating these gen-

era. It includes an introduction to the study as a whole; an outline of my proposed

reclassification of the group, with keys to the segregrate genera and subgenera and a

list of the nominal species in each; and a revision of the smallest genus and the only

one requiring a new name.

I am indebted to many persons whose assistance and cooperation Hinde this study

possible. I am especially grateful to John N. Belkin for encouragement throughout

the duration of this study, for informative discussions on sabethine taxonomy, for

reading and criticizing this manuscript, for notes on type specimens in Brazilian mu-

seums, and for opening his home to me during my visits to Los Angeles.

I also thank J. Hal Arnell, Sandra J. Heinemann and Peter F. Mattingly for stimu-

lating discussions related to this study; Curtis W. Sabrosky and E. L. Peyton for help

and consideration during my visit to the Smithsonian Institution; Sharon L. Belkin

for drawings of type specimens deposited in South America; Sandra J. Heinemann

and William A. Powder for preparation of much of the material studied; Nobuko

Kitamura and L. Margaret Kowalczyk for preparation of the preliminary drawings

and final illustrations; Thomas S. Acker for use of his microscope with phase con-

trast and Normarski interference optics; Norma M. Washington for typing rough

drafts of the manuscript; and Sandra J. Heinemann for preparing the text copy for

lithoprinting.

And I thank the following for generously loaning me mosquitoes in their care or

for providing information about type specimens: John N. Belkin, Sandra J. Heine-

mann and William A. Powder, University of California, Los Angeles; Curtis W. Sabro-

sky, Systematic Entomology Laboratory, U. S. Department of Agriculture, Washing-

ton, D. C.; Ronald A. Ward and E. L. Peyton, Medical Entomology Project, Smith-

sonian Institution, Washington, D. C.; Peter F. Mattingly, British Museum (Natural

History), London; Paul H. Arnaud, Jr., California Academy of Sciences, San Francis-

co; Alfonso Diaz Najera, Instituto de Salubridad y Enfermedades Tropicales, Mexico

Zavortink: Reclassification of Trichoprosopon 3

City; Miguel Garcia, Instituto Nacional de Microbiologia, Buenos Aires; Pedro Galin-

do, Gorgas Memorial Laboratory, Panama, R. P.; L. L. Pechuman, Cornell University;

Pedro W. Wygodzinsky, American Museum of Natural History, New York; Oswaldo

P. Forattini and Maria das Dores Cotrim, Faculdade de Saude Publica, Universidade

de Sao Paulo; S. L. Tuxen, Universitetets Zoologiske Museum, Kobenhavn; R. Krau-

se, Staatliches Museum fiir Tierkunde, Dresden; Oswaldo Cruz, Filho, Instituto Os-

waldo Cruz, Rio de Janeiro; George W. Byers, University of Kansas; Ferenc Mihalyi,

Természettudomanyi Mazeum Allattara, Budapest; Charles L. Hogue, Los Angeles

County Museum of Natural History; and D. C. Geijskes, Stichting Surinaams Muse-

um, Paramaribo.

MATERIAL AND METHODS

The majority of the specimens examined during this study were borrowed from

the University of California at Los Angeles [UCLA], and were collected for the pro-

ject “Mosquitoes of Middle America’”’ (Belkin, Schick et al. 1965). Additional speci-

mens were borrowed from or studied at the United States National Museum of Natu-

ral History [USNM], including the primary types of cerqueirai, culicivora, dicella-

phora, homotina, lampropus, leucopus, mogilasia, moralesi, rapax, schedocyclia,

shropshirei, townsendi, trichopus, trichorryes, ulopus, vonplesseni and wilsoni; the

British Museum (Natural History) [BM], including the primary types of espini, fluvi-

atilis, frontosa, lunata, nivipes, pallidiventer and perturbans; the California Academy

of Sciences [CAS] ; the Medical Entomology Project [MEP] ; the Instituto de Salubri-

dad y Enfermedades Tropicales [ISET] ; the Instituto Nacional de Microbiologia

[INM]; the Gorgas Memorial Laboratory [GML] ; Cornell University [CU] ; the Los

Angeles County Museum of Natural History [LACM]; the American Museum of Nat-

ural History [AMNH]; the Faculdade de Saude Publica, Universidade de Sao Paulo

[FH] ; the Instituto Oswaldo Cruz [IOC] ; the Stichting Surinaams Museum [SSM] ;

the Universitetets Zoologiske Museum [ZMC], in particular the holotype of longipes;

and the Staatliches Museum fiir Tierkunde [SMT], in particular the holotype of sub-

splendens.

The taxonomic procedures used in this study are the classical comparative mor-

phological ones that I outlined elsewhere (Zavortink 1974). The form of presenta-

tion, terminology and abbreviations used in the descriptions follow Belkin (1962a)

for the most part; terminology for the legs, however, is from Schick (1970). The ex-

tent of the light tarsal markings is measured from the apex of the segments, so ‘‘Tar-

sus 4-III white scaled in apical 0.1-0.4” means the white scales of hind tarsal segment

4 are found in a narrow to broad band in the apical 0.1 to apical 0.4 of the segment.

In the original citations, an asterisk marks those taxa for which the type specimen

has been seen. Only the most important literature relative to the genus being treated

is listed following the generic citation or synonymy, but all references seen are cited

under each species. Descriptions of species and drawings are based on topotypic ma-

terial whenever possible. The descriptions are usually composite, that is, based on

more than one specimen. Drawings were usually prepared from a single specimen

and then corrected to show the modal condition of several specimens for the taxo-

nomically important features. Besides the usual illustrations of the fourth instar lar-

va, pupa and whole genitalia of each species and an adult representative of each

group, several other drawings are included. For each species these are the larval man-

dible, or maxilla, or both; claws of the adult male; lateral aspect of the basal mesal

4 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

lobe and proctiger of the male genitalia; and dorsal and ventral aspects of the aedea-

gus or phallosome. In addition, for some species younger instar larvae or notable ex-

tremes in variation in any stage or structure are illustrated. On the distribution maps,

solid symbols indicate localities from which I have examined specimens and hatched

symbols indicate reliable literature records. The Official Standard Names Gazetteers

of the United States Board on Geographic Names have been used in compiling the

distribution lists.

Collection data of collections made expressly for the project ““Mosquitoes of Mid-

dle America”’ are being published in a series of papers. For general information the

first publication of this series should be consulted (Belkin and Heinemann 1973).

For more information on collections cited in this paper, the following publications

of the series should be consulted: for Costa Rica, Heinemann and Belkin 1977a; for

Guatemala, Honduras and Nicaragua, Heinemann and Belkin 1977b; for Mexico, Hei-

nemann and Belkin 1977c; for Panama and the Canal Zone, Heinemann and Belkin

1978a; for Venezuela and the Guianas, Heinemann and Belkin 1978b; and for Co-

lombia, Heinemann and Belkin 1978c.

Throughout the text, the word “larva’’ refers to the fourth instar larva unless pre-

ceded by an adjective indicating otherwise and the word “‘trichoprosopon”’ is used

for convenience to refer to all the species included in this study, i.e. the ones includ-

ed in the single genus 7Jrichoprosopon by Lane and Cerqueira (1942:484-529).

SYSTEMATICS

TAXONOMIC HISTORY. The previously described nominal species included in

the genus 7richoprosopon by Lane and Cerqueira (1942:484-529) have had a long

and varied history of generic placement. This is due in part to the large number of

independent workers in the early twentieth century, in part to changing generic con-

cepts and in part to strictly nomenclatural matters. Three species that came to re-

side in Trichoprosopon were described in the nineteenth century: longipes Fabricius

1805 and digitatus Rondani 1848 in the genus Culex and perturbans Williston 1896

in Aedes.

The period of greatest descriptive activity was the first decade of the twentieth

century, with Theobald describing five species of trichoprosopons, Lutz four, and

Dyar and Knab nine. Three of Theobald’s species were the types of new genera pro-

posed by him. These combinations were: Trichoprosopon nivipes Theobald 1901,

Goeldia fluviatilis Theobald 1903 and Runchomyia frontosa Theobald 1903. His

other two species were Wyeomyia lunata Theobald 1901 and Phoniomyia magna

Theobald 1905. One of Lutz’s species, pallidiventer Lutz 1904, was the type of his

new genus Hyloconops; his other three species were Trichoprosopon splendens Lutz

1904, Trichoprosopon compressum Lutz 1905 and Hyloconops longipalpis Lutz

1905. Most of Dyar and Knab’s species were described in the genus Lesticocampa:

leucopus, rapax, ulopus and vonplesseni in 1906; culicivora in 1907; and schedocyc-

lia in 1908. When Dyar and Knab proposed the genus Lesticocampa in 1906, they

stated the type species was Wyeomyia lunata Theobald 1901; but the species before

them was actually rapax (see Dyar and Knab 1906a:137 and Howard, Dyar and

Knab 1915:165, 166). Other species described by these authors were Phoniomyia

homotina Dyar & Knab 1906, Joblotia mogilasia Dyar & Knab 1907 and J. trichor-

ryes Dyar & Knab 1907. The genus Joblotia Blanchard 1901 had been proposed as

a replacement for Theobald’s Trichoprosopon, which was erroneously thought to be

Zavortink: Reclassification of Trichoprosopon 5

preoccupied. Several other generic names applied to species of Trichoprosopon were

published during this period; Rhynchomyia Lutz 1904 as an unjustified emendation

of Runchomyia Theobald 1903; Binotia Blanchard 1904 as a replacement for Runch-

omyia Theobald 1903, which Blanchard also believed was preoccupied; and /sosto-

myia Coquillett 1906. This latter name is of particular interest, for it is also based

on a misidentified type species. Coquillett stated the type to be Aedes perturbans

Williston 1896, but the specimens before him were actually Culex conservator Dyar

& Knab 1906 (see Dyar and Knab 1906b:221-222; Howard, Dyar and Knab 1915:

187, 219-220, 310; and Edwards 1930:301; 1932:217). An important synonymy

established during this decade was that of Trichoprosopon nivipes with Culex digita-

tus by Dyar and Knab (1907:206, 207).

During the decade from 1910 to 1919, several additional specific names and one

generic name were proposed. The names were: Lynchiaria paranensis Brethes 1910,

Lesticocampa dicellaphora Howard, Dyar & Knab 1913, L. lampropus Howard, Dyar

& Knab 1913, L. espini Martini 1914, Trichoprosopon wilsoni Ludlow 1918, L. trich-

opus Dyar 1919 and L. moralesi Dyar & Knab 1919. In the third volume of their

classical study of mosquitoes, Howard, Dyar and Knab (1915) placed the Central

American species of this group known to them in three genera, Wyeomyia, Lestico-

campa and Joblotia. Although they were unacquainted with the West Indian /sosto-

myia perturbans (Williston), they believed it probable that /sostomyia was a syno-

nym of Lesticocampa. Fabricius’s longipes was first associated with this group of

mosquitoes in this work (as Lesticocampa longipes).

In the decade of the twenties, only a single species, 7richoprosopon shropshirei

Ludlow 1920, was described. However, changes in the classification of this group ap-

peared in several works during this time. Bonne-Wepster and Bonne (1921:12-18) es-

tablished the synonymy of Jsostomyia, Binotia, Runchomyia, Hyloconops and Lesti-

cocampa with Goeldia Theobald 1903. The following year Dyar (1922:99) synony-

mized Lynchiaria with this genus. In their reclassification of American Culicidae,

Dyar and Shannon (1924:482-483) placed all the trichoprosopons known to them in

three genera, /sostomyia, Goeldia and Joblotia. Isostomyia included the species with

short palpi in both sexes, Joblotia the species with hairs [setae] on the clypeus, and

Goeldia the unspecialized species. Finally, in his ““Mosquitoes of the Americas,”

Dyar (1928:90-109) treated every previously described nominal species of this group

in the same three genera, /sostomyia, Goeldia and Joblotia.

In the 1930’s several names were proposed for South American taxa. These

were: Joblotia splendens var. subsplendens Martini 1931, Goeldia leucopus var. hy-

perleuca Martini 1931, Isostomyia brevipes Lima 1931, Goeldia (Isogoeldia) lueder-

waldti Lane 1936 and Goeldia lanei Antunes 1937. Several additional changes were

made in the supraspecific classification. Edwards (1930:301) proposed Ctenogoeldia

as a subgenus of Goeldia, with Lesticocampa dicellaphora as the type species. Ed-

wards did not accept the usage of Isostomyia for a sabethine, and proposed Jsogoel-

dia with Aedes perturbans as the type for the remaining species with short palpi in

both sexes. He also treated Jsogoeldia as a subgenus of Goeldia. In his world catalog

of mosquitoes, Edwards (1932:70-73) placed the species with hairs [setae] on the

clypeus in Trichoprosopon, and all remaining species in Goeldia. Within the latter

genus, he recognized three subgenera, Ctenogoeldia for the species with short palpi

in both sexes and silver scales on the scutellum, Jsogoeldia for the less ornamented

species with short palpi in both sexes, and Goeldia. He (Edwards 1932:71) consid-

ered it possible that the subgenus Goeldia should be further divided into additional

subgenera. In his catalog of neotropical Culicidae, Lane (1939: 158-166) adopted

6 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Edwards’ classification of Trichoprosopon and Goeldia.

In their reclassification of American Sabethini, Lane and Cerqueira (1942:484-

529) united the genera Trichoprosopon and Goeldia, and recognized seven subgenera

within the resulting composite genus. These were: Trichoprosopon, for the species

with hairs [setae] on the clypeus; Limamyia, a monotypic subgenus based on Jsosto-

myia brevipes; Vonplessenia, another monotypic subgenus, based on Lesticocampa

vonplesseni; Ctenogoeldia, for the species with silver scales on the scutellum; /sogoel-

dia, for the dull species with short male palpi; Shannoniana, for fluviatilis, the type,

and related species, with this name to be used in place of Goeldia, which was now

known to be preoccupied; and Hyloconops, for the remaining species. Lane and Cer-

queira also described 10 new species in 1942, obscurum and soaresi in Trichoproso-

pon sensu stricto; walcotti in the subgenus Ctenogoeldia; and castroi, edwardsianus,

evansae, humboldti, reversus, similis and theobaldi in the subgenus Hyloconops.

Stone (1944:335-341) used the name Runchomyia in place of Hyloconops and de-

scribed two additional taxa, Trichoprosopon (Runchomyia) cerqueirai and T. (T.) di-

gitatum var. townsendi.

The year 1953 was the last in which studies significant to the history of Trichopro-

sopon appeared. Levi-Castillo (1953:63-70) described two new species from Ecua-

dor, andinus in the nominate subgenus and cotopaxensis in the subgenus Hyloconops.

Lane (1953:813-861), in his ““Neotropical Culicidae,” followed his earlier work with

Cerqueira in the treatment of Trichoprosopon, except that Rhunchomyia (sic) was

used in place of Hyloconops and the taxonomic changes suggested by Stone (1944)

were incorporated. Unfortunately, some errors were introduced into Lane’s 1953

treatment of Trichoprosopon in English from the original 1942 reclassification by

Lane and Cerqueira in Portuguese. Included among these are the reversing of the spe-

cies names on the drawings of the male genitalia of humboldti and lunatum.

In their synoptic catalog of mosquitoes, Stone, Knight and Starcke (1959:73-77)

followed Lane’s 1953 treatment of Trichoprosopon, except that Isostomyia was used

in place of [sogoeldia.

Neither the generic limits nor the internal classification of Trichoprosopon pro-

posed by Lane and Cerqueira (1942:484-529) have been challenged before, and these

authors’ treatment of Trichoprosopon is followed in the new world catalog of mos-

quitoes by Knight and Stone (1977:312-316) except for a single change in subgeneric

placement made by Belkin (1968b:34, 35). However, the stage for major changes in

sabethine taxonomy was set by Belkin (1962a:486) when he pointed out that “The

internal classification of the tribe is in a chaotic condition. It has been based almost

entirely on very superficial external adult characters and, to a lesser extent, on male

genitalia. Larval and pupal characters have been neglected almost entirely; it is be-

coming increasingly evident, however, that they will be of greatest value in arriving at

a natural classification of the group.”’ Later, writing specifically about 7richoproso-

pon, Belkin (1968b:34, 35) stated that the subgeneric classification of Lane and Cer-

queira was “artificial and unsatisfactory” because these authors “‘completely disregar-

ded excellent characters in the immature stages.’’ Because he had not studied the en-

tire genus thoroughly, Belkin made only one subgeneric change at that time, the

transfer of pallidiventer from the subgenus Runchomyia to Trichoprosopon sensu

stricto.

The American trichoprosopons have always been considered to be generically dis-

tinct from the generalized Old World sabethines now placed in the genera 77ipteroti-

des and Maorigoeldia. However, according to Lee (1946:223-224), Trichoprosopon

and Tripteroides are very close and there is little to separate them. He points out

Zavortink: Reclassification of Trichoprosopon 9

that the range of variation of adult and male genitalic characters in Tripteroides falls

within the broader range of variation of Trichoprosopon. Furthermore, he notes that

there is parallel development in larval features in both groups. Lee argues for separa-

ting the genera, though, on the basis of the presence of a pectin in the larva of Trip-

teroides and its absence in Trichoprosopon, and on the basis of the allopatric distri-

butions of the groups. Belkin (1962a:495-496) apparently disagreed with Lee’s as-

sessment of variation in these genera, for he considered Tripteroides and Trichopro-

sopon to be amply distinct in all stages.

TAXONOMIC CHARACTERS. The trichoprosopons are a very difficult group

taxonomically. The species in a particular complex are often distinguished by char-

acteristics so minor they would not be considered significant in other genera. This is

true not only in the case of the adults, but frequently also in the case of the male

genitalia and immatures. Another factor contributing to the difficulty of the group

is that in some instances species easily distinguished in one stage are indistinguishable

in another. Because the differences between the species may be so subtle or may be

limited to only some stages, accurate identification requires excellent quality mater-

ial consisting of adults of both sexes with associated larval and pupal exuviae.

Adult characters. Most taxonomic characteristics are found in the distribution of

setae on various parts of the body, particularly the sclerites of the pleuron; various

details of the scales, especially on the thorax; and the markings of the legs and abdo-

men. Important setal characters are: presence or absence of setae on clypeus and

ppn; number of setae on coxa I; position of setae on stp and mep; presence or ab-

sence of setae on psp, and their number and strength when present; and presence or

absence of setae on upper calypter of wing, and details of their distribution when

present. Setae are sometimes developed in unexpected places; one undescribed spe-

cies of Shannoniana has long, hooked setae on the abdominal sternites. Important

characteristics related to the scales include: size, shape, color, and strength and color

of iridescence of scales on dorsum of head and thorax; and distribution, color,

strength and color of iridescence, density, and direction of imbrication of scales on

pleuron. Some of the pleural scale characteristics are very difficult to use, either be-

cause they require excellent optics to be seen, as in the case of the direction of imbri-

cation, or because the differences are comparative or relative and difficult to describe

objectively, as in the case of subtle differences in color and density. The markings of

the legs, particularly the distribution of light scales on the tibiae and tarsi, are ex-

tremely important, but do present some problems. These light markings may be con-

spicuous and clearly demarked, or they may be rather inconspicuous and poorly set

off, or visible at only some angles of observation or present on only one surface of

the leg. Also, the development of the light tarsal markings may be extremely variable

in some species. Markings of the abdomen include: shape of light patches on tergites

and presence or absence of dark scales on sternites, and intensity of pigmentation

and distribution of these dark scales when present. The sternal markings are often

difficult to use because the sternites may be completely hidden by the tergites, or be-

cause the markings themselves may be weakly developed and poorly demarked or vis-

ible at only some angles of observation.

Other features that may be important in distinguishing species include: length of

palpus in both sexes; length of proboscis; length of thorax, for it is shortened or

lengthened in some species; intensity of pigmentation of scutal integument and parts

of pleural integument; relative lengths of particular leg segments; and development of

outstanding scales on legs. Additional characteristics of the male include: plumosity

of flagellum and size, shape and dentition of claws of legs I and II.

8 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Male genitalic characters. There are many excellent characters in the male genita-

lia, especially in minute details of tergite IX and the aedeagus. Both whole mounts

and carefully made dissections must be studied in order to see all the important char-

acteristics. Except in Shannoniana, where the male genitalia of many species are vir-

tually identical, most species can be separated on the basis of the genitalia.

Briefly, characters I have found to be of value are as follows: for tergite VIII: pre-

sence or absence of strong setae; for tergite IX: development of basal apodeme, num-

ber, size, and shape of lobes, shape of separation between lobes, number and curva-

ture of rows of setae on lobes, and number, length, strength, and shape of setae; for

the sidepiece: length, shape, distribution and development of setae, and presence or

absence of spicules on tergal surface; for the basal mesal lobe: distinctness and shape

of lobe, and number, length, strength and curvature of its setae; for the clasper:

length, strength, shape, and length, shape and dentition of spiniform; for the aedea-

gus: size, shape, details of tergal arms, and development of a beak, crest, flanges,

teeth or spicules apically; and for the paraproct: number, size and shape of teeth,

and number and distribution of cercal setae.

I think it is worth mentioning here that I believe the various components of the

phallosome of the culicids have been misinterpreted and confused, and should be

carefully restudied in all major groups of the family. Not only would a better under-

standing of the structure and functioning of the genital apparatus result from such a

study, but probably also a better understanding of the relationships of the higher

taxa. Briefly, I am of the opinion that the “‘aedeagus” of most sabethines and of

some other mosquitoes, as for example the aedines I studied previously (Zavortink

1972), is homologous to the opisthophallus of Maorigoeldia argyropus (Walker 1848)

(Belkin 1962a:492, 493; 1968a:8, 9, 107, 108), and not the aedeagus of that primi-

tive mosquito. It is probably only the “‘sternal plates” of the ‘“‘aedeagus’’ of some

sabethines that are derived from the true aedeagus. Since my opinions on genitalic

homologies are based, admittedly, on limited casual observations, I choose to contin-

ue using the existing terminology until such time as a careful comparative study can

be completed.

Pupal characters. There are many excellent taxonomic characters in the pupae

and, with only a few exceptions, the pupa of every species is distinctive. In fact, in

Shannoniana and Runchomyia sensu stricto, pupae of related species may be remark-

ably different. There are too many characters relating to chaetotaxy to be men-

tioned here; it must suffice to say that the strength, length, number of branches, and

position of many of the dorsal setae are important in distinguishing species or groups.

Other important characters are: pigmentation of integument; presence or absence of

dorsal sensillum; shape and pigmentation of trumpet; shape of tergite VIII; size of

male genital lobe; and size, shape, serration, spiculation and development of midrib

of paddle.

In many of the species of this study, and in many other New World and Old World

sabethines, seta 1-C of the pupa is long, double, sigmoidally curved and has the bran-

ches hooked at the apex. This specialized seta is apparently used to anchor the pupa

to objects below the surface of the water (Iyengar and Menon 1948:41, 42). At this

time it is not known if this specialization has arisen repeatedly, or if it has evolved

only once and therefore indicates a close relationship for those groups that possess it.

Belkin (1962b:359) erroneously shows seta 1-C as unmodified in Malaya solomonis

(Wharton 1947).

I have not seen seta 14-ITI-VII in the-pupa or larva of any specimens examined dur-

ing this study, and I think it can be safely concluded that it is not developed in these

mosquitoes.

Zavortink: Reclassification of Trichoprosopon 9

Larval characters. The number of excellent taxonomic characters in sabethine lar-

vae is truly incredible. There are numerous structural features of the head capsule,

mouthparts and terminal segments that are, I believe, of great phylogenetic signifi-

cance. And, there are literally countless setal characters that are important in delimi-

ting both species and groups.

Setae on the thorax and siphon are sometimes very obviously specialized morpho-

logically, but the biological significance of these specializations is largely unknown.

Iyengar and Menon (1948:39-41) observed that the larva of Malaya genurostris Leice-

ster 1908 (as Harpagomyia genurostris) attaches itself to the side of its habitat with

specialized siphonal setae, and supports itself with these same setae and setae of the

lateral prothoracic groups when it rests on its back at the bottom. It is unfortunate

that so few observations have been made on living mosquitoes that the functional sig-

nificance of almost all characters used in classifying the group is unknown. In order

to erect a natural classification, it is essential that systematists be able to distinguish

among similarities due to relationship, parallelism and convergence, and such distinc-

tions can be made only when the functional significance of the characters in question

is known.

It will also be necessary to study the younger instar larvae of the sabethines before

a natural classification of the tribe can be achieved. There are often remarkable shifts

in position and changes in development of setae from one instar to the next in a par-

ticular species. It is clear in groups like Shannoniana, where the fourth instar larvae

of most species differ conspicuously in many setal characteristics, that many of the

specific differences of the mature larvae result from only differing degrees of neoteny

in the various species. In groups like this, major changes in larval morphology may

evolve very quickly in response to relatively few changes in genes controlling develop-

ment, so that the amount of morphological divergence in the fourth instar larvae is

not an accurate reflection of overall genetic divergence.

In at least Johnbelkinia ulopus there is a remarkable change in the development of

the maxilla from one instar to the next. In the second instar larva (fig. 7), the maxil-

la is much longer than the head and has no apical spines; in the third instar (fig. 6), it

is shorter than the head and has both a rigid and an articulated apical spine.

Structural features of the head capsule that are important taxonomically are: de-

velopment of the foramen magnum, which may be circular and bounded by a distinct

collar or slitlike and without a collar, and course of the maxillary suture, which may

be straight or curved, and which may or may not extend to posterior tentorial pit.

The phylogenetic significance of the development of the foramen magnum is un-

known; the slitlike character state was first noticed only recently (Belkin, Heinemann

and Page 1970:19) and its distribution in the family is unknown. I have observed

that the slitlike foramen magnum occurs also in some Old World sabethines and in

Aedeomyia squamipennis (Lynch Arribalzaga 1878). Structural features of the

mouthparts that are important are: modification of mandible for grasping; modifica-

tion of maxilla for grasping; form of maxillary palpus; and articulation of maxilla

with head capsule. The modifications of the maxilla seem to be particularly numer-

ous and significant. In some sabethines the maxilla is more or less normal, whereas

in others it is variously modified, either extremely elongate or with strong apical

spines, apparently for grasping. In some groups the maxillary palpus is an elongate,

cylindrical structure completely free from the body of the maxilla; in others the pal-

pus is partly to nearly completely adnate to the body of the maxilla. In Shannoniana

there is a very strong articulation between the maxilla and head capsule far ventrad

of the palpus that is not developed in the other groups studied.

10 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Important structural features of the terminal segments include: presence or ab-

sence of comb scales, and their shape, number and arrangement when present; shape,

length and spiculation of siphon; development of accessory midventral setae on si-

phon; and presence or absence of a pecten on siphon. The trichoprosopons have gen-

erally been regarded to lack a true pecten. However, the fringe of long spicules on

the ventral surface of the siphon in Runchomyia represents the pecten. In R. magna

the first instar larva has a normal pecten of enlarged teeth in a pair of ventrolateral

rows; the second instar larva has a single ventral row of elongate, filamentous teeth;

and the third and fourth instars have a single ventral row of very numerous, long, fil-

amentous spicules. Further, in fourth instar larvae of R. lunata and R. perturbans

the distal elements of the pecten are normally developed teeth in separate rows.

Other larval characters that I have found useful are: pigmentation of sclerotized

parts of body; shape of mental plate; shape, length and dentition of mandible and

maxilla; and presence or absence of dorsal sensillum.

For the setae, the strength, length, type of development, number of branches, pig-

mentation, position and presence or absence of an alveolar plate may all be impor-

tant in particular cases. Specializations in setae 4-P and 2,6-S often provide good

taxonomic characters. Seta 8-M, which is a large multiple seta in most mosquito lar-

vae, is completely absent in all species of Trichoprosopon, and weakly developed in

all species of Shannoniana.

The immatures of the Sabethini seem to be particularly favorable material for ob-

serving the nerve connections between setae of one instar or stage and the developing

subsequent instar or stage, and I have been able to unquestionably homologize many

larval and pupal setae in all major groups of this study. Setae that I have been able to

homologize in each group will be listed in the revision of that group. At this time,

though, I would like to indicate one correction that needs to be made in a previously

published illustration. In the larva of Malaya solomonis illustrated by Belkin (1962b:

360), the labels on setae 3 and 4 on segments III and IV should be reversed. Tracing

of the nerves in Runchomyia magna, the larvae of which is similar to that of M. solo-

monis, shows the first seta mesad of seta 1 to be seta 4, not seta 3, on these segments.

PROPOSED CLASSIFICATION. Forty-five of the 48 nominal species included in

Trichoprosopon at the onset on my study (see Knight and Stone 1977:312-316) and

19 undescribed related species known to me at this time fall into six distinct groups

apparently representing four separate phyletic lines. Of the three remaining nominal

species, one belongs in the genus Wyeomyia, and two are considered to be nomina

dubia.

The four phyletic lines are most clearly demarked from each other in the larval

stage, but they are amply distinct in the adult and pupal stages, and usually also in

the male genitalia. Because of the distinctness of these lines, and because some may

have close affinities with mosquitoes presently placed in other genera, I am recogni-

zing each as a separate genus at this time. The four genera, some of the more impor-

tant features for distinguishing them from each other, their possible affinities and the

previously described nominal species belonging to each are summarized below.

I consider my reclassification of this group to be tentative only, and subject to

change when more information is available. Biologists, including even some systema-

tists, often lose sight of the fact that a biological classification is a scientific hypothe-

sis, and, like all other hypotheses, it may need to be changed or discarded when new

data are discovered. The tribe Sabethini is a large one that has been poorly collected

and little studied in both the New and Old Worlds, so that many additional data of

taxonomic value await discovery, and as they are discovered, changes in the classifica-

Zavortink: Reclassification of Trichoprosopon 1]

tion of the group will be inevitable. A classification, again like any other hypothesis,

has heuristic value, so that its very existence should stimulate further research that

may ultimately bring about its modification or demise.

Genus Johnbelkinia. This is the smallest segregate genus, with but three taxono-

mic species, all of which were placed previously in Trichoprosopon (Runchomyia).

The larvae are distinguished by a long row of multibranched accessory midventral se-

tae on the siphon; adults by the combination of a long proboscis and light markings

on tarsi IT and III; pupae by presence of a dorsal sensillum on only segments IV and

V; and male genitalia by a large patch of very dense, strongly developed setae on side-

piece. The affinities of Johnbelkinia are unknown, as I have seen no sabethines from

either the New World or the Old World that are similar. Nominal species belonging

to this genus are: Lesticocampa culicivora Dyar & Knab 1907, Trichoprosopon (Hy-

loconops) edwardsianus Lane & Cerqueira 1942, Lesticocampa leucopus Dyar &

Knab 1906, Culex longipes Fabricius 1805, Lesticocampa trichopus Dyar 1919 and

Lesticocampa ulopus Dyar & Knab 1906.

Genus Trichoprosopon. This is the largest genus, with 21 taxonomic species, sev-

eral of which are undescribed. The group includes not only the species with setae on

the clypeus that have always been included in it, but also species lacking such setae

that have been placed in the subgenera Limamyia, Vonplessenia and Runchomyia.

Larvae of the group are distinguished by the unmodified maxilla and absence of seta

8-M; adults by a short proboscis in concert with absence of a postmedian light band

on tibia III; and pupae by weak and short seta 5-II-VII. The male genitalia are not

much differentiated from those of Shannoniana or Runchomyia sensu stricto. This

genus seems not to have any close relatives. Nominal species belonging to Trichopro-

sopon are: Trichoprosopon (Trichoprosopon) andinus Levi-Castillo 1953, Isostomyia

brevipes Lima 1931, Trichoprosopon (Hyloconops) castroi Lane & Cerqueira 1942,

Trichoprosopon compressum Lutz 1905, Culex digitatus Rondani 1848, Trichoproso-

pon (Hyloconops) evansae Lane & Cerqueira 1942, Lesticocampa lampropus Howard,

Dyar & Knab 1913, Goeldia lanei Antunes 1937, Joblotia mogilasia Dyar & Knab

1907, Trichoprosopon nivipes Theobald 1901, Trichoprosopon (Trichoprosopon) ob-

scurum Lane & Cerqueira 1942, Hyloconops pallidiventer Lutz 1904, Trichoproso-

pon (Hyloconops) similis Lane & Cerqueira 1942, Trichoprosopon (Trichoprosopon)

soaresi Lane & Cerqueira 1942, Trichoprosopon splendens Lutz 1904, Joblotia splen-

dens var. subsplendens Martini 1931, Trichoprosopon (T.) digitatum var. townsendi

Stone 1944, Joblotia trichorryes Dyar & Knab 1907, Lesticocampa vonplesseni Dyar

& Knab 1906 and TJrichoprosopon wilsoni Ludlow 1918.

Genus Shannoniana. This is a large genus, with 15 species. Most of these are un-

described, and I suspect scores more await discovery in the field. The previously de-

scribed species were all correctly associated in the subgenus Shannoniana by Lane

and Cerqueira (1942:502-506). Larvae are distinguished by the modified maxilla

and its unusual articulation with the head capsule; adults by the postmedian light

band on tibia III; pupae by absence of a dorsal sensillum on any segment in combina-

tion with a well developed seta 5 on at least segments III-V; and male genitalia by

large preapical teeth on the aedeagus in combination with a pair of lobes on IX-T. I

believe Shannoniana is closely related to the subgenus Rachisoura of Tripteroides

from the Australian zoogeographical region. There are several striking similarities in

structure of the larval head capsule in these groups, including the virtually identical

development and articulation of the maxilla. Belkin (1962a:529) earlier noted the

striking similarity between Rachisoura and some Trichoprosopon. Included nominal

species are: Goeldia fluviatilis Theobald 1903, Hyloconops longipalpis Lutz 1905,

12 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Lesticocampa moralesi Dyar & Knab 1919 and Lesticocampa schedocyclia Dyar &

Knab 1908.

Genus Runchomyia. This is a complex and poorly known group comprising at

least 11 species in three subgenera, Runchomyia sensu stricto, Ctenogoeldia and Iso-

stomyia. My treatment is possibly too conservative, and the subgenera perhaps de-

serve generic rank. The larvae of Runchomyia are distinguished by the slitlike fora-

men magnum and a filamentous pecten on the siphon; adults by long proboscis to-

gether with dark tarsi; and pupae by presence of a dorsal sensillum on segments III-V.

The male genitalia of Runchomyia exhibit a wide range of variation; those of Cteno-

goeldia and Isostomyia are unusual and unlike any other New World sabethines, but

those of the nominate subgenus are generalized and can not be distinguished as a

group from those of Trichoprosopon. The affinities of Runchomyia are problematic

at this time. The Old World Malaya and Topomyia, the New World Sabethes, Lima-

tus and Phoniomyia, and most of the New World Wyeomyia have the slitlike foramen

magnum, so the affinities most likely lie with these groups. The resemblance in adult

ornamentation, male genitalia and chaetotaxy of the immatures between Runchomy-

ia (Ctenogoeldia) magna and some species of Malaya is great enough to suggest a rela-

tionship. The three subgenera of Runchomyia are distinguished by characters given

in the key that follows. Included species are: for Runchomyia sensu stricto: Tricho-

prosopon (Runchomyia) cerqueirai Stone 1944, Runchomyia frontosa Theobald

1903, Trichoprosopon (Hyloconops) humboldti Lane & Cerqueira 1942, Lesticocam-

pa rapax Dyar & Knab 1906, Trichoprosopon (Hyloconops) reversus Lane & Cerque-

ira 1942 and Trichoprosopon (Hyloconops) theobaldi Lane & Cerqueira 1942; for

subgenus Ctenogoeldia: Lesticocampa dicellaphora Howard, Dyar & Knab 1913,

Phoniomyia homotina Dyar & Knab 1906, Phoniomyia magna Theobald 1905 and

Trichoprosopon (Ctenogoeldia) walcotti Lane & Cerqueira 1942; for subgenus Isosto-

myia: Lesticocampa espini Martini 1914, Wyeomyia lunata Theobald 1901, Lynchi-

aria paranensis Brethes 1910, Aedes perturbans Williston 1896 and Trichoprosopon

(Joblotia) shropshirei Ludlow 1920.

Excluded Species. I have examined a female specimen of Goeldia (Isogoeldia) lue-

derwaldti Lane 1936:6-8 from the FH collection. This specimen agrees with the ori-

ginal description of luederwaldti and was determined as that species by Lane himself.

Since I find this specimen to be similar to, and perhaps even the same as, Wyeomyia

(Dendromyia) personata (Lutz 1904), this nominal species should be transferred to

Wyeompia.

Nomina Dubia. The holotype of Goeldia leucopus var. hyperleuca Martini 1931:

201 has been lost (see Belkin 1971:31), and it is not certain from the brief original

description what known taxonomic species Martini had before him. Therefore I

think it best to consider the name a nomen dubium. I have seen the single female

specimen from Restrepo, Colombia, in the FH collection that Lane and Cerqueira

(1942:526-527) and Lane (1953:858) treated as Trichoprosopon hyperleucus, and I

consider it to be Johnbelkinia ulopus. Martini’s description of hyperleuca does not

quite fit known specimens of ulopus, and since the type was collected about 1600km

south of the known range of that species along the eastern slope of the Andes, I am

hesitant to simply synonymize hyperleuca with ulopus.

The location of the holotype male of Trichoprosopon (Hyloconops) cotopaxensis

Levi-Castillo 1953:66-68 is unknown, and in all probability the specimen is nonexis-

tent. The original description of cotopaxensis in based on specimens representing at

least two, and most likely three, different species. The male genitalia described and

illustrated by Levi-Castillo are Johnbelkinia ulopus and the pupa illustrated is Tricho-

Zavortink: Reclassification of Trichoprosopon La

prosopon digitatum (see next paragraph). According to the description, the legs of

the adult were dark, but both ulopus and digitatum have conspicuous light markings

on the legs, which indicates that yet a third species was involved. The lengths given

for the palpi of both the male and the female are too great for ulopus, and the habi-

tat (bamboo stump) in which the pupae of the type series were collected is not one

normally utilized by ulopus. Two dark-legged trichoprosopons, Runchomyia magna

and Trichoprosopon andinum, are known from coastal Ecuador, and a third, T. palli-

diventer, might logically occur there, but Levi-Castillo’s description seems not to be

based on any of these. The culicid fauna of the Pacific lowlands of Ecuador is the

most highly endemic in South America, and cotopaxensis may be a distinct species

unknown elsewhere. Since this can not be determined without the holotype male, I

consider the name to be a nomen dubium.

Levi-Castillo described numerous new species of mosquitoes from Ecuador in the

period 1944-1955; included were two trichoprosopons, Trichoprosopon cotopaxensis

and T. andinus, described in 1953. The holotypes of most of his species were depos-

ited in the Centro Ecuatoriano de Investigaciones Entomologicas in Guayaquil, but

the location of the types of some of his species, including the two trichoprosopons,

was never known. In the late 1950’s Alan Stone wrote Levi-Castillo several times,

each time asking for the location of the types of andinus and cotopaxensis (and oth-

ers). These letters were never answered. In the summer of 1961 Richard F. Darsie,

Jr., visited Levi-Castillo in Ecuador, and obtained from him 564 slide preparations of

mosquitoes. Included among these were supposed to be the slides of types. Darsie

did not obtain any adult mosquitoes, as these were allegedly packed away. The slides

obtained by Darsie were subsequently sent to the USNM. In 1961 Stone wrote to

Levi-Castillo and asked if he could provide more information about the slides, as

more than half were unlabeled, and suggested that he consider depositing his adults

in the USNM also. Although Levi-Castillo responded the following year, he did not

provide any additional information about the slides or the whereabouts of his adult

collection. I have located 54 slides of trichoprosopons in Levi-Castillo’s material at

the USNM. Only three species are represented, T. digitatum, T. andinum and J. ulo-

pus. The material of ulopus consists of five slides of dissected male genitalia. Four

are totally unlabeled. The fifth bears two standard slide labels, one pasted over the

top of the other. The upper label states simply “‘T. / cotopaxensis.”” The lower can

be deciphered by holding the slide up to a bright light, and it states ‘““Trichoprosopon

/ (T.) obscurum / Valencia / Los Rios.” Levi-Castillo’s genitalia preparations are

among the worst I have ever seen. The genitalia are so irregularly fragmented, and

the pieces so disoriented and squashed, that they are almost unrecognizable. One re-

sult, though, is that each slide is unique, with its own peculiar set of artifacts. On

this basis, I believe it can be stated with certainty that the genitalia illustration in the

original description of cotopaxensis was prepared from the slide of ulopus labeled T.

cotopaxensis. For digitatum there are 29 slides, mostly of dissected male genitalia,

and mostly unlabeled. There are, however, four slides of pupal skins. Of these, only

one pupa has the abdomen intact and bearing most of its setae; this slide is labeled

“Trichopro / sopon / V.P.”’ I believe the pupal illustration accompanying the original

description of cotopaxensis was made from this slide of digitatum. The specimen has

the float hairs (setae 1-I) rotated caudad and partly covering segment II, seta 6-VI

forked near the middle on the left side, the paddles slightly overlapped and is female,

and the illustration accurately reflects these peculiarites.

14 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

BIONOMICS

Under natural conditions the immature stages of all trichoprosopons occur in liq-

uids associated with plants. Specific habitats include rot holes in trees, fallen fruits

and nuts, bamboo stumps, punctured bamboo internodes, fallen leaves and palm

spathes, leaf axils of many kinds of living plants and flower bracts of species of Cala-

thea and Heliconia. Most species are specific to one or a few similar habitats, but

others are more catholic. In most cases the breeding sites hold a relatively large quan-

tity of fluid that is largely or entirely accumulated rain water. However, some of the

leaf axil breeding sites do not contain open water, but only a thin mucilaginous film

that may be of plant origin between the appressed leaf bases.

In all species of trichoprosopons the larval mouthparts are modified. In the genus

Trichoprosopon it is the mandible that is enlarged, while in the genera Johnbelkinia,

Shannoniana and Runchomyia it is the maxilla. The modifications of the mouth-

parts have generally been considered to be for predation, and indeed several species

have been reported as being predaceous, especially on Wyeomyia, or cannibalistic.

However, according to Galindo, Carpenter and Trapido (1951:133) many sabethine

larvae that have been reported to be predaceous are only facultatively so, and do not

as a rule feed on other mosquito larvae. These authors also state “the larvae of these

sabethines are not predaceous in the same sense as Toxorhynchites, Psorophora

(Psorophora) spp. and Culex (Lutzia) spp. since they do not swallow the prey, but

usually grab it by the mid-abdomen, breaking it in two, and apparently sucking the

body juices.”’ Assem (1959:36-37), writing about New Guinean species of Tripter-

oides (Rachisoura), reports that the modified maxillae of these larvae are used as clas-

ping organs to hold large food particles as well as in aggressive behavior between lar-

vae. As for the latter, he reports that larvae may fight until the death of one, which

is then consumed by the other. However, not all such fights end in cannibalism,

since one duel he observed lasted two hours before ending in a draw with both larvae

surviving.

In light of the above, the supposed predatory behavior of all trichoprosopons

needs to be reexamined. It should be stressed also that, in so far as possible, observa-

tions should be made under natural conditions. I collected large numbers of Runch-

omyia magna larvae from individual Calathea bracts in Venezuela; only this one spe-

cies of mosquito was present in these bracts at this stage in their development. When

these same larvae were placed together in vials in the laboratory, many were subse-

quently found dead, having been bitten in two. The simple environment within a vial

is in such stark contrast to the complex one within a Calathea bract, where there are

countless cavities and crevices among the flowers and their parts, that I am not pre-

pared to argue that this species is normally cannibalistic on the basis of my observa-

tions in the laboratory.

Very little is known about the biology or ecology of adult trichoprosopons. Sev-

eral species are attracted to and bite man and other mammals, and several are known

to be diurnal.

As already mentioned, the trichoprosopons are a difficult group taxonomically,

and misidentification of the species has been very common. This difficulty in accu-

rately identifying the species has undoubtedly hindered the gathering and reporting

of biological information about them. The improved classification and keys in the

present revision should facilitate identification of the species, and thereby stimulate

studies of their biology.

2(1).

3(1).

4(3).

5(4).

2(1).

Zavortink: Reclassification of Trichoprosopon 15

KEYS TO GENERA AND SUBGENERA

FORMERLY PLACED IN TRICHOPROSOPON

ADULTS

Proboscis 0.85-1.20 length of femur I; row of lower stp setae extending dor-

sad to above level of lower edge of mep . . . Ae

Proboscis 1.20-1.40 length of femur I; row of lower. stp setae Gepally not ex-

tending dorsad to level of lower edge Of MGs tee tee sd

Tibia III without postmedian light band; laterotergite without scales basad

and sparsely scaled distad, its lower margin visible; pcx without scales

Trichoprosopon

Tibia, na with broad complete o or incomplete postmedian light band; latero-

tergite densely scaled, its lower margin hidden; pcx with scales

iad i Shannoniana

Tarsi II, III partly light scaled; scutal scales moderately broad and flat and

scales of vertex and occiput with brilliant silver and azure blue reflections

Johnbelkinia

ne TL, Tl dad scaled: scutal scales narrow and curved or scales of vertex

and occiput without silver reflections and with only weak to moderate

green or blue reflections (Runchomyia) 29...

Scutal scales moderately broad to broad and flat; scales of vertex without sil-

ver reflections and with only weak to moderate green or blue reflections;

ppn without posterior seta . . . . . subgenus I[sostomyia

Scutal scales narrow and curved; scales of vertex with brilliant silver and azure

blue reflections; ppn with.1 or more posterior setae ....... .5

Scales on base of midlobe of scutellum with brilliant silver reflections ;

ee subgenus Ctenogoeldia

eles on Ricioke ap cumilam witout ae) reflections ‘

subgenus Pe heanie

MALE GENITALIA

Sidepiece without distinctly separated basal mesal lobe (in part Runchomyia)

wivewi te yh derbies ante S tam ieee Wouek Meer h Wht cm eh subgenus Ctenogoeldia

Sidepiece with distinctly separated basal mesallobe .........2

Basal mesal lobe developed as an elongate, weakly or strongly sclerotized

plaque (in part Runchomyia) . . . . . . . Subgenus Isostomyia

Basal mesal lobe developed as a more or less crescentic or semicircular lobe,

the apical part of which may be strongly produced. . ...... .3

16

3(2).

4(3).

5(4).

2(1).

3(2).

Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Basal tergal area of sidepiece with large, very dense patch of strongly devel-

oped, moderately long to long setae; [X-S largely membranous, only base

and sides sclerotized, distal margin of sclerotized portion strongly concave

. Johnbelkinia

Basal tergal ai area , of sidepiece without patch of specialized setae; [X-S largely

or entirely sclerotized, more or less semicircular or crescentic, distal mar-

gin strongly convex throughout or with median apical emargination . .4

Aedeagus with large preapical teeth and IX-T with pair of distinctly separated

submedian lobes bearing a single or he double row of apically curved

and/or flattened setae . . . . . Shannoniana

Aedeagus without large preapical teeth or TX-T with single broad median

lobe, sometimes notched in middle, bearing several rows of straight or

nearly straight spinsdorm setae medially .-. . w S

With the following combination of characters: sidepiece moderately long to

long; aedeagus without large teeth, serrations or spicules apically but with

narrow sharp apical beak; spiniform short to moderately long; paraproct

with 2-6 strong apical teeth; and apex of submedian tergal arm of aedeagus

narrow (in part Runchomyia) ee . . . . subgenus Runchomyia

Without the above combination of sharacters: either sidepiece short; or aede-

agus with large teeth, marginal serrations, or small spicules apically: or ae-

deagus without narrow apical beak; or spiniform long; or paraproct with 1

strong apical tooth; or apex of submedian tergal arm of aedeagus broad .

. Trichoprosopon

PUPAE

Seta 5-III-VI weakly developed and very short . . . . . Trichoprosopon

Seta 5-IV and usually also 5-III,V or 5-V,VI pkia h to strongly devel-

oped and moderately longtolong ...... 7

Seta 5-C weakly developed and short; seta 3-III at the same level as or cepha-

lad of 4-III, far from caudal margin of tergite; dorsal sensillum absent on

tergites III- V ae . . Shannoniana

Seta 5-C moderately to stronsly developed and moderately long to long; seta

3-III caudad of 4-III, near caudal margin of tergite; dorsal sensillum pres-

enon vertes IY Of IVY oe Ne OP re ee

Inner part of paddle spiculose; seta 5-II,III moderately to strongly developed

and moderately long to long and seta 6-VII moderately to strongly devel-

oped, 3-18b, and usually cephalad and laterad of 9-VII; dorsal sensillum

present on tergitesIV,V .. . . Johnbelkinia

Inner part of paddle glabrous; seta 5. -II, I weakly developed and short or seta

6-VII weakly developed, single or double, and caudad and mesad of 9-VII;

dorsal sensillum present on tergites III-V (Runchomyia). ..... . 4

4(3).

5(4).

2(1).

3(2).

4(1).

5(4).

Zavortink: Reclassification of Trichoprosopon 17

Paddle strongly produced apically or seta 5-V,VI weakly developed and short;

caudolateral angle of tergite VIII strongly to very strongly produced, ex-

tending at least 0.50 distance to apex of segment IX

Bid tebe Rastevis

Paddle vl fo triangular pees ete re V VI moderately to strongly developed

and moderately long to long; caudolateral angle of tergite VIII not or only

slightly produced, not extending more than 0.25 distance to apex of seg-

ment IX

Seta 5-C much longer and stronger than 7-C and 3-10b; seta 6-I moderately

strong, weaker than 3-I and/or 6-II. . . . . . subgenus Ctenogoeldia

Seta 5-C subequal in length and strength to 7-C or at least 40b; seta 6-I

strongly developed, as strong as or stronger than 3-I and 6-II . :

A : subgenus Runchomypyia

LARVAE

Head capsule with normal circular foramen magnum bounded by distinct col-

lar; siphon without pecten .. . ae Oe a ae

Head capsule with short to long abate: siilikee foramen wyacng not

bounded by distinct collar; siphon with filamentous pecten (Runchomyia)

4

Maxilla normal, not modified for grasping, never extremely long or with

strong apical spines; maxillary suture extending to posterior tentorial pit;

seta S-Moabsent @ e730 7: . . . . . Irichoprosopon

Maxilla modified for grasping, either extremely long or with strong apical

spines; maxillary suture not extending to posterior tentorial pit; seta 8-M

presents: (aioe, BER Sie oe ee ae te Wea. SO

Siphon without accessory midventral setae; maxilla with strong articulation

with head a far ventrad of palpus; seta 8-M weakly developed

Shannoniana

Siphon with iongt row y of multibranched accessory midventral setae; maxilla

without strong articulation with head capsule far ventrad of palpus: seta 8-

Mostronsiy developed (227 te et Pee ee ohnbelkinia

Seta 3-X 2-8b; seta 6-S strongly developed, rigid, hooked at apex

: subgenus Kovotaiin

Seta 3-X sinele: ‘seta 6i S ently apreioueds Alexie: momnorwedou ss faa 88

Maxilla with rigid apical spine shorter than articulated apical spine; most

comb scales arising from single large plate; seta 5-T strongly developed,

moderately lone: .. 6.54 . . . subgenus Ctenogoeldia

Maxilla with rigid apical spine longer than articulated apical spine; comb

scales free; seta 5-T weakly developed, short . . subgenus Runchomyia

18 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

TAXONOMIC TREATMENT

JOHNBELKINIA Zavortink, gen. n.

TYPE SPECIES: *Culex longipes Fabricius 1805, French Guiana.

Trichoprosopon (Runchomyia) in part of Stone, Knight and Starcke (1959:76-77); Knight and

Stone (1977:313-314). 3

Trichoprosopon (Rhunchomyia) in part of Lane (1953:855-858, 860); Cerqueira (1961a:459-464).,

Trichoprosopon (Hyloconops) in part of Lane and Cerqueira (1942:525-527, 528).

Goeldia (Goeldia) in part of Dyar (1923a:84-85, 86); Edwards (1932:72); Lane (1939:161, 162-

163).

Goeldia in part of Bonne-Wepster and Bonne (1921:17-18; 1922:38); Dyar and Shannon (1924:

482-483); Bonne and Bonne-Wepster (1925:141-146, 152); Dyar (1925: 128-130; 1928:96-98).

Lesticocampa in part of Dyar and Knab (1906a: 137-138); Theobald (1910:558, 622); Howard,

Dyar and Knab (1915:168-174). |

FEMALES. Medium sized to large mosquitoes. Moderately ornamented; scales of

vertex and occiput with brilliant silver or azure blue reflections; pleural scales metal-

lic silver; dark scales largely moderately iridescent.

Head: Eyes contiguous above antennae. Frontal tuft absent. Orbital setae numer-

ous, middle pair elongate. Vertex and occiput densely covered with broad flat scales

with brilliant silver or azure blue reflections; occiput with row of dark erect scales

posteriorly. Lateral scales dense, broad, flat, silver. Clypeus moderate-sized, promi-

nent; brown to dark brown, shiny; without setae or scales. Proboscis not modified;

slender, long, usually 1.20-1.40 length of femur I; dark scaled. Palpus 0.16-0.20

length of proboscis; apparently 2-segmented; dark scaled. Antenna 0.55-0.70 length

of proboscis; torus without scales; flagellar segment 1 with small scales dorsally.

Thorax: Scutal integument brown to dark brown. Acrostichal and dorsocentral

setae present on anterior promontory; supraalar setae numerous; a few anterior dor-

socentral setae and prescutellar setae sometimes present. Scutellum with 4 long setae

on midlobe, usually 3 on lateral lobe. Postnotum with numerous setae. Scutum

without extensive bare areas. Scutal scales dense; most moderately broad, flat,

straight, some narrower, curved; brown, dull to moderately iridescent; broader, al-

ways iridescent posteriorly. Scutellar scales broad, flat, brown, moderately irides-

cent. Postnotum with or without scales. Paratergite bare. Apn lobes moderately

large, not approximated dorsally. Meron small, its upper edge in line with base of

hindcoxa. Pleural integument light straw yellow to tan and usually partly brown.

Apn with anterior row of strong setae. Ppn with 1,2 strong posterior setae. Ppl, sp

and upper mep with setae. Pra setae 2-5. Stp setae restricted to lower part, below

level of lower edge of mep. Psp sometimes with weak, fine setae. Apn scales dense,

broad, colorless to smoky, translucent, usually with metallic silver or pale gold reflec-

tions. Ppn scales dense, broad, flat, colorless to dark; with metallic gold reflections,

some upper moderately blue and purple iridescent. Acx, ssp, psp, all but lower ante-

rior margin of stp and entire mep with dense, broad, flat, transparent, metallic silver

scales; ppl with a few similar scales; pcx, meron, metapleuron and metameron with-

out scales.

Legs: Long; femora I-III progressively shorter; tibiae I-III progressively shorter;

tarsus I-III longer than tibia III. Leg III with or without conspicuous outstanding

scales on tibia and tarsus. Coxae with dense, broad, flat, metallic silver scales. Femo-

Zavortink: Reclassification of Trichoprosopon 19

ra dark scaled with short to long streak of pale scales on lower or posterior surface.

Tibiae entirely dark scaled. Tarsi dark scaled with light scales on 2 or 3-4 or 5-II and

3 or 4-5-III. Claws simple; small on I, II; very small, often appearing single on III.

Wing: Dark scaled. Scales broad; wide spreading dorsally on Rs, R2,3, R,, R3

and M. Vein 1A ending far distad of branching of Cu. Alula with a few narrow mar-

ginal scales distad. Upper calypter without setae or with short row of setae basad.

Haltere: Dark scaled.

Abdomen: Tergite I completely scaled. Laterotergite without scales basad, its

lower margin visible. Setae conspicuous on tergite I and distal segments. Tergites I-

VII dark scaled with broad lateral patch of lustrous creamy to pale gold scales from

base to apex, the patches broader apically on distal segments. Sternites II-VII com-

pletely scaled, scales entirely lustrous creamy to pale gold or dark in median line

from base to apex of all segments.

FEMALE GENITALIA. As illustrated; not studied in detail.

MALES. Essentially as in females except for sexual characters. Head: Clypeus

small. Palpus long, 0.70-0.85 length of proboscis; 5-segmented; slender; setae few;

dark scaled. Antenna 0.53-0.70 length of proboscis; torus moderately enlarged, with

a few very small setae and scales medially; flagellum sparsely to moderately plumose,

segments 12,13 only slightly longer than in female. Legs: Anterior claw enlarged on

I or L,I; with submedian swelling or distinct tooth on I. Abdomen: Tergites with

setae laterally.

MALE GENITALIA. Large, exserted. Segment VIII: Unmodified. Segment IX:

Tergite strongly developed, with pair of distinctly separated, prominent, divergent

submedian lobes. Setae of each tergite lobe 4-7 (3-8), moderately to very strongly

developed, long, curved laterad apically, arising in a single or partly double curved

row. Sternite strongly developed; basal and lateral parts strongly sclerotized, forming

V- or U-shaped sclerite, disc membranous; without setae. Sidepiece: Main articula-

tion apparently with apex of lateral sclerotization of sternite IX; apodeme of side-

piece moderately strong, nearly perpendicular to long axis of genitalia. Moderately

long to long, conical, slightly curved mesad apically. Mesal surface not membranous.

Basal tergal area with large, very dense patch of strongly developed, moderately long

to long setae. Apical tergolateral, lateral and sternal surfaces with numerous strongly

developed, long setae; apical tergomesal area bare. Lateral and sternal surfaces dense-

ly scaled. Basal Mesal Lobe: Well developed, distinctly separated from sidepiece.

Crescentic to nearly semicircular, or with conical body and long narrow dorsal strip.

With numerous strongly developed setae. Clasper: Simple. Moderately long to long,

curved mesad apically, uniformly slender beyond base or swollen apically. Spiniform

short. Phallosome: Simple. Aedeagus with lateral sclerotizations weakly connected;

apical tergal arms not joined; lateral sclerotizations not approximated sternally; apex

of submedian tergal arm narrow, simple; broadest near base; index 1.7-2.5; without

large preapical teeth or sclerotized apical beak. Sternal aedeagal plates absent. Gon-

opore apical. Proctiger: Strongly developed. Paraproct with moderately large apical

teeth. Cercal sclerite present, elongate, or absent. Cercal setae numerous, fine.

PUPAE. Cephalothorax: Seta 1-C strongly developed, long, double, sigmoidally

curved. Seta 5-C moderately to strongly developed, long, usually 4-6b (3-7); stronger

and longer than 7-C.

Trumpet: Moderately long, narrow; pinna short; appearing slightly urceolate in

dorsal aspect, nearly cylindrical beyond basal curve in lateral aspect.

Abdomen: Tergite VIII with posterior edge straight laterad, caudolateral angle not

produced. Dorsalsensillum present on tergites IV,V only. Seta 1-I strongly devel-

20 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

oped, moderately long to long, dendritically branched; seta 1-I] moderately to strong-

ly developed, moderately long, usually 2-5b or f (1-9); seta 1-III,[V weakly to moder-

ately developed, short, usually single or double (1-3b). Seta 2-II-VII usually near cau-

dal edge of tergite, laterad of seta 1-II, mesad of seta 1-III-VII. Seta 3-I-III strongly

developed, long, usually single (1,2b), near caudal edge of segment and caudad of lev-

el of seta 4 on II,III. Seta 5-II-VI strongly developed, long, single on II, single or dou-

ble on III, usually double (2-6b) on IV-VI; seta 5-VII weakly developed, short, single

or double. Seta 6-I,II strongly developed, long, usually 2,3b (1-3) on I, usually single

(1,2b) on II; 6-VII moderately to strongly developed, 3-18b, usually cephalad and lat-

erad of seta 9-VII. Seta 7-1 moderately strong, moderately long, 2-5b; seta 7-II usual-

ly ventral. Seta 9-VII 15-24b (13-26); seta 9-VIII 14-24b (12-28).

Paddle: Large, subequal in length to or longer than seta 9-VIII, oval, apical part

acute. Midrib distinct, straight or curved laterad. Inner and outer margins usually

with small submarginal spicules, apex with larger submarginal and marginal spicules.

Surface of inner part and mesal portion of outer part with numerous small spicules.

LARVAE. Head: Foramen magnum circular, bounded by distinct collar. Posteri-

or tentorial pit removed from collar. Maxillary sutures straight or nearly so, parallel,

not extending to posterior tentorial pits. Labial sclerite broad; anterior margin con-

cave, with short, sharp spicules, moderately produced laterally. Mental plate very

short; apical margin deeply emarginate laterad of central teeth and tripartite. Mandi-

ble short. Maxilla modified for grasping, elongate, with short nonarticulated spine

and long, distally toothed articulated spine; without strong articulation with head

capsule far ventrad of palpus or basomesal tuft of elongate spicules. Maxillary palpus

adnate to body of maxilla basally, apical 0.3-0.5 free, short, conical. Distinct sclero-

tized palpifer absent. Seta 1-C strongly developed, stout. Seta 5-C laterad of 6-C.

Setae 8-10-C weakly developed, 3-5b (3-7). Seta 14-C weakly developed, usually sin-

gle (1-3f), near anterior ventral margin of head capsule. Seta 15-C weakly developed,

2-11f, very near anterior margin of labial sclerite. Seta 6-MP short, peglike, arising at

very base of body of maxilla.

Thorax: Seta O0-P moderately developed, 11-27b, not closely associated with 2,3-P

or 4-P. Seta 1-P single, closely associated with 2,3-P and slightly laterad of them. Se-

ta 4-P strongly developed, long, usually 14-21b (11-22), displaced laterad, on com-

mon tubercle with 5-7-P. Seta 5-T weakly developed, short. Seta 6-P strongly devel-

oped, long, usually 3,4b (2-5). Seta 7-P strongly developed, long, usually 18-27b (15-

28). Seta 8-M strongly developed, long, usually 6-10b (3-12). Setae 7,13-T not aris-

ing from sclerotized bars connected to tubercle of 9-12-T.

Abdomen: Dorsal sensillum present on IV,V only. Seta 1-I-VII without alveolar

plate; 1-I,11,VI weakly developed; 1-III-V,VII moderately developed; 1-III mesad of

4-III; 1-IV laterad of 4IV. Seta 2-I-VII mesad of seta 1 of same segment. Seta 3-VII

moderately developed, moderately long, usually 2,3b (1-4), without alveolar plate.

Seta 5-I, VI, VII and sometimes V weakly developed, usually 3-7b (2-8); 5-II-IV or V

moderately developed, usually 3-8b (2-11). Seta 6-I-III strongly developed, long, us-

ually 6-14b (4-16); 6-IV-VI strongly developed, long, single or double on IV, single

on V,VI.

Segment VIII: Strongly produced dorsocaudally, forming long cylindrical portion

beneath siphon. Comb scales weakly pigmented, usually 33-57 (9-76), free, in large,

irregular, oblique patch; each scale small, narrow, evenly fringed or with apical spine.

Setae 1-5-VIII displaced dorsad. Seta 3-VIII weakly developed, short, 4-13b. Seta 5-

VIII moderately developed, moderately long, 3-9b.

Siphon: Long, slender, index 6.6-10.2, straight or curved dorsad distally. Con-

Zavortink: Reclassification of Trichoprosopon ps |

spicuously spiculose; spicules short, simple, becoming stronger on apical portion of

siphon. Pecten absent. Seta 1-S moderately developed, usually single (1,2b), arising

near middle of siphon. Accessory midventral setae (1a-S) 20-37, not distinctly paired,

in long row extending entire length of siphon; moderately developed; medial setae us-

ually 4-7b (3-11). Seta 2-S strong, laterally compressed, apex curved and pointed;

arising on tubercle beyond body of siphon. Accessory dorsolateral setae (2a-S) 4-7

pairs; weakly developed, 1-3b. Seta 6-S weakly developed, flexible, nonhooked.

Anal Segment: Saddle small, dorsal. Seta 2-X usually 10-16b (7-17). Seta 3-X us-

ually 3-9b (2-10). Seta 4-X usually 5-15b (3-17); without sclerotized bar connecting

alveoli of this pair of setae.

DISCUSSION. The genus Johnbelkinia is distinguished from the other groups for-

merly placed in Trichoprosopon as follows: in the adult by the combination of: (1)

a long, slender proboscis, which is 1.20-1.40 length of femur I, (2) brilliant silver or

azure blue reflections of scales on vertex and occiput of head, (3) moderately broad

and flat scutal scales, (4) restriction of stp setae to area below level of lower edge of

mep and (5) partly light scaled tarsi II and III; in the male genitalia by: (1) a large,

very dense patch of strongly developed setae in basal tergal area of sidepiece, (2) api-

cal gonopore of aedeagus, (3) absence of sternal aedeagal plates, (4) weakly connec-

ted lateral sclerotizations of aedeagus, apical tergal arms not joined and (5) combina-

tion of strongly developed, V- or U-shaped IX-S and bare apical tergomesal area of

sidepiece; in the pupa by: (1) presence of a dorsal sensillum on only segments IV,V,

(2) oval, acute paddle with a distinct midrib and spicules on surface, (3) combination

of a strongly developed, long, 2,3b (1-3) seta 6-I, a strongly developed, long, single

(1,2b) seta 6-II and a moderately to strongly developed, 3-18b seta 6-VII that is usu-

ally cephalad and laterad of seta 9-VII and (4) combination of a moderately to

strongly developed, long, 4-6b (3-7) seta 5-C, a strongly developed, long seta 5-II-VI

that is single on II, single or double on III, and usually double (2-6b) on IV-VI and a

strongly developed, long, usually single (1,2b) seta 3-II,IIJ; and in the larva by (1) us-

ually large patch of comb scales, (2) long row of multiple, accessory midventral setae

on siphon, (3) tripartite mental plate, (4) strongly developed, long, usually 6-14b (4-

16) seta 6-I-III, (5) straight, parallel maxillary sutures that do not extend to posterior

tentorial pits, (6) combination of a strongly developed, long, 6-10b (3-12) seta 8-M

and absence of a pecten on siphon, (7) combination of a circular foramen magnum

with a distinct collar and a grasping maxilla without a strong articulation with head

capsule far ventrad of palpus and (8) maxillary palpus adnate to grasping maxilla for

0.5-0.7 of its length.

The affinities of Johnbelkinia are unknown. The larva resembles Shannoniana and

Tripteroides (Rachisoura) in development of the collar and maxillary sutures, but it

resembles Runchomyia in structure of the maxilla and in chaetotaxy. The larva has

additional unique features not found in any of these other groups, like the usually

large patch of comb scales and the resemblance of seta 6-III to 6-I,1I. The male geni-

talia are very unusual and, as far as is known, are unlike those of any other sabethine

mosquitoes. The adults resemble some species of Trichoprosopon in ornamentation,

but resemble Runchomyia again in development of the proboscis and in position of

the stp setae.

I am recognizing three species, leucopus, longipes and ulopus, in this genus. These

Species are strongly differentiated in the male genitalia and adult stage, but only

weakly differentiated in the immature stages. Dyar (1928:96-98) recognized the

same three species, but with the names Goeldia leucopus, G. trichopus and G. longi-

pes, respectively, and recognized also their close relationship. Lane and Cerqueira

22 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

(1942:525, 526, 528) and Lane (1953:855-858, 860) recognized the same species,

but with the names Trichoprosopon leucopus, T. edwardsianum and T. longipes, re-

spectively. Even though the same three taxonomic species have been recognized in

this group for more than half a century, misidentifications are extremely common.

There has been a particularly great amount of confusion in identification of males;

the genitalia illustrated as Trichoprosopon leucopus by Lane (1953:fig. 849) are ac-

tually ulopus, and the genitalia illustrated as Jongipes by Lane (1953:fig. 847) are the

true longipes, not Lane’s longipes, which is ulopus.

Several of the characters used by previous authors to separate species in this group

of mosquitoes are not reliable. Howard, Dyar and Knab (1915:163) used differences

in length of the female palpus to separate Lesticocampa longipes and culicivora.

However, this character is of no value, as there are no significant differences in palpal

length among the species of Johnbelkinia. Dyar himself (1923a:84) had also come to

this same conclusion. Dyar (1928:95) used white scaling of tarsus II to separate

Goeldia trichopus and longipes. This character is so variable, even at one locality,

that it can not be used to distinguish the species of Johnbelkinia. However, it is of

limited value in characterizing the species, and in general tarsus II is darkest in leuco-

pus and lightest in longipes. The presence or absence of outstanding scales on tarsus

III has been used by several authors. Again, this character is too variable to be used

alone to separate the species; part of the variation is biological and part is artifactual,

being related to the age and degree of rubbing of the specimens. The outstanding

scales are least conspicuous, in fact absent, in leucopus and most conspicuous in long-

ipes. In ulopus, they vary from being absent to nearly as conspicuous as in longipes.

Lane and Cerqueira (1942:488) used the presence or absence of prescutellar setae to

separate TJrichoprosopon longipes and edwardsianus. This character is of very limited

value, as previously pointed out by Cerqueira (1961a:459-460). Both prescutellar se-

tae and dorsocentral setae are more often developed in males than in females, and

both are more often developed in longipes and leucopus than in ulopus.

The following setae of ulopus larvae and pupae have been unquestionably homolo-

gized by tracing the appropriate nerve from a larval seta to a developing pupal seta in

one or more of eight specimens showing such connections from collections BRA 49,

COL 152, CR 541, CR 556, PA 41 and PA 596.

Thorax: Larval seta 1-T = pupal seta 10-C

' Larval seta 2-T = pupal seta 11-C

Larval seta 3-T = pupal seta 12-C

Abdominal segment I: Setae 2, 3, 4,

Abdominal segment V: Setae 3, 4,

Abdominal segment VI: Setae 3, 4, 5,

Abdominal segment VII: Setae 3, 4, 6, 7, 8, 10, 11

In numbering setae on the prothorax of the larva, I consider the more lateral of

the setae between 1-3-P and 5-7-P to be 4-P and the more mesal one to be O-P, regard-

less of which is more posterior. Setae 5-7-P have been assigned numbers on the basis

of their sequence from the midline of the body only, so 5-P is most mesal, 7-P most

lateral, and 6-P between these. This interpretation of these prothoracic setae is in

agreement with Belkin, Heinemann and Page’s (1970:191-229) for Wyeomyia, but at

odds with Belkin’s (1962b:360) for Malaya solomonis. Since these setae have no

3

129

Abdominal segment II: Seige 1, 2.3. 4, 5, 6,.7, 9, 11

Abdominal segment III: Setae 1, 2,3 C7. 0. 4, 1,4

Abdominal segment IV: Setae ie 2, 3,4,.5, 7, 9 aso “S”

4,5 9

5

a0,

4,5,

4, 5,

4,5,

Me

16

Zavortink: Reclassification of 7richoprosopon 23

demonstrated pupal homologies, neither interpretation can be shown to be correct or

incorrect.

Because misidentification of the species has been so common in this group, it is

frequently difficult or impossible to decide which zoological species actually formed

the basis for observations reported in the literature. I have tried to infer which spe-

cies various authors had before them when I compiled the lists of citations, but I

have undoubtedly made many mistakes in doing this. In some cases there are no cri-

teria for deciding, and so these citations are listed with a question mark under more

than one species. Except for the localities listed by Cerqueira (1961b), the distribu-

tions attributed to the species in this paper are based solely upon specimens exam-

ined by me.

This genus is dedicated to John N. Belkin, one of the world’s leading culicidolo-

gists, who happens also to be my mentor and friend.

A total of 1978 specimens, including 302 males, 658 females, 627 larvae, 391 pu-

pae and 349 individual rearings (182 larval, 135 pupal, 32 incomplete), has been ex-

amined for this revision of Johnbelkinia.

BIONOMICS. The immatures of Johnbelkinia are normally found in leaf axils and

flower bracts of tropical plants, usually of the families Araceae, Marantaceae and

Strelitziaceae, but also of the Bromeliaceae, Musaceae and Palmae. The larvae have

been reported to be predaceous on those of Wyeomyia. Adults are sylvan and pre-

dominantly diurnal. Females bite humans and various other mammals.

MEDICAL IMPORTANCE. Relatively large numbers of female Johnbelkinia have

been attracted to human subjects in studies conducted in various parts of Middle and ~

South America, so the species are apparently strongly anthropophilic. Arboviruses

have been isolated from ulopus in Trinidad and Colombia. Since these mosquitoes

are known to harbor viruses, and are attracted to and bite humans, they are of po-

tential importance as vectors of disease. Species of this genus have also been found

carrying the eggs of the human bot fly, Dermatobia hominis (Linnaeus, Jr., 1781).

DISTRIBUTION. The genus Johnbelkinia is widespread in tropical America, from

southern Mexico to coastal Ecuador, eastern Bolivia and central coastal Brazil. It is

apparently absent from all islands of the West Indies except Trinidad and Tobago.

KEYS TO SPECIES

ADULTS

l. Upper calypter usually without setae, rarely with 1 seta; scutal scales usually

moderately iridescent blue, ee bac purple, yellow-green or green .

I. ulopus

Cigna eaiv pier with short row of setae basally: scutal scales usually dull

brown or weakly iridescent copper, bronze, gold, green, blue or purple . 2

2(1). Tarsus 4-III completely white scaled; leg III with conspicuous outstanding

scales on distal part of tibia and on tarsus 1-3 or 4; integument of mep and

meron light . . . . 2. longipes

Tarsus 4-IIT white scaled it in apical 0. 1 ig 4: leg i without outstanding scales;

integument of mep and meron strongly markenca es PS leucopus

24

2(1).

1906.

Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

MALE GENITALIA

Basal mesal lobe crescentic to nearly semicircular and with setae that are di-

rected caudad and curved or weakly sinuous at apex; sidepiece with den-

sest region of basal cacie: setae distad of level of apex of basal mesal lobe

I. ulopus

Basal. metal lobe with long narrow dorsal strip and onic body bearing setae

that are directed dorsad and very strongly curled at apex; sidepiece with

densest region of basal tergal setae proximad of level of apex of basal me-

Bal OG eye el ee ee = oe

Tergite IX lobe much longer than wide; clasper constricted basally and slight-

ly to conspicuously swollen apically; apical tergal arm of aedeagus with

patch of 4-11 small, strongly sclerotized, dorsal preapical teeth and serru-

late or striate, laterally curved, sclerotized apical process . . 2. longipes

Tergite [X lobe about as long as wide; clasper without basal constriction and

uniformly slender beyond base; apical tergal arm of aedeagus without dor-

sal preapical teeth or sclerotized apical process . .. . . .3. leucopus

PUPAE

(3. leucopus unknown)

Sternite II-VII uniformly very weakly pigmented, or median part of anterior

margin of II-IV only slightly more intensely pigmented than remainder of

ventral surface. . . . I. ulopus

Sternites II-[V (II-VIT) atk eee part of eee? bees ede pig-

mented, conspicuously darker than the very weakly pigmented remainder

OF Ventral SURACes ey os ee ee le es Ge oe, loneipes

LARVAE

(3. leucopus unknown)

Seta 4-X usually 5-9b (3-12), rarely with alveolar plate; seta 6-I usually 6-10b

(4-14); most comb scales usually with even fringe or with 1 or 2 apical ele-

ments of fringe enlarged into smallspines . . . . . tL. ulopus

Seta 4-X usually 10-15b (6-17), with alveolar plate; seta a I usually 11-14b

(7-14); most comb scales usually with moderately large apical spine

. 2. longipes

1. Johnbelkinia ulopus (Dyar & Knab)

Figs. 1, 3-7

Lesticocampa ulopus Dyar and Knab 1906a:137-138. *TYPE: Holotype female, Blue-

fields [Zelaya] , Nicaragua, W. F. Thornton [USNM, 10004]. Synonymized with longipes

by Howard, Dyar and Knab (1915:172-174). NEW COMBINATION AND NEW STATUS.

Zavortink: Reclassification of Trichoprosopon 25

1907. Lesticocampa culicivora Dyar and Knab 1907:207. *TYPE: Holotype female (11.30), Ta-

bernilla, Canal Zone, Panama, reared from larva from flower bracts of Heliconia, 28 Apr

1907, A. Busck [USNM, 10849]. Synonymized with longipes by Dyar (1923a:84-85).

TRANSFERRED SYNONYMY.

Trichoprosopon (Runchomyia) ulopus of Belkin, Schick and Heinemann (1965:42); Valencia

(1973:14).

Lesticocampa ulopus of Busck (1908:75-76); Theobald (1910:622); Stone and Knight (1957:119).

Trichoprosopon (Runchomyia) culicivora of Belkin, Schick and Heinemann (1965:45, 58).

Goeldia culicivora of Bonne-Wepster and Bonne (1921:17; 1922:38).

Lesticocampa culicivora of Busck (1908:76); Theobald (1910:558); Martini (1914:65, 66); How-

-ard, Dyar and Knab (1915:168-170); Stone and Knight (1957:119).

Trichoprosopon (Runchomyia) longipes of Stone, Knight and Starcke (1959:76-77, in part); Faur-

an (1961:14-15, in part); Stone (1963:121); Forattini (1965:169, 170, 172, 173, 174-175, 176,

217, 291, in part); Cova Garcia, Sutil and Rausseo (1966a:66, 68, in part; 1966b:67, 131, 384-

385, in part); Barreto-Reyes and Lee (1969:415); Belkin, Heinemann and Page (1970:14, 16,

18, 19, 191); Kruijf (1972:50); Diaz Najera and Vargas (1973:124); Knight and Stone (1977:

314, in part).

Trichoprosopon (Rhunchomyia) longipes of Lane (1953:855-857, in part); Barreto-Reyes (1955:

80); Trapido, Galindo and Carpenter (1955:531, 535, 537, 538, 539); Trapido and Galindo

(1955:670); Floch (1956:35, 221, in part); Trapido and Galindo (1957:122, 125, 131); Cerque-

ira (1961a:460; 1961b:154); Diaz Najera (1961:204); Morales and Vidales (1962:5, 10, 11);

Diaz Najera (1966:63); Xavier and Mattos (1970:452; 1975:256; 1976:403).

Trichoprosopon (Rhynchomyia) longipes of Diaz Najera (1963:192).

Trichoprosopon (Hyloconops) longipes of Lane and Cerqueira (1942:525, in part); Anduze (1943:

191); Lane (1945:133); Anduze, Pifano and Vogelsang (1947:12); Anduze (1947:351); Arnett

(1949:244); Galindo, Carpenter and Trapido (1951:129).

Trichoprosopon longipes of Galindo, Trapido and Carpenter (1950:549, 554); Horsfall (1955:317);

Galindo (1958:436); Aitken (1960:4, 8); Maia (1963:91); Spence, Anderson et al. (1964:114);

Porter (1967:39); Barr and Barr (1969:196); Forattini, Rabello and Cotrim (1970:79).

Goeldia (Goeldia) longipes of Dyar (1923a:84-85; 1923b:174); Edwards (1932:72, in part); Lane

(1939:162-163, in part); Floch and Abonnenc (1947:2-3, in part).

Goeldia longipes of Bonne-Wepster and Bonne (1921:17; 1922:38); Evans (1923:110); Bonne-Wep-

ster and Bonne (1923:124, in part); Dyar and Shannon (1924:483); Bonne and Bonne-Wepster

(1925:144-146, in part); Dyar (1925:128-129; 1928:96-97, in part); Dunn (1929:500); Lima

(1931:66-67); Edwards (1934:632); Komp (1936:60, in part); Lane (1936:7); Antunes (1937a:

72-73, in part; 1937b:72); Patino Camargo (1937:242, in part); Boshell-Manrique (1938:415, in

part); Kumm, Komp and Ruiz (1940:397); Anduze (1941:9); Floch and Abonnenc (1942b:2-3,

in part); Floch (1955:315, in part); Horsfall (1955:319).

Lesticocampa longipes of Howard, Dyar and Knab (1915:172-174).

Goeldia trichopus of Bonne (1923:128-129); Bonne and Bonne-Wepster (1923:124, in part; 1925:

141-143, in part); Shannon (1931:7).

Trichoprosopon (Runchomyia) leucopus of Stone, Knight and Starcke (1959:76, in part); Cova

Garcia, Sutil and Rausseo (1966a:67, 68, in part); Barreto-Reyes and Lee (1969:415); Diaz Na-

jera and Vargas (1973:124); Knight and Stone (1977:314, in part).

Trichoprosopon (Rhunchomyia) leucopus of Lane (1953:857-858, in part); Trapido and Galindo

(1955:670); Trapido, Galindo and Carpenter (1955:531, 535, 537, 538, 539); Diaz Najera

(1961:204; 1966:63).

Trichoprosopon (Hyloconops) leucopus of Vargas and Martinez-Palacios (1953:300); Vargas (1956:

33); Diaz Najera (1963:192).

Trichoprosopon leucopus of Forattini (1965:172).

Goeldia (Goeldia) leucopus of Dyar (1923a:86, in part).

Goeldia leucopus of Bonne and Bonne-Wepster (1925:152, in part); Dyar (1925:129, in part; 1928:

97-98, in part).

Lesticocampa leucopus of Howard, Dyar and Knab (1915:170-172, in part).

26 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Trichoprosopon (Runchomyia) hyperleucum of Stone, Knight and Starcke (1959:76, in part);

Knight and Stone (1977:313, in part).

Trichoprosopon (Rhunchomyia) hyperleucus of Lane (1953:858).

Trichoprosopon (Hyloconops) hyperleucus of Lane and Cerqueira (1942:526-527).

Trichoprosopon hyperleucum of Forattini, Rabello and Cotrim (1970:79).

Lesticocampa rapax of Dyar and Knab (1906:137, in part).

?Trichoprosopon (Runchomyia) longipes of Martinez and Prosen (1953:31); Morales-Ayala (1971:

13},

?Goeldia longipes of Bonne (1923:129).

?Trichoprosopon (R.) edwardsianus of Suarez (1963:239).

?Trichoprosopon (Hyloconops) trichopus of Anduze, Pifano and Vogelsang (1947:12).

?Goeldia (Goeldia) trichopus of Chagas, da Cunha et al. (1938:198, 202); Floch and Abonnenc

(1947:3).

?Goeldia trichopus of Kumm and Novis (1938:503, 512); Floch and Abonnenc (1942a:3; 1942b:3).

?Trichoprosopon (Runchomyia) leucopus of Fauran (1961:14); Stone (1961:33); Morales-Ayala

(1971:139); Kruijf (1972:50).

?Trichoprosopon (Rhunchomyia) leucopus of Trapido and Galindo (1957:122, 124, 125, 128,

131, 134, 135, 138); Xavier and Mattos (1975:256).

?Trichoprosopon (Hyloconops) leucopus of Anduze (1947:351); Anduze, Pifano and Vogelsang

(1947:12).

?Trichoprosopon leucopus of Galindo and Trapido (1957:146).

?Goeldia (Goeldia) leucopus of Floch and Abonnenc (1947:3).

?Goeldia leucopus of Anduze (1941:9); Floch and Abonnenc (1942a:3; 1942b:3).

FEMALE (fig. 3). Wing: 4.20mm. Proboscis: 4.02mm. Femur I: 3.27mm. Abdo-

men: about 3.9mm. Thorax: Scutal scales usually moderately iridescent blue or

blue-green, sometimes purple, yellow-green or green. Postnotum usually without

scales. Pleural integument strongly darkened on ssp, psp, stp, mep and meron, or

only weakly to moderately darkened on ssp and psp, or ssp, psp and stp. Legs: Leg

III with or without outstanding scales; if present, varying from inconspicuous and re-

stricted to tarsus 2-III to conspicuous and extending from apical 0.5 of tarsus 1-III

to apex of 3-III. Tarsus II with streak of white scales on anterior and dorsal surfaces

starting between base of 2-II and apical 0.6 of 3-II and extending to apex of 4-II or

5-II; scales on ventral and posterior surfaces of same segments dark, tanish, partly

white or entirely white. Tarsus III white scaled from apical 0.0-0.5 of 3-III or apical

0.2-1.0 of 4-III to apex of 5-III. Wing: Upper calypter usually without setae, rarely

with | seta. Abdomen: Sternites II-VII entirely light scaled or with some scales

moderately to strongly darkened.

MALE (fig. 3). Essentially as in female except for sexual characters. Head: Fla-

gellum sparsely plumose, middle segments with about 14-18 bristles. Legs: Anterior

claw I enlarged, with submedian crestlike swelling; anterior claw II enlarged, simple.

MALE GENITALIA (figs. 4, 7). Segment IX: Tergite lobe about as long as wide.

Setae of lobe 5-7 (4-8), moderately to strongly developed, flattened apically, with

long, filamentous apical prolongation. Sternite U-shaped. Sidepiece: Slender. Den-

sest region of basal tergal setae distad of level of apex of basal mesal lobe. Basal ster-

nomesal setae 2-5, short to long, curved at apex; strongly developed medial sterno-

mesal setae 1-7. Basal Mesal Lobe: Crescentic to nearly semicircular; setae directed

caudad, curved or weakly sinuous at apex. Clasper: Moderately long to long; with-

out basal constriction; uniformly slender beyond base; with several fine setae apical-

ly. Spiniform straight, stout or slender. Phallosome: Aedeagus weakly pigmented;

distal lateral margin partly concave in dorsal aspect; distal end of submedian tergal

arm nearly parallel with mate; apical tergal arm without dorsal preapical teeth or

Zavortink: Reclassification of Trichoprosopon Pe

sclerotized apical process. Proctiger: Paraproct with 4 or 5 (3-6) moderately large,

dorsally curved apical teeth and with serrulate or crenulate, moderately sclerotized

apical lamella mesad of apical teeth that does not project beyond teeth. Cercal scler-

ite present, elongate. :

PUPA (fig. 4). Abdomen: 4.7mm. Trumpet: 0.55mm. Paddle: 0.68mm. Cepha-

lothorax: Very weakly pigmented. Trumpet: Very weakly to weakly pigmented,

sometimes becoming slightly darker distad. Abdomen: Very weakly pigmented;

sternites II-VII uniformly pigmented or median part of anterior margin of II-IV only

_ slightly more intensely pigmented than remainder of ventral surface. Terminal Seg-

ments: Male genital lobe moderately large, its width usually less than 0.50 (0.41-

0.54) of segment VIII. Paddle: Very weakly pigmented. Index usually 1.22-1.42

(1.11-1.84).

LARVA (fig. 5). Head: 1.30mm. Siphon: 1.21mm. Anal Saddle: 0.28mm.

Head: Very weakly pigmented, collar darker. Mental plate with 10-12 (10-13) teeth

on each side of median tooth. Antenna: Often with a few to numerous small spic-

ules medially. Abdomen: Seta 6-I usually 6-10b (4-14); seta 6-II usually 8-11b (5-

15); seta 6-III usually 7-10b (5-12). Segment VIII: Comb scales usually 37-57 (21-

76); most comb scales of patch usually with even fringe or with 1 or 2 apical ele-

ments of fringe enlarged into small spines. Siphon: Very weakly pigmented, becom-

ing slightly darker distad. Straight or weakly to moderately curved dorsad distally.

Seta 1-S inserted 0.42-0.61 distance from base of siphon. Accessory midventral setae

(la-S) usually 23-28 (20-30). Anal Segment: Saddle very weakly pigmented, indis-

tinct. Seta 3-X usually 3-6b (2-8). Seta 4-X moderately to strongly developed, usual-

ly 5-9b (3-12), rarely with alveolar plate.

SYSTEMATICS. Johnbelkinia ulopus may be distinguished by the following char-

acters: in the adult from longipes and leucopus by: (1) usual absence of setae on up-

per calypter of wing and (2) moderately iridescent blue, blue-green, green, purple or

yellow-green scales on scutum; in the male genitalia from Jongipes and leucopus by:

(1) crescentic to nearly semicircular shape of basal mesal lobe, (2) caudally directed

setae of basal mesal lobe that are curved or only weakly sinuous at apex and (3) loca-

tion of densest region of basal tergal setae distad of level of apex of basal mesal lobe;

in the pupa from Jongipes by: (1) uniformly or nearly uniformly very weakly pig-

mented sternites and (2) smaller male genital lobe that is usually less than 0.50 width

of segment VIII; and in the larva from longipes by: (1) usually fewer branches, 5-9

(3-12), in seta 4-X, (2) usual absence of an alveolar plate for seta 4-X, (3) usually

fewer branches, 6-10 (4-14), in seta 6-I and (4) shape of comb scales, which are usual-

ly evenly fringed or have one or two apical elements of fringe enlarged into small

spines. Johnbelkinia ulopus may be further distinguished from longipes, with which

it is sympatric in parts of South America, by the following additional adult charac-

ters: (1) less plumose flagellum in male, (2) submedian crestlike swelling on anterior

claw I of male, (3) enlarged anterior claw II of male, (4) usually lighter abdominal

sternites in female and (5) usual absence of outstanding scales on tibia III and basal

0.5 of tarsus 1-III.

There is a considerable amount of variation in this species, with much of it being

geographical. In the adults, the integument of mep and the meron is strongly dark-

ened in specimens from Nicaragua, Costa Rica, Panama, Colombia and the Pacific ver-

sant of Ecuador; tarsus II is darkest in specimens from Nicaragua, Costa Rica, western

Panama and the Atlantic versant of Colombia; leg III is without outstanding scales or

with only inconspicuous to moderately conspicuous outstanding scales in specimens

from Nicaragua, Costa Rica and western Panama; and tarsus III is darkest, with seg-

28 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

ment 4 dark in the basal 0.3-0.6, in specimens from Nicaragua, Costa Rica, western

Panama and Colombia. Specimens from north of the area indicated in Central Amer-

ica and south and east of the areas indicated in South America have a lighter integu-

ment on mep and the meron, more extensively light scaled tarsi II and III, and usual-

ly more conspicuous outstanding scales on leg III. All four of the variations mention-

ed reach their most extreme expression in Nicaragua, Costa Rica and western Panama,

precisely the area in which ulopus is sympatric with leucopus, a species with strongly

darkened integument on mep and the meron, with the darkest tarsi II and HI known

in the genus, and without outstanding scales on leg III. This appears to be another

example of species being more similar in their area of sympatry than elsewhere. I

have reported similar observations in the genera Orthopodomyia (Zavortink 1968:12-

13) and Aedes (Zavortink 1972:22, 51). Now, as in the past, I consider the most rea-

sonable explanation for this phenomenon to be introgressive hybridization.

In the male genitalia, the basal sternomesal setae of the sidepiece are 3 or 4, mod-

erately strong, short and do not arise on a swelling in specimens from Nicaragua, Cos-

ta Rica, western Panama, Trinidad and French Guiana; these setae are more strongly

developed and longer in specimens from central and eastern Panama and Venezuela;

and are 4-7, strongly developed, long and arise on a swollen lobe in specimens from

the Pacific versant of Colombia and Ecuador. The medial sternomesal setae of the

sidepiece are usually 4-7 in all areas mentioned above except the Pacific versant of

Colombia and Ecuador, where they are absent or reduced to 1. The clasper is rela-

tively long in all known populations except the one in Pacific coastal Ecuador, where

it is shortened. Setae of the basal mesal lobe, which are usually relatively short and

only curved at the apex, are longer and weakly sinuous at the apex in specimens from

the Pacific part of Colombia and Ecuador. The two extremes in variation in the male

genitalia are illustrated, the topotypic Nicaraguan population in Fig. 4, and the Paci-

fic Ecuadorian population in Fig. 7. I consider the Nicaraguan form to be the more

primitive, because of its disjunct distribution in Central America and northeastern

South America, and the Ecuadorian to be the most derived. This interpretation is

subject to change, though, when the genitalia become known for the northern Cen-

tral American populations and the more southern South American populations. It is

perhaps significant that the Nicaraguan form, with short basal sternomesal setae, 4-7

medial sternomesal setae, and short and slightly curved basal mesal lobe setae, is

known from areas where ulopus occurs with or near leucopus or longipes, both of

which have longer basal sternomesal setae, 0-4 medial sternomesal setae and longer,

sinuous basal mesal lobe setae. Such morphological divergence in areas of sympatry

suggests character displacement. I believe it fully possible that both character dis-

placement and introgressive hybridization could occur, in different characters of

course, between the same species.

In the larva, many specimens from Trinidad, Tobago and the Guianas have an en-

larged spine at the apex of the comb scales, an alveolar plate at the base of seta 4-X

and more than the usual number of branches in setae 6-I and 4-X. In these features,

these larvae resemble longipes, which occurs throughout much of this same region.

Again, introgressive hybridization is a possible explanation for this pattern of varia-

tion.

The third instar larva of ulopus, illustrated in Fig. 6, differs from the fourth instar

in several notable features. The maxilla is relatively longer, the prothorax has a large

sclerotized plate anterolaterally, setae 5,6-P are very strongly developed and the ter-

minal abdominal segments are spiculose. The second instar larva, parts of which are

shown in Fig. 7, has some even more remarkable features. The maxilla is extremely

Zavortink: Reclassification of Trichoprosopon 29

long and lacks apical spines and the spicules of the siphon are enlarged, resemble pec-

ten teeth and are restricted to a broad stripe of several rows on each side. This instar

resembles the third in development of the prothoracic plate, strength of 5,6-P and

presence of spicules on the body. As in the case of the second instar larva of Runch-

omyia magna, setae 8-M and 7-T are absent.

Several artifacts are shown in the illustration of the male genitalia of ulopus from

coastal Ecuador (Fig. 7). In the undissected genitalia the appearance of the aedeagus

and proctiger is unusual because the phallosome in partly everted. In the case of the

- dissected parts, the lateral aspect of the proctiger is unusual because this piece is part-

ly rotated.

BIONOMICS. The immatures of ulopus are represented in more than 125 collec-

tions made for the project ““Mosquitoes of Middle America.’ Eighty percent of these

collections are from leaf axils, 17% from flower bracts of Heliconia and 3% total

from treeholes, fallen cacao fruits and fallen palm spathes. Of the more than 100

collections from leaf axils, 62% are from axils of plants in the family Araceae (vari-

ously listed on the record forms as Colocasia, Dieffenbachia, Montrichardia, Xantho- —

soma, alispata, elephant ear, taro or Araceae), 25% are from Strelitziaceae (listed as

Heliconia or platanillo), 8% are from Marantaceae (listed as Calathea or Marantaceae),

3% are from Bromeliaceae (listed as bromeliad) and 2% are from Musaceae (listed as

wild banana). Galindo, Carpenter and Trapido (1951:129) reported the larvae (as

longipes) from many of these same habitats and also from the leaf axils of Caladium

(Araceae) and the flower bracts of Calathea. Bonne’s report (1923:129) of longipes

larvae from the leaf bases of Heliconia and Ravenala (probably Phenakospermum gui-

anense, Strelitziaceae) may refer to ulopus in part.

Larvae of ulopus have been reported as being predaceous, particularly on the lar-

vae of Wyeomyia, by Busck (1908:75-76, as ulopus and culicivora), Howard, Dyar

and Knab (1915:168-170, as culicivora) and Galindo, Carpenter and Trapido (1951:

129, as longipes). Additional reports of predation on Wyeomyia by larvae of longi-

pes by Bonne (1923:129) and Bonne-Wepster and Bonne (1925:144-146) may refer

to ulopus in part.

Most adults collected for the project “Mosquitoes of Middle America’’ were ob-

tained resting, particularly in forested areas, between the hours of 0700 and 1300 or

in biting-landing collections with human bait between the hours of 0800 and 1800.

A few have been obtained in Shannon, light or ‘“mouse-baited”’ traps.

Edwards (1934:632) reported the capture of an adult [male] of ulopus (as longi-

pes) from the flowers of Matayba inelegans in Guyana and Bonne (1923:128-129) re-

ported the capture of a male (as trichopus) in a house in Surinam. Barreto-Reyes and

Lee (1969:415) noted this species (as Jongipes and leucopus) carrying the eggs of

Dermatobia hominis in Colombia. Antunes’s report (1937a:72-73) that one of 150

adults of Jongipes caught at Restrepo, Colombia, carried the eggs of D. hominis and

that the species had a “special predilection” for biting the noses of horses refers in

part to ulopus. The reports of Floch and Abonnenc (1947:2-3) that longipes adults

are sometimes found in agglomerations in the forest in French Guiana and of Kumm

and Novis (1938:503, 512) that Jongipes females have been taken in traps baited with

agoutis and “‘other wild animals” in Brazil may refer in part to ulopus.

Large numbers of ulopus have been reported (as Jongipes and leucopus) as being

attracted to human subjects during the day and predominantly at ground level in

studies conducted in western Panama and Costa Rica by Trapido, Galindo and Car-

penter (1955:531, 535, 537, 538, 539), in western Panama by Trapido and Galindo

(1957:122, 124, 125, 128, 131, 134, 135, 138), in Honduras by Trapido and Galin-

30 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

do (1955:670), in Nicaragua by Galindo and Trapido (1957:146) and in Colombia

by Barreto-Reyes and Lee (1969:415). Although the single specimen previously de-

termined as Jeucopus from the Gorgas Memorial Laboratory collection that I exam-

ined was actually ulopus, it is possible that Jewcopus is represented in the material

collected in some of the studies by Galindo and associates.

Aitken (1960:4, 8) reported the isolation of the arbovirus TR8349 from ulopus

(as longipes) in Trinidad and Spence, Anderson et al. (1964:114) reported the isola-

tion of Triniti virus from a mixed pool of trichoprosopons that included two ulopus

(as longipes) that had been captured at human bait in the Arena Forest of Trinidad,

31 May 1955, between 1745 and 1915 hours. Barreto-Reyes and Lee (1969:415) al-

so reported the isolation of arbovirus from this species (as Jongipes and leucopus) in

Colombia.

DISTRIBUTION. Johnbelkinia ulopus extends from Veracruz and Oaxaca in

southern Mexico southward into the northern half of South America. In South

America, I have seen specimens from near the Pacific coast in Colombia and Ecuador,

near the Caribbean coast in Colombia and Venezuela, and along the Atlantic coast in

Trinidad, the Guianas and the Brazilian states of Para and Bahia. Inland, I have seen

specimens from only Boyaca and Meta, Colombia, but Cerqueira (1961b:154) re-

cords this species (as Jongipes) from the Brazilian states of Amazonas and Mato Gros-

so. Lane (1953:857) recorded ulopus (as longipes) from Cuba, and this record has

been repeated by others (Stone, Knight and Starcke 1959:76; Porter 1967:39). I

have not seen the specimen upon which this record is based, and, like Belkin, Heine-

mann and Page (1970:191), I believe this record is doubtful. Material examined:

1681 specimens; 276 males, 477 females, 578 larvae, 350 pupae; 311 individual rear-

ings (153 larval, 129 pupal, 29 incomplete).

BRAZIL. Bahia: Ilheus, July 1930, N. C. Davis, R. C. Shannon, 1F [USNM]. Para: Belem,

IPEAN, Mocambo Forest, 1970, T.H.G. Aitken (BRA 49), 1L [UCLA]. Curralinho, Nov 1935, 1F;

Jan 1936, 1F; date unspecified, 1F, 1L [USNM] ; date unspecified, H.W. Kumm, 5F [BM]. Curra-

linho, Campinho, H. W. Kumm, 1F [BM]. Curralinho, Furo Pruana, 1F [USNM]. Curralinho, Rio

Canaticu, 1F [USNM]. Curralinho, Rio Pariacasinho, 2F [USNM]. Vigia, Vigia, June 1935, 1F

[BM].

COLOMBIA. Boyaca: Chiquinquira, Pauna, Feb 1937 (COR 1346), 1F; ?Feb 1937 (COR

1347, 1350), 2F [UCLA]. ?Muzo, 1936 or 1937 (COR 1249, 1252, 1254, 1255, 1256, 1257,

1259, 1260), 19F [UCLA]. Meta: ?Chipadera, Nov 1936 (COR 107), 1F [UCLA]. Restrepo, Dec

1941, P.C.A. Antunes, 1F [FH]. Villavicencio, Bosque Ocoa, 14 June 1944 (COK 44), 1F [UCLA].

Locality unspecified, 20 Nov 1936 (COR 119), 2F; 20 Oct 1936 (COR 128), 1F; 21 Oct 1936

(COR 130), 8F; 23 Oct 1936 (COR 133), 1F; 27 Nov 1936 (COR 161), 1F;4 Dec 1936 (COR 164,

165), 2F [UCLA]. Narino: Gorgona I., July 1924, L. E. Cheesman, 1F [BM]. Norte de Santan-

der: Villamizar, Zulia River, L.H. Dunn, 1F [USNM]. Valle del Cauca: Buenaventura (20km S),

Rockefeller Foundation Virus Field Station (W), 24 Mar 1966, V. H. Lee (COL 152), 1 IpM (152-

10), 3 pM (152-100-102), 5L [UCLA]. Darien, New Calima Dam (200m W), 31 Aug 1965, J. N.

Belkin, V. H. Lee (COL 120), 1 IpF (120-10), 1 pF (120-100), 1P [UCLA]. Locality unspecified:

(COR 487, 491, 521, 523), 6F [UCLA].

COSTA RICA. Alajuela: Higuito, near San Mateo, P. Schild, 3M, SF [USNM]. Orotina, 1F

[USNM]. Cartago: La Suiza, P. Schild, 1M [USNM]. Turrialba, 5 Aug 1963, C. L. Hogue, 1F

[UCLA] ; 5 June 1951, O. L. Cartwright, 5F [USNM] ; June 1954, P. A. Buxton, 1F [BM]. Limon:

Limon (8km S), Beverly (W), 15 Dec 1971, D. and K. Schroeder (CR 556), 1 IpM (556-10), 6 IpF

(556-11, 14-18), 1 Ip (556-12), 3 pM (556-101, 103, 104), 2P, 3L [UCLA]. Limon (8km S), Bom-

ba (1.8km E), 3 Oct 1971, D. and K. Schroeder (CR 466), 3 IpM (466-11, 14, 15), 3 IpF (466-10,

12, 13), 2 pM (466-100, 101), 2L [UCLA]. Liverpool, H. W. Kumm, 1F [BM]. Pandora, 29, 30

Apr 1964, F. S. Truxal, 1F [LACM]. Siquirres, 1F [USNM]. Zent (8km E), 17 Dec 1971, D.

Schroeder (CR 565), 1 lpM (565-10), 1 IpF (565-11), 4 pM (565-100-102, 104) [UCLA]. Zent,

Zavortink: Reclassification of Trichoprosopon 31

about half way to Boston, 12 Dec 1971, D. and K. Schroeder (CR 541), 1 IpM (541-15), 3 IpF

(541-11, 14, 16), 2 lp (541-12, 13), 1 pM (541-102), 3 pF (541-100, 103, 104), 2L [UCLA].

Puntarenas: Golfito (near), 9 July-1 Aug 1972, A. Menke, F. Truxal (CR 172), 1M [UCLA]. Osa

Peninsula, Rincon, 1 July 1963, C. L. Hogue (CR 140), 1 lpF (140-301) [UCLA].

ECUADOR. Bolivar: Balzapamba, F. Campos, 1M [USNM]. Canar: Cochancay, km 86 Hwy

8, 13 Feb 1966, J. N. Belkin, E. J. Gerberg (ECU 164), 1 lpF (164-20), 1M; same data (ECU 165),

1 lpF (165-11), 1L [UCLA]. Los Rios: Pichilingue, 15-19 Oct 1944, E. J. Hambleton, 1F [USNM].

Valencia, 1M [USNM]. Locality unspecified: Aug 1938, Hanson, 1F; 26 July 1938, 1M [UCLA];

4M [USNM].

FRENCH GUIANA. Guyane: Cayenne, foret de Cabassou, 28 Nov 1967, J. Clastrier (FGC

3287), 1F; 18 July 1968, J. Clastrier (FGC 429-1), 1M; 12 July 1968, J. Clastrier (FGC 3911), 1F

[UCLA]. Cayenne, foret de la Chaumiere, 13 Mar 1968, J. Clastrier (FGC 3318), 1 pF (3318-10)

[UCLA]. Remire, near Lac du Rorota, 8 Mar 1967, R. X. Schick (FG 110), 1 lpF (110-10), 1 pM

(110-100) [UCLA]. Inini: Saul, upper Fleuve Mana, near Placer Souvenir, 29 June 1952 (FGA

189), 4F [UCLA]. Saut Bief, Apr 1940, 3F [UCLA]. Locality unspecified: J. Clastrier, 1F

[UCLA].

GUYANA. Demerara: Georgetown, 1.5km N of Central Horticultural Station on road from

Atkinson Field, 10 Nov 1967, R. X. Schick, P. Rauch, R. Hansell (GUY 53), 2 IpF (53-10, 24)

[UCLA]. Essequibo: Essequibo River, Moraballi Creek, 23 Aug 1929, 1M [BM]. Issororo, G. E.

Bodkin, G. M. Vevers, 1F [BM]. Locality unspecified: Nov 1909, L. D. Cleare, 1F [BM].

HONDURAS. Atlantida: Tela, Lancetilla, Tela Railroad Co. grounds, 13 Aug 1964, A. Quino-

nez (HON 26), 1 lpF (26-11), 1L; 17 Aug 1964, A. Quinonez, V. P. Cowsill (HON 44), 2 pF (44-

100, 101), 7L [UCLA].

que, 5 Sept 1959, J. Rene Zurita, 3F [ISET] ; 19 July 1970, D. and K. Schroeder (MEX 580), 2F

[UCLA]. Oaxaca: Valle Nacional (above), 12 May 1963, 1F [USNM]. Tabasco: Teapa, ?Samun,

8 Dec 1954, M. Figueroa, 1F [ISET]. Veracruz: Cardenas, Texistepec, 24 Mar 1960, R. Zurita, 1F

[ISET]. Moloacan, 26 Nov, 10 Dec 1959, J. Lopez B., 3F; 5 Jan 1960, J. R. Zurita, 1F [ISET].

St. Lucrecia, 21 June 1905, F. Knab, 1F [USNM].

NICARAGUA. Zelaya: Bluefields, W. F. Thornton, 8F (includes ulopus holotype) [USNM].

Bluefields, Bluefields Airport, 15 July 1964, A. Quinonez (NI 49), 1 IpF (49-20), 1 lp (49-21), 2L

[UCLA]. Bluefields, Cemetery (1.5km W), 23 Nov 1971, D. and K. Schroeder (NIC 82), 2F

[UCLA]. Bluefields (4km S), Punta Masaya, 27 Nov 1971, D. and K. Schroeder (NIC 113), 1F;

same data (NIC 116), 1 lpM (116-21), 1 IpF (116-20) [UCLA]. Bluefields, Rio Escondido, 17 July

1964, A. Quinonez (NI 52), 2 pM (52-103, 107); same data (NI 54), 1P, 3L; same data except 18

July 1964 (NI 56), 3 lpF (56-15, 18, 19), 1 pM (56-103), 1 pF (56-112), 1 lp (56-16), 1P, 4L

[UCLA].

PANAMA AND CANAL ZONE. Bocas del Toro: Almirante, 30 Apr 1963, A. Quinonez (PA

289), 2 IpF (289-107, 108), 2 pM (289-101, 103), 2 pF (289-104, 106), 1 lp (289-102), 2P, 4L

[UCLA]. Almirante (15km SW), 2 May 1963, A. Quinonez (PA 303), 1 pM (303-102), 1 pF (303-

101), 1P, 2L [UCLA]. Almirante, Crematory, 27 Apr 1963, A. Quinonez (PA 265), 1 pF (265-

114) [UCLA]. Almirante, farm near, 14, 15 Apr 1964, A. Quinonez (PA 665), 1L [UCLA]. Al-

mirante, 14% mile swamp, 4 May 1963, A. Quinonez (PA 310), 1F [UCLA]. Almirante, Milla 2, 9

May 1963, A. Quinonez (PA 342), 1F [UCLA]. Almirante, Oil Tank, 27 Apr 1963, A. Quinonez

(PA 261), 1 lp (261-103), 2P, 3L [UCLA]. Almirante (1-4km W), Quebrada Nigua, 27 Apr 1963,

A. Quinonez (PA 256), 1 IpF (256-105), 3 pM (256-103, 104, 110), 1 pF (256-109), 1 Ip (256-

112), 1L [UCLA]. Almirante, Repressa, 15 Apr 1964, A. Quinonez (PA 667), 1M [UCLA]. Chir-

iqui Grande, Mata de Cacao, 17 Apr 1963, A. Quinonez (PA 220), 7M; same data (PA 221), 1 pM

(221-101), SL; same data (PA 224), 1 lp (224-105); same data except 21 Apr 1963 (PA 246), 1M,

1F [UCLA]. Chiriqui Grande, Punta de Pena, 14 Apr 1963, A. Quinonez (PA 193), 2 lpF (193-

101, 102), 5L, 1P [UCLA]. Isla Colon, Bocas del Toro (2km N), Big Creek, 10 Apr 1964, A. Qui-

nonez (PA 655), 1 IpF (665-102), 1P, 1L; same data except 11 Apr 1964 (PA 657), 1 IpF (657-

112), 1 pM (657-113), 2 pF (657-107, 114), 1L [UCLA]. Punta de Pena, Chiriquicito, 14 Apr

1963, A. Quinonez (PA 192), 1 pM (192-101), 4L; same data except 19 Apr 1963 (PA 232), 1 lpF

32 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

(232-101), 3L, 1P; same data except 22 Apr 1963 (PA 249), 3F; same data except 22 Apr 1963

(PA 250), 1F; same data except 23 Apr 1963 (PA 251), 1M, 5F; same data except 23 Apr 1963

(PA 252), 2F [UCLA]. Punta de Pena (3km S), El Guabo, 14 Apr 1963, A. Quinonez (PA 196),

1 IpM (196-121), 1P; same data (PA 199), 2 IpF (199-101, 103), 1 pM (199-102), 2F, 1P; same da-

ta (PA 201), 1 IlpM (201-103), 1 lpF (201-105), 2 pF (201-101, 102), 1 lp (201-106), 7L, 3P; same

data except 15 Apr 1963 (PA 202), 2 pM (202-101, 102), 1L; same data except 16 Apr 1963 (PA

208), 1F [UCLA]. Punta de Pena, Kaysan, 16 Apr 1963, A. Quinonez (PA 214), 2L; same data

except 18 Apr 1963 (PA 227), 1 lpM (227-103), 1 pM (227-102), 1 pF (227-101), 5L; same data

except 19 Apr 1963 (PA 239), 1 pM (239-102), 1 lp (239-101), 2L; same data except 20 Apr 1963

(PA 244), 5M, 2F [UCLA]. Locality unspecified, 25 Sept 1903, P. Osterhout, 1M, 1F [USNM].

Canal Zone: Ancon, 15 Jan 1916, L.H. Dunn, 1M [USNM]. Balboa, Corozal Dam site, 18 Sept

1943 (KO 30-21), 1F [UCLA]. Balboa, Orchid Gd., 3 Jan 1943, 1M [USNM]. Buenos Aires, San-

ta Rosa, 19 July 1972, J. H. Arnell, R. Hinds (PA 1135), 1 lp (1135-11), 1L [UCLA]. Camp Gail-

lard, 3 Jan-26 Dec 1925, D. Baker, 3M, 9F; 1 Jan 1925, J. B. Shropshire, 2F; thru C. S. Ludlow,

1M [USNM]. Cano Saddle, Gatun Lake, 10, 15 Aug 1923, H. G. Dyar, R. C. Shannon, 1M, 1F;

May-Sept 1923, R. C. Shannon, 4F [USNM]. Chilibre, Juan Mina, 20 July 1972, J. H. Arnell, R.

Hinds (PA 1138), 1F [UCLA]. Chilibre, Rio Chilibre, 21 Jan 1963, A. Quinonez (PA 26), 1 IpF

(26-106) [UCLA]. Corozal, 19 Aug 1913, 6 Dec 1919, J. Zetek, 1M, 1F; 24 Jan 1925, D. Baker,

1F [USNM]. Culebra, Nov 1924, 19 Dec 1925, D. Baker, 1M, 5F; 1918, L. H. Dunn, 2M, 1F; 14

Jan 1923, 24 Nov 1924, J. B. Shropshire, 1M, 3F; 30 Dec 1920, thru C. S. Ludlow, 1M [USNM].

Empire, 29 Nov 1925, D. Baker, 1M, 1F;6 Jan, 10 Dec 1923, J. B. Shropshire, 2M, 2F; 1918, L. H.

Dunn, 1M [USNM]. Fort Clayton, 17 Jan 1925, D. Baker, 1F [USNM];5 Nov 1944, K. E. Frick,

1M [CAS]; 5 Mar, 13 Jan, 2M, 6F [USNM]. Fort Clayton (4km N), Chiva Chiva, 10 Nov 1965, A.

Quinonez (PA 765), 1F, 1P, 1L; same data (PA 766), 1P [UCLA]. Fort Clayton, Chiva Chiva, 3.4

km N of Hwy C2 on Hwy C21, 12 Nov 1965, R. X. Schick, A. Quinonez (PA 776), 4 lpM (776-10,

11, 13, 20), 1 lp (776-14), 1L [UCLA]. Fort Davis, 10 Oct 1924, 18 Mar, 25 July 1925, D. Baker,

1M, 2F; 23 Dec, thru C. S. Ludlow, 1M, 2F; 23 Dec, 4M, 4F [USNM] ; 12 Mar, 28 May 1951, 3F

[UCLA]. Fort Randolph, 2F [USNM]. France Field, 20 Aug 1923, H. G. Dyar, R. C. Shannon,

1F; 18 Jan 1925, D. Baker, 1M, 1F [USNM]. Frijoles, Barro Colorado Island, 3 Dec 1965, A. Qui-

nonez (PA 851), 1 lpM (851-10); same data (PA 853), 2 lpM (853-10, 12), 1F, 1P, 1L; same data

except 4 Dec 1965 (PA 862), 1 lpM (862-15) [UCLA]. Gamboa, 22 Jan 1963, J. Labastid (CZ 4),

1 lpF (4-101); same data (CZ 5), 1 lpF (5-101); same data (CZ 6), 1 IpM (6-102), 2 lpF (6-103,

105), 3 pM (6-106, 115, 126), 1 pF (6-107), 1 P, 3L; same data (CZ 9), 1 IpF (9-101), 3L; same da-

ta (CZ 10), 1 pM (10-103), 1L; 24 Jan 1963, M. Keenan (CZ 11), 1 pF (11-102), 4L [UCLA].

Gamboa (15km NW), Frijoles, 1 Dec 1965, R. X. Schick, A. Quinonez (PA 843), 1 IpM (843-13), 1

IpF (843-12), 1 pM (843-100), 2 lp (843-10, 11); same data (PA 846), 1 lpM (846-10), 1 pM (846-

101), 1 pF (846-100), 1M, 1P [UCLA]. Gamboa, Summit, 12 Jan 1943 (KO 23-9), 1M [UCLA].

Gatun, 11 Mar 1908, June 1909, A. H. Jennings, 2F; 10 Dec 1921-Sept 1924, J. B. Shropshire, 2M,

SF; 21 Aug 1913, J. Zetek, 1M; 20 Sept, 6 Dec 1924, D. Baker, 1M, 2F; 24 Oct 1921, 29 Oct, 7

Jan, 2M, 3F [USNM]. Gorgona, 7 Feb 1908, A. H. Jennings, 1M, 1L [USNM]. La Pita, 26 Jan

1922, J. B. Shropshire, 1M [USNM]. Las Cascadas, 2 Feb 1908, Bath, 1M [USNM]. Lion Hill, A.

Busck, 2F [USNM]. Majagual, 20 Jan, 3 Feb 1923, J. B. Shropshire, 1M,3F [USNM]. Margarita,

27 Jan 1923, J. B. Shropshire, 1F [USNM]. Miraflores, 10 Oct 1921, J. B. Shropshire, 1M, 3F

[USNM]. Monte Lirio, 10 Dec 1921, J. B. Shropshire, 1F [USNM]. Nuevo Emperador (2km NE),

Hwy K16 1.1km NW of Hwy K19, 23 Nov 1965, A. Quinonez (PA 831), 2 IpM (831-10, 11), 1 IpF

(831-12), 1L; Nuevo Emperador (5km E), Hwy K10 3.5km NW of Hwy K15, 23 Nov 1965, A. Qui-

nonez (PA 822), 1 IpF (822-10), 1 pM (822-100), 1P, 1L; Nuevo Emperador (5km NE), junction of

Hwys K10 and K19, 23 Nov 1965, A. Quinonez (PA 828), 1 lpF (828-10); same data (PA 829), 1

pF (829-100); Nuevo Emperador (6km N), Hwy K16 2.7km N of Hwy K16A, 7 Dec 1965, A. Qui-

nonez (PA 869), 1L; Nuevo Emperador (7km N), old Hwy K16 5.3km N of Hwy K16A, 7 Dec

1965, A. Quinonez (PA 874), 1 lpM (874-10) [UCLA]. Paraiso, 1916, 1918, L. H. Dunn, 1M, 1F;

6 Oct 1913, J. Zetek, 1F [USNM]. Paraiso (across Canal from), Hwy K15 between Hwys K2 and

K6, 21 Nov 1965, A. Quinonez (PA 804), 1 lpF (804-10) [UCLA]. Paraiso (3km NW), Culebra, 13

Dec 1965, R. X. Schick (PA 899), 1 lpF (899-10); Paraiso (2km NW), Culebra, 13 Dec 1965, R. X.

Schick, A. Quinonez (PA 900), 1 IpM (900-10), 1L [UCLA]. Porte Grande, 13, 27 Jan 1923, J. B.

Zavortink: Reclassification of Trichoprosopon 33

Shropshire, 3M [USNM]. San Pablo, 12 Feb 1908, A. H. Jennings, 1F [USNM]. Santa Rosa,

Orange Plantation, 17 Nov 1936, L. E. Rozeboom (PAR 37), 2M [UCLA]. Summit, Oct 1946, N.

L.H. Krause, 1F [USNM]; 1 Nov 1939-17 Aug 1941, 4M [USNM] ; 12 Jan 1943, W.H.W. Komp

(KO 43), 2L [USNM] ; 1 Oct-3 Nov 1939, 11M, 10F [UCLA]. Summit (across Canal from), Em-

pire Firing Range, 18 Nov 1965, A. Quinonez, R. X. Schick (PA 796), 1L [UCLA]. Summit, Mad-

den Forest Preserve, Las Cruces Trail, 17 Sept 1964, A. Quinonez (PA 708), 1 lpF (708-10), 1 pM

~ (708-100); Summit, Madden Forest Preserve, Hwy C25 at Las Cruces Trail, 31 Oct 1965, J. N. Bel-

kin, R. X. Schick, M. Moody (PA 735), 1M, 1P, 1L; Summit, Madden Forest Preserve, S of junc-

tion of Hwy C25 and Las Cruces Trail, 31 Oct 1965, J. N. Belkin, R. X. Schick, M. Moody (PA

736), 1 IpM (736-20), 1 pM (736-102); Summit, Madden Forest Preserve, Hwy C25 3.7km N of

Hwy C2, 15 Nov 1965, R. X. Schick, A. Quinonez (PA 787), 2 lpM (787-10, 20), 2 lpF (787-22,

24), 2 pM (787-103, 105), 2 pF (787-102, 104), 2P, 18L; Summit, Madden Forest Preserve, S of

Green Park, 16 Nov 1965, M. Moody, A. Quinonez (PA 794), 2 pM (794-100, 101), 8L [UCLA].

Tabernilla, 28 Apr 1907, A. Busck, 1F (culicivora holotype); 4 Feb, 12 Mar 1908, A. H. Jennings,

2M, 1F; A. Busck, 1M [USNM]. Toro Point, 22 July 1922, 12 Jan 1923, J.B. Shropshire, 2M, 2F

[USNM], 1F [CU]. Locality unspecified, “Atlantic side,” 1F [UCLA]. Locality unspecified,

1920, 1M; A. H. Jennings, 1M; 14 Feb 1924, 1M, 1F; 14 Feb 1924, thru C. S. Ludlow, 1M, 1F

[USNM]; 1916, L. P. Dunn, 1F [CU]. Chiriqui: David, 19 July 1920, 1F [USNM]. The Falls, 11

Nov 1949, SF [UCLA]. Colon: Colon, Cativa, 22 Nov 1965, R. X. Schick, A. Quinonez (PA 806),

1 IpF (806-10) [UCLA]. Palmas Bellas, Pina, 30 Nov 1963, A. Quinonez (PA 575), 2L; same data

(PA 576), 3L [UCLA]. Portobelo, Mar 1941, A. Busck, 1F; 1 Sept 1923, H. G. Dyar, R. C. Shan-

non, 1F [USNM]; 4 Dec 1963, A. Quinonez (PA 580), 1 lpF (580-113); same data (PA 583), 1 pM

(583-101), 1 pF (583-102); same data (PA 584), 1 lp (584-101); same data except 6 Dec 1963 (PA

593), 1 lpM (593-106), 1 pM (593-107), 1 pF (593-105) [UCLA] ; same data except 8 Dec 1963

(PA 596), 2 lpF (596-106, 107), 2 pM (596-105, 109), 4 pF (596-101, 103, 104, 108), 2 lp (596-

102, 112), 2M, 2F, 5P, 57L [UCLA], 2L [BM] ; same data except 9 Dec 1963 (PA 599), 1 pF (599-

107) [UCLA]. Portobelo, Isla Caldera, 5 Dec 1963, A. Quinonez (PA 587), 1 IpF (587-102), 1L

[UCLA]. Salud, 6 Sept 1945, K. E. Frick, 3L [CAS]. Darien: El Real, Boca Yaviza, 9 Jan 1964,

A. Quinonez (PA 619), 4 IpM (619-107, 109, 113, 115), 5 IpF (619-119, 122-125), 4 pM (619-112,

117,118, 121), 4 pF (619-101, 104, 110, 120), 2 lp (619-122, 123), 33L [UCLA], 1 IpM (619-

114) [BM]. El Real, Pirre, 14 Jan 1964, A. Quinonez (PA 623), 2 lpM (623-115, 116), 1 pM (623-

103), 3 pF (623-106, 110, 113), 1 lp (623-114), 2F, 2P, 13L [UCLA], 1 pF (623-108) [BM] ; same

data except 15 Jan 1964 (PA 625), 1 pM (625-106), 1 lp (625-107), 1P, 13L; same data except 17

Jan 1964 (PA 628), 1 lpM (628-120), 2 IpF (628-103, 104), 2 pM (628-105, 108), 3 pF (628-109,

117, 119), 33L [UCLA]. La Palma (30km N), 0.8km from Santa Fe, 22 Nov 1966, O.G.W. Berlin

(PA 946), 1 IpM (946-10), 2 IpF (946-11, 12), 1L; La Palma (30km N), near Santa Fe, 9 Dec 1966,

O.G.W. Berlin (PA 993), 1 lpF (993-10); same data except 10 Dec 1966, O.G.W. Berlin, M. Mena

(PA 996), 1 lpF (996-10) [UCLA]. Morte (4km NE), Morti Hydro, 3 Dec 1966, O.G.W. Berlin (PA

976), 1L [UCLA]. Pucro, Camino Tapalisa, 24 Feb 1964, A. Quinonez (PA 629), 2 lpM (629-106,

116), 2 IpF (629-107, 110), 4 pM (629-102-105), 7 pF (629-108, 111-115, 117), 1 lp (629-109), 1

P, 55L [UCLA]. Pucro, La Laguna, 10 July 1963, A. Quinonez (PA 449), 1L; same data (PA 451),

1 pF (451-108), 1L [UCLA]. Pucro, Rio Tacarcuna Valley, 14 June 1963, A. Quinonez (PA 384),

1 IpF (384-110), 3L; same data except 15 June 1963 (PA 392), 1 lpM (392-103); same data except

20 June 1963 (PA 405), 1 lp (405-101); same data except 22 June 1963 (PA 414), 1 pF (414-108);

same data except 23 June 1963 (PA 416), 1F; same data except 8 July 1963 (PA 443), 1M [UCLA].

Pucro, Rio Tuira, 6 Mar 1958 (GG 78), 1 pM (78-103) [UCLA]. Pucro, Rio Tuira, mouth of Rio

Paya, 27 Mar 1958 (GG 38), 1 IpF (38-105), 1 pM (38-103), 1 pF (38-104) [UCLA]. Pucro, Tacar-

cuna, 1 Sept 1958 (GG 111), 1 lpM (111-113), 2 IpF (111-103, 107), 1 pF (111-109) [UCLA].

Santa Fe, 13 Apr, 19 May, 17, 19 Aug 1967, 2M, 3F [GML]. Sasardi, 7 Aug 1967, 1M [GML].

Locality unspecified, 7 July 1958 (GG 108), 1 lpM (108-123), 1 lpF (108-122), 2 pM (108-104,

112) [UCLA]. Panama: Chame, Nueva Gorgona, Playa Gorgona, 22 Aug 1963, A. Quinonez (PA

532), 1F [UCLA]. Chepo, 16 Oct 1939, 1F [UCLA]. Chiva (14km N), Cerro Campana, 29 Aug

1963, A. Quinonez (PA 539), 1F [UCLA]. Chilibre, Juan Mina, 18 Jan 1963, A. Quinonez (PA 6),

4 IpM (6-101, 104, 105, 108), 4 IpF (6-102, 103, 106, 107), 1L; same data except 20 Jan 1963 (PA

17), 6 IpM (17-106, 117, 119-121, 125), 1 IpF (17-127), 1 pF (17-108), 1L; same data except 24

34 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Jan 1963 (PA 41), 1 IpM (41-101), 1 pM (41-102), 10L [UCLA]. Huile (near, 3km N Nuevo Em-

perador), 4.8km NW junction Hwys K16 and K19, 23 Nov 1965, A. Quinonez (PA 835), 1 lpF

(835-10), 1 pF (835-102), 1P, 1L [UCLA]. La Chorrera, 5 Nov 1944 (ASM 260-2), 1F [UCLA].

Pacora, 15 Dec 1944, 14, 20 Aug 1945, K. E. Frick, 9M, 8F, 2P, 15L [CAS]. Pacora, Station B, 5

Sept, 7 Nov 1950, 7F [UCLA]. Panama City, 13 Sept 1926, D. P. Curry, 1F [USNM]. Pedregal,

near Tocumen, 19 Sept 1963, A. Quinonez (PA 556), 1 lpF (556-101) [UCLA]. Pequeni, Rio Pe-

queni, 9 May 1966, A. Quinonez (PA 941), 1 lp (941-10) [UCLA]. Pequeni (4km NE), Rio Pe-

queni, Estacion Hidro el Candelaria, 6 May 1966, A. Quinonez (PA 938), 2L [UCLA]. San Juan

de Pequeni, D. P. Curry, 9M, 3F [USNM]. Tocumen, Cerro Azul, La Zumbadora, 7 Feb 1963, A.

Quinonez (PA 66), 1 IM (66-103); same data except 8 Feb 1963 (PA 72), 1 IpF (72-101), 1L; same

data except 8 Feb 1963 (PA 74), 1 IpM (74-101); same data except 10 Feb 1963 (PA 84), 1 pM

(84-104), 1 lp (84-108); same data except 10 Feb 1963 (PA 85), 1 IpF (85-101); same data except

14 Feb 1963 (PA 92), 2L; same data except 15 Feb 1963 (PA 94), 1 pM (94-101); same data ex-

cept 18 Feb 1963 (PA 112), 1L [UCLA]. San Blas: Rio Cuadi, 17 June, 6 Nov 1967, 1M, 1F

[GML]. Province Unknown: Ahorea Lagarta, 31 Aug 1926, D. P. Curry, 1F [USNM]. Locality

Unspecified: 28 Apr 1907, A. Busck, 1 lp [USNM];1 July 1937, L. E. Rozeboom (PAR 118), 1M;

Hopkins, 1F; (Arnett 126-1), 1M, 1L [UCLA].

SURINAM. Para: Zanderij (Skm N), Republiek, 5 Nov 1963, D. C. Geijskes (SUR 34), 1F

[UCLA]. Paramaribo: Paramaribo, Charlesburg, 16 Feb 1940, D. C. Geijskes, 1F [SSM]. Locality

Unspecified: J. Bonne-Wepster, 1F [USNM].

TRINIDAD AND TOBAGO. TOBAGO. St. George: Mason Hall, Caledonia, 17 Nov 1965, T.

Aitken, R. Martinez, A. Guerra (TOB 48), 4F [UCLA]. Mason Hall, Trafalgar Estate, Easterfield, 2

% milepost, 29 Nov 1965, R. Martinez, A. Guerra (TOB 137), 3F; same data except 29, 30 Nov

1965 (TOB 147), 1F [UCLA]. Roxborough, 23 Nov 1965, R. Martinez, A. Guerra (TOB 91), 1

IpM (91-12) [UCLA]. Roxborough, Roxborough-Bloody Bay Road, 10 milepost, 23 Nov 1965, R.

Martinez, A. Guerra (TOB 97), 5F [UCLA]. TRINIDAD. Nariva: Mayaro, Bush Bush Forest, Na-

riva Swamp, 3 Nov 1964 (TR 806), 1 IpM (806-114), 1 pF (806-101) [UCLA]. Tabaquite, Charu-

ma Forest, 27 Aug 1964, A. Guerra (TR 635), 1 IpM (635-105), 1 pF (635-102), 1L; same data ex-

cept 8 Oct 1964 (TR 750), 1 lpM (750-105); same data except 8 Oct 1964 (TR 761), 1F [UCLA].

St. Andrew: Arima, Cumaca, 22 Oct 1964, A. Guerra (TR 784), 1F [UCLA]. Arima, Cumaca

Road, 14 Jan 1965, A. Guerra (TR 943), 1F [UCLA]. Rio Claro, 20 Aug 1964, A. Guerra (TR

616), 1 IpM (616-104); same data (TR 617), 1 lpM (617-100) [UCLA]. Sangre Grande, Biche, 9

July 1964, F. Powdhar (TR 556), 1 IpM (556-122) [UCLA]. Sangre Grande, Cumaca, 3 Sept 1964,

A. Guerra (TR 650), 2 pM (650-110, 113) [UCLA]. Sangre Grande, Cumaca Road, Pamponette

Trace Junction, 20 May 1965, A. Guerra (TR 1170), 1F [UCLA]. Sangre Grande, Mt. Harris, 16

July 1964, F. Powdhar (TR 573), 1F [UCLA]. Sangre Grande, Nestor Village, 12 June 1964, A.

Guerra (TR 484), 3F [UCLA]. Sangre Grande, Nestor, 6% milepost, 24 June 1965, F. Powdhar

(TR 1232), 1F [UCLA]. Sangre Grande, Turure Forest, Oct 1966, F. Guerra (TR 1618), 1F; same

data except Nov 1966 (TR 1619), 1F [UCLA]. Sangre Grande, Valencia Old Road at 4.5 milepost,

24 Sept 1964, A. Guerra, 1F [UCLA]. Sangre Grande, Valencia Old Road, 1 Oct 1964, A. Guerra

(TR 744), 1F [UCLA]. St. David: Toco, Matelot, 12 Mar 1964, A. Guerra (TR 182), 1 lpM (182-

125) [UCLA]. Toco, Tompire Southbank Road, 27 Feb 1964, A. Guerra (TR 114), 1F [UCLA].

St. George: Arima, Andrews Estate, 25 Mar 1965, A. Guerra (TR 1060), 1L [UCLA]. Arima,

Arena Forest, 13 Aug 1965, A. Guerra (TR 1326), 1 lpM (1326-20); same data except 20 Aug 1965

(TR 1355), 1M [UCLA]. Arima, Aripo Valley, 19 Mar 1964, A. Guerra, 3F [UCLA]. Arima,

Calvary Hill, 30 Aug 1965, A. Guerra (TR 1379), 1F [UCLA]. Arima, Las Hermanas Estate, 18

Aug 1965, A. Guerra (TR 1336), 1 pF (1336-100), 1L [UCLA]. Arima, Mt. Becke, 29 Apr 1965,

A. Guerra (TR 1132), 1F [UCLA]. Arima, Verdant Vale, 10 Sept 1964, A. Guerra (TR 686), 1F;

same data except 12 Nov 1964 (TR 831), 2F [UCLA]. St. Augustine, 20 Feb 1925, C. L. Withy-

combe, 1F [BM]. County Unknown: Tuencha, 25 May 1926, C. L. Withycombe, 1F [BM]. For-

est Reserve, 5 Dec 1965, T.H.G. Aitken (TR 1430), 1L [UCLA].

VENEZUELA. Aragua: Choroni (1.0km N), 16 July 1969, T. J. Zavortink (VZ 219), 2 lpF

(219-14, 16), 1 pF (219-102), 3L [UCLA]. Guamitas, 12 Aug 1927, M. Nunez Tovar, 1M [USNM].

Maracay, M. Nunez Tovar, 1F [USNM]. Maracay, Guamitas, road to picnic area, 15 July 1969, T.

J. Zavortink, J. A. Bergland (VZ 199), 1 lpM (199-11), 1 IpF (199-10) [UCLA]. Carabobo: Cara-

Zavortink: Reclassification of Trichoprosopon 35

bobo, 30 July 1927, M. Nunez Tovar, 1M [USNM]. Puerto Caballo, Borburata, 24 July 1969, J.

Pulido, J. Valencia (VZ 263), 1 lpF (263-17), 1 pF (263-101) [UCLA]. Monagas: Caripito, 13

Mar, 2 June 1942, 2F [AMNH]. Sucre: Guanoco, Green House Bay, 1 Jan 1929, E. de Verteuil,

W. Cook, F. W. Urich, 1F [USNM]. Zulia: Colon Dist., 18 Dec 1933, P. Anduze, 1F; May 1934,

1F [CU]. La Rivera, July 1934, P. Anduze, 1F [CU]. Locality Unspecified: 10 Feb 1973, CDC

(VZ 420), 2F [UCLA]; P. Anduze, 1M; No. 2345, 1M [USNM].

NO DATA. D. P. Curry, 2M, 2F; No. 1447, 1M; 2M, 1L [USNM]; No. 168, 1P, 1M [CU].

2. Johnbelkinia longipes (Fabricius)

Figs. 2, 8-10

1805. Culex longipes Fabricius 1805:34. *TYPE: Holotype male, America meridionali, Smidt

[J. C. Schmidt] [ZMC; type locality restricted to Cayenne [Guyane] , French Guiana, by

Belkin, Schick and Heinemann 1965:41-42]. NEW COMBINATION.

1919. Lesticocampa trichopus Dyar 1919: 10-11. *TYPE: Holotype female, Teffé [Tefe], Ama-

zonas, Brazil, June 1906, A. Ducke [USNM, 21996]. Considered as doubtfully distinct

from longipes by Dyar (1928:97); synonymized with longipes by Lima (1931:66-67).

1942. Trichoprosopon (Hyloconops) edwardsianus Lane and Cerqueira 1942:528. TYPE: Holo-

type female, Belterra, Santarem, Pard, Brazil, Sept 1938 [IOC]. NEW SYNONYMY.

Trichoprosopon (Runchomyia) longipes of Stone, Knight and Starcke (1959:76-77, in part); Faur-

an (1961:14-15, in part); Forattini (1965:169, 170, 172, 173, 174-175, 176, 217, 291, in part);

Belkin, Schick and Heinemann (1965:41-42); Cova Garcia, Sutil and Rausseo (1966a:66, 68,

in part; 1966b:67, 131, 384-385, in part); Belkin (1968:36, 46, 58); Knight and Stone (1977:

314, in part).

Trichoprosopon (Rhunchomyia) longipes of Lane (1953:855-857, in part).

Trichoprosopon (Rhynchomyia) longipes of Floch (1956:35, 221, in part).

Trichoprosopon (Hyloconops) longipes of Lane and Cerqueira (1942:525, in part).

Goeldia (Goeldia) longipes of Edwards (1932:72, in part); Senevet and Abonnenc (1939:248-250);

Lane (1939: 162-163, in part); Floch and Abonnenc (1947:2-3, in part).

Goeldia longipes of Bonne-Wepster and Bonne (1923:124, in part); Bonne and Bonne-Wepster

(1925:144-146, in part); Dyar (1928:96-97, in part); Lima (193 1:66-67, in part); Komp (1936:

60, in part); Antunes (1937a:72-73, in part); Patino Camargo (1937:242, in part); Boshell-Man-

rique (1938:415, in part); Floch and Abonnenc (1942b:2-3, in part); Floch (1955:315, in part).

Culex longipes of Wiedemann (1828:7, 546); Macquart (1838:34-35, in part); Zimsen (1964:451).

Culex longipede of Macquart (1834:36).

Sabethes longipes of Robineau-Desvoidy (1827:412); Lynch Arribalzaga (1891:167-168, in part);

Hunter (1900:281, in part).

Trichoprosopon (Runchomyia) edwardsianum of Stone, Knight and Starcke (1959:76); Cova Gar-

cia, Sutil and Rausseo (1966a:67, 68; 1966b:131, 384); Belkin, Schick and Heinemann (1971:

9, 36, 47, 53); Knight and Stone (1977:313).

Trichoprosopon (Rhunchomyia) edwardsianum of Cerqueira (1961a:459-464).

Trichoprosopon (Rhunchomyia) edwardsianus of Lane (1953:860); Barreto-Reyes (1955:81); Cer-

queira (1961b:154); Xavier and Mattos (1970:452; 1975:255; 1976:402).

Trichoprosopon (Hyloconops) edwardsianus of Lane (1945:133).

Trichoprosopon edwardsianum of Forattini, Rabello and Cotrim (1970:78).

Trichoprosopon edwardsianus of Maia (1963:91).

Goeldia edwardsianus of Horsfall (1955:319).

Trichoprosopon (Runchomyia) trichopus of Belkin, Schick and Heinemann (1971:9, 34, 50, 51).

Goeldia trichopus of Bonne-Wepster and Bonne (1921:18); Dyar (1921:27); Bonne-Wepster and

Bonne (1922:38); Dyar (1923a:81); Bonne and Bonne-Wepster (1923:124, in part); Dyar and

Shannon (1924:483); Bonne and Bonne-Wepster (1925:141-143, in part); Dyar (1928:97); Mar-

tini (1931:201), Evans and Walker (1935:465-466); Lane (1936:7); Antunes (1937b:72); An-

duze (1941:9).

36 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

Goeldia aff. trichopus of Townsend (1934:492).

Lesticocampa trichopus of Stone and Knight (1957:119).

?Trichoprosopon (Runchomyia) longipes of Martinez and Prosen (1953:31); Morales-Ayala (1971:

139).

?Goeldia longipes of Bonne (1923:129).

?Trichoprosopon (R.) edwardsianus of Suarez (1963:239).

?Trichoprosopon (Hyloconops) trichopus of Anduze, Pifano and Vogelsang (1947:12).

?Goeldia (Goeldia) trichopus of Chagas, da Cunha et al. (1938:198, 202); Floch and Abonnenc

(1947:3).

?Goeldia trichopus of Kumm and Novis (1938:503, 512); Floch and Abonnenc (1942a:3; 1942b:3).

?Trichoprosopon (Runchomyia) leucopus of Fauran (1961:14); Stone (1961:33); Morales-Ayala

(1971:139); Kruijf (1972:50).

?Trichoprosopon (Rhunchomyia) leucopus of Xavier and Mattos (1975:256).

?Goeldia (Goeldia) leucopus of Floch and Abonnenc (1947:3).

?Goeldia leucopus of Floch and Abonnenc (1942a:3; 1942b:3).

FEMALE. Wing: 4.60mm. Proboscis: 4.20mm. Femur I: 3.30mm. Abdomen:

about 4.4mm. Thorax: Scutal scales usually dull brown or weakly iridescent copper,

bronze, gold, green, blue or purple. Postnotum usually without scales. Pleural integ-

ument usually slightly to moderately darkened on ssp and stp, sometimes strongly

darkened. Legs: Leg III with conspicuous outstanding scales on apical part of tibia

and on tarsus 1-3 or 4-III, these most conspicuous on tarsus 2-III. Tarsus 2-II with

yellowish-cream to white scales in streak in apical 0.1-0.8 of anterior and dorsal sur-

faces, or in complete band in apical 0.1-0.8, or completely covering segment; 3,4-II

completely yellowish-cream to white scaled; 5-I] with yellowish-cream to white scales

restricted to streak at base of anterior and dorsal surfaces or completely covering seg-

ment. Tarsus 3-III dark scaled or white scaled on apical 0.1-0.6; 4,5-III completely

white scaled. Wing: Upper calypter with short row of setae basally. Abdomen:

Sternites II-VII usually with dark scales in median line from base to apex or in medi-

an apical patch.

MALE (fig. 8). Essentially as in female except for sexual characters. Head: Fla-

gellum moderately plumose, middle segments with about 24 bristles. Legs: Anterior

claw I enlarged, with distinct, long submedian tooth; anterior claw II not enlarged,

simple.

MALE GENITALIA (figs. 8, 10). Segment IX: Tergite lobe much longer than

wide. Setae of lobe 4 or 5 (3-5), strongly to very strongly developed, somewhat flat-

tened apically, with short filamentous apical prolongation. Sternite U-shaped. Side-

piece: Slender to stout. Densest region of basal tergal setae proximad of level of

apex of basal mesal lobe. Basal sternomesal setae 2-4, moderately long, curved or an-

gularly bent at apex; strongly developed medial sternomesal setae 2-4. Basal Mesal

Lobe: Body conical, with long narrow dorsal strip; setae directed dorsad, very

strongly curled at apex. Clasper: Long; constricted basally; slightly to conspicuously

swollen apically; with several fine setae apically. Spiniform bent at apex; slender.

Phallosome: Aedeagus moderately pigmented; distal lateral margin usually not con-

cave in dorsal aspect; distal end of submedian tergal arm angled or curved toward

mate; apical tergal arm with patch of 4-11 small, strongly sclerotized, dorsal preapical

teeth and serrulate or striate, laterally curved, sclerotized apical process. Proctiger:

Paraproct with 4-7 (3-7) moderately large, dorsally curved, apical teeth and with

crenulate, moderately sclerotized lamella mesad of apical teeth that may project dis-

tad beyond more ventral teeth. Cercal sclerite absent or present, elongate.

Zavortink: Reclassification of Trichoprosopon 37

PUPA (fig. 8). Abdomen: 4.4mm. Trumpet: 0.51mm. Paddle: 0.68mm. Cepha-

lothorax: Very weakly pigmented. Trumpet: Very weakly to weakly pigmented,

sometimes becoming slightly darker distad. Abdomen: Predominantly very weakly

pigmented; sternites II-IV (II-VII) with median part of anterior margin moderately

pigmented, conspicuously darker than remainder of ventral surface. Terminal Seg-

ments: Male genital lobe large, its width usually greater than 0.50 (0.47-0.60) of seg-

ment VIII. Paddle: Very weakly pigmented. Index usually 1.26-1.59 (1.21-1.63).

LARVA (fig. 9). Head: 1.18mm. Siphon: 1.21mm. Anal Saddle: 0.23mm.

Head: Very weakly pigmented, collar darker. Mental plate with 11,12 (10-14) teeth

on each side of median tooth. Antenna: Without spicules or with very few small

spicules medially. Abdomen: Seta 6-I usually 11-14b (7-14); seta 6-II usually 10-14

b (8-16); seta 6-III usually 9-12b (8-14). Segment VIII: Comb scales usually 33-54

(9-65); most comb scales of patch usually with moderately large apical spine. Si-

phon: Very weakly pigmented, becoming darker distad. Moderately to strongly

curved dorsad distally. Seta 1-S inserted 0.51-0.67 distance from base of siphon. Ac-

cessory midventral setae (la-S) usually 30-36 (25-37). Anal Segment: Saddle very

weakly pigmented, indistinct. Seta 3-X usually 6-9b (5-10). Seta 4-X strongly devel-

oped, usually 10-15b (6-17), with alveolar plate.

SYSTEMATICS. The adult of Johnbelkinia longipes differs from ulopus and leu-

copus by: (1) more strongly plumose flagellum of male, (2) distinct tooth on anterior

claw I and small anterior claw II of male, (3) usually extensive dark scaled areas on

sternites and (4) more extensive distribution of outstanding scales on leg III, these

being present on apical part of tibia and all of tarsus 1-III. The adult differs addition-

ally from leucopus by: (1) lighter integument on mep and meron and (2) entirely

white scaled tarsus 4-III. It differs additionally from ulopus by: (1) presence of setae

on upper calypter of wing and (2) duller brown scales on scutum. The male genitalia

differ from ulopus and leucopus by: (1) long tergite IX lobe, (2) shape of clasper,

which is constricted basally and swollen apically and (3) presence of small but strong-

ly sclerotized teeth and sclerotized apical process on apical tergal arm of aedeagus.

The pupa differs from ulopus by: (1) darkened integument on median anterior part

of sternites IJ-IV and (2) larger male genital lobe, which is usually broader than 0.50

width of segment VIII. The larva differs from ulopus by: (1) seta 4-X usually 10-15b

(6-17) and with alveolar plate, (2) seta 6-I 11-14b (7-14) and (3) moderately large

apical spine of comb scales.

There is variation in tarsus III markings, color of the pleural integument and male

genital structure in this species. Specimens from the Guianas and Colombia have tar-

sus 3-III entirely dark scaled or light scaled in the apical 0.1 only. Most specimens

from Para, Brazil, have tarsus 3-III light scaled in the apical 0.2-0.4, but one from

Santarem has it dark scaled. Most specimens from Ecuador, Peru and Bolivia have

this segment white scaled in the apical 0.3-0.6, but again one specimen from Ecuador

has it dark scaled. Two males from the Rio Madeira drainage of Brazil and Bolivia

have the integument of ssp and stp much more strongly darkened than any other

specimens I have seen. These same males exhibit some unusual variations in the male

genitalia, for the sidepiece is slender, the clasper less curved and less swollen apically,

the aedeagus has fewer preapical teeth but a more distinctly serrulate apical process,

the cercal sclerite is present and the apical lamella of the paraproct does not extend

beyond the ventral paraproct teeth. It is quite possible these two males represent an-

other species, but since this is not absolutely clear at this time, I prefer to include

them in /ongipes until more material is available.

Much of the Amazon basin was flooded or very arid at various times in the Pleisto-

38 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

cene, and populations of longipes that exist there now must have invaded the area

from forest refugia in mountains surrounding the basin only relatively recently. In

fact, all the variants known in longipes probably differentiated in separate refugia in

times past when the tropical rain forest was not continuous.

I have studied two slide mounted third instar larvae of Jongipes, and these resem-

ble ulopus closely.

BIONOMICS. The larvae of longipes have been reported in the literature as occur-

ring in the leaf axils of Mauritia sp. (Palmae) by Cerqueira (1961a: 459-464, as ed-

wardsianum) and Musa bihai |=Heliconia bihai, Strelitziaceae] by Senevet and Abon-

nenc (1939:248-250). Immatures collected for the project “Mosquitoes of Middle

America” in the Guianas were found in the leaf axils of Heliconia sp.and Phenako-

spermum guianense (Strelitziaceae) and once in the flower bracts of Heliconia sp.

The immatures were collected from leaf axils of banana (Musaceae) in Brazil by John

F. Reinert.

On two separate occasions males of Jongipes were observed flying backwards and

forwards about three feet above the ground in a patch of heliconias in Brazil by

Evans and Walker (1935:465-466, as trichopus). Adults have been collected in rest-

ing traps and malaise traps in the forest and females have been collected biting hu-

mans.

Numerous papers reporting observations on longipes or trichopus in South Amer-

ica probably refer at least in part to ulopus, and they are summarized under that spe-

cies.

DISTRIBUTION. Johnbelkinia longipes is apparently restricted to the portion of

the northern half of South America that drains into the Atlantic Ocean. In the east-

ern part of the continent, I have seen specimens from the state of Monagas, Venezue-

la, in the north to the state of Para, Brazil, in the south. In the western part, I have

seen specimens from the department of Meta, Colombia, in the north to the depart-

ment of Beni, Bolivia, in the south. Cerqueira (1961b:154) records this species (as

edwardsianus) in the south from the state of Mato Grosso, Brazil. Every specimen of

Johnbelkinia from Trinidad that I have examined is ulopus. However, since longipes

in known from directly across the Gulf of Paria in Venezuela, it may occur also in

Trinidad. Material examined: 281 specimens; 25 males, 166 females, 49 larvae, 41

pupae; 38 individual rearings (29 larval, 6 pupal, 3 incomplete).

BOLIVIA. Beni: Guajaramerim, Vaca Diez, Mar 1939, 1F [USNM]. Locality unspecified:

May 1943, H. P. Carr, 1M [USNM].

BRAZIL. Amazonas: Humaita, Nov 1937, 1F [BM]. Manaus, W. M. Mann, Baker, 1F [USNM].

Tefé, June 1906, A. Ducke, 1F (trichopus holotype) [USNM]. Para: Altamira (160km W), 6 Nov

1974, J. F. Reinert (81-A), 1L [MEP]. Belterra, Santarem, Sept 1938, 1F [USNM]. Boa Vista,

Tapajos, C.H.T. Townsend, 29 Sept 1932, 1M [USNM]. Cameta, Jan 1936, 1F [USNM]. Curralin-

ho, H. W. Kumm, 1F [BM]. Curralinho, Rio Itauca, 1F [USNM]. Jatobal, Bacuri, 28 Oct 1974, J.

F. Reinert (63), 1F [MEP]. Marituba, Sta. Isabel, Feb 1938, 1F [USNM]. Rondonia: Guajara

Mirim, Rio Madeira, May 1931, R. C. Shannon, 1M [USNM].

COLOMBIA. Meta: ?Chipadera, Nov 1936 (COR 107), 2F; 12 Nov 1936 (COR 109), 1F

[UCLA]. ?Maragal, 14 Nov 1936 (COR 113), 1F [UCLA]. Restrepo, 1935 (KO 30-7), 1F;3 Sept

1935 (KO 4-16), 1F; 10 July 1935 (KO 30-36), 1F [UCLA]. Restrepo, Vega Grande, 27 Dec 1936

(COR 74, 75), 3F [UCLA]. Villavicencio, Bosque Ocoa, 6 June 1944 (COK 42), 1F; 14 June 1944

(COK 44), 11F; 20 June 1944 (COK 47), 5F [UCLA]. Villavicencio, Palmichal, 1936 (COR 91),

1F; 9 Nov 1936 (COR 98), 1F [UCLA]. Locality unspecified, 12 Nov 1936 (COR 108), 1F; 19

Nov 1936 (COR 115), 2F; Nov 1936 (COR 123, 125), 8 F; 20 Oct 1936 (COR 128), 1F; 21 Oct

1936 (COR 130), 10F; 23 Oct 1936 (COR 132, 133), 2F; 8 Nov 1936 (COR 136), 6F; 9 Nov 1936

(COR 138), 1F; 10 Nov 1936 (COR 140), 1F; 27 Nov 1936 (COR 161), 6F; 22 Dec 1936 (COR

175, 176), 4F [UCLA].

Zavortink: Reclassification of Trichoprosopon 39

ECUADOR. Napo: Coca and Napo Rivers, 23 Apr-12 May 1965, L. E. Pena (ECU 8), 1M, 4F

[UCLA]. Napo, Pompeya Island, Napo River, 19-26 May 1965, L. E. Pena (ECU 19), 1F [UCLA].

FRENCH GUIANA. Guyane: Cayenne, Smidt, 1M (longipes holotype) [ZMC]. Cayenne, Apr

1943, 2F [UCLA]. Cayenne, foret de Cabassou, 14 Nov 1967, J. Clastrier (FGC 3278), 1F; 12

July 1968, J. Clastrier (FGC 3911), 1F; 8 Sept 1968, J. Clastrier (FGC 3636), 1 IpF (3636-40);

same data (FGC 3639), 3L; same data (FGC 3640), 1L [UCLA]. Cayenne, foret de la Chaumiere,

7 Aug 1968, J. Clastrier (FGC 3580), 1L; same data (FGC 3581), 1 IpF (3581-31); same data (FGC

3592), 1 IpM (3592-32) [UCLA]. Cayenne (near), Ile de Cayenne (FGA 154), 1F [UCLA]. Cay-

enne (24km SW), Le Gallion, 1 Feb 1965, T.H.G. Aitken, R. Martinez, A. Guerra (FG 20), 2 lpF

(20-20, 21); same data except 2 Feb 1965 (FG 30), 1 IpM (30-10), 1 IpF (30-13), 1 lp (30-11)

[UCLA]. Cayenne, Roura, 13 Sept 1968, J. Clastrier (FGC 3648), 1L; same data (FGC 3652), 1

IpM (3652-44); same data (FGC 3653), 1 pM (3653-40); same data (FGC 3656), 1 lpF (3656-31)

[UCLA]. Couachi, Mana (10km E), 15 Mar 1969, J. Clastrier (FGC 4021), 1L; same data (FGC

4023), 1L [UCLA]. Le Gallion (3km W), Pont des Cascades, 16 Mar 1967, R. X. Schick, J. Fred-

erick (FG 151), 1 IpM (151-10), 1 lpF (151-30); same data (FG 152), 1 lIpM (152-10), 1L [UCLA].

Matoury, 2F [UCLA]. Matoury, Camp Rochambeau (E edge), 19 July 1968, J. Clastrier (FGC

3538), 1 pF (3538-30); same data (FGC 3540), 1 pM (3540-12); same data (FGC 3543), 1 IpF

(3543-34); same data (FGC 3544), 1 IpF (3544-30); same data (FGC 3545), 1L; same data (FGC

3553), 1 IpM (3553-30) [UCLA]. Matoury, foret de Cogneau, 28 May 1968, J. Clastrier (FGC

3459), 1 pF (3459-10); 25 Sept 1968, J. Clastrier (FGC 3663), 1 pF (3663-41); same data (FGC

3665), 1M; same data (FGC 3666), 1L [UCLA]. Matoury, Matoury A, 10 July 1968, J. Clastrier

(FGC 3503), 1L [UCLA]. Matoury, Matoury C, 20 Nov 1968, J. Clastrier (FGC 3802), 1 lpF

(3802-54); same data (FGC 3803), 1 IpF (8803-26) [UCLA]. Montjoly, Cabassou, Ile de Cayenne,

2 July 1952 (FGA 191), 1F [UCLA]. Montjoly, Raban, Ile de Cayenne, base of Montagne Tigre

(FGA 43), 1 pF (43-10), 1L [UCLA]. Montsinery, Tonegrande, 6 Sept 1968, J. Clastrier (FGC

3628), 1 lpM (3628-43) [UCLA]. Remire (SW), Chemin Vidal, 14 Mar 1967, R. X. Schick (FG

129), 2 IpM (129-10, 11); same data (FG 131), 1 IpF (131-10) [UCLA]. Rochambeau, 1F [UCLA].

GUYANA. Demerara: Madewiri Creek, Mar 1941, 1F [UCLA].

PERU. Huanuco: Tingo Maria, 21 Oct 1946, J.C. Pallister, 1F [AMNH]; date unspecified, E.

J. Hambleton, 3M, 1F [USNM]. Loreto: Chimbote, Amazon, Mar 1931, R. C. Shannon, 1F

[USNM]. Iquitos, Mar-Apr 1931, R. C. Shannon, 45F, 1 Ip, lp, 2L [USNM]. Pachitea Mund, 6

Nov 1903, 1F [BM].

SURINAM. Para: Onverwacht (S5km S), Wegnaar Overtoon, 7 Apr 1967, R. X. Schick, H.A.M.

de Kruijf (SUR 228), 1L; same data (SUR 229), 1 lp (229-10) [UCLA]. Zanderij (2km N), 13 Apr

1967, R. X. Schick, H.A.M. de Kruijf (SUR 242), 2 IpM (242-10, 11); same data (SUR 242A), 1P;

same data (SUR 243), 1P; same data (SUR 245), 1 IpM (245-11), 1 lpF (245-10); same data (SUR

246), 2 IpM (246-10, 12), 1 IpF (246-11); same data (SUR 247), 1 lpF (247-10) [UCLA]. Locality

unspecified: 25 Sept 1946, D. C. Geijskes, 1F [SSM].

VENEZUELA. Monagas: Caripito, 10 Mar 1942, 1F [AMNH].

3. Johnbelkinia leucopus (Dyar & Knab)

Figs. 2, 10

1906. Lesticocampa leucopus Dyar and Knab 1906:137. *TYPE: Lectotype female, Bluefields

[Zelaya] , Nicaragua, W. F. Thornton [USNM, 10003; designation by Stone and Knight

1957:118]. NEW COMBINATION.

Trichoprosopon (Runchomyia) leucopus of Stone, Knight and Starcke (1959:76, in part); Belkin,

Schick and Heinemann (1965:42); Cova Garcia, Sutil and Rausseo (1966a:67, 68, in part);

Knight and Stone (1977:314, in part).

Trichoprosopon (Rhunchomyia) leucopus of Lane (1953:857-858, in part).

Trichoprosopon (Hyloconops) leucopus of Lane and Cerqueira (1942:526).

Trichoprosopon leucopus of Arnett (1949:245).

40 Contrib. Amer. Ent. Inst., Vol. 17, no. 1, 1979

Goeldia (Goeldia) leucopus of Dyar (1923a:86, in part; 1923b:174); Edwards (1932:72); Lane

(1939:161).

Goeldia leucopus of Bonne-Wepster and Bonne (1921:17; 1922:38); Dyar and Shannon (1924:

483); Bonne and Bonne-Wepster (1925:152, in part); Dyar (1925:129, in part; 1928:97-98, in

part); Lane (1936:7); Antunes (1937b:72); Kumm, Komp and Ruiz (1940:397); Horsfall (1955:

319).

Lesticocampa leucopus of Busck (1908:76); Theobald (1910:622); Howard, Dyar and Knab (1915:

170-172, in part); Stone and Knight (1957:118).

?Trichoprosopon (Rhunchomyia) leucopus of Trapido and Galindo (1957:122, 124, 125, 128,

131, 134, 135, 138).

?Trichoprosopon leucopus of Galindo and Trapido (1957:146).

FEMALE. Wing: 4.60mm. Proboscis: 4.04mm. Femur I: 3.47mm. Abdomen:

about 3.7mm. Thorax: Scutal scales usually dull brown or weakly iridescent copper,

bronze, gold, green, blue or purple. Postnotum often with dark iridescent scales.

Pleural integument strongly darkened on ssp, psp, stp, mep and meron. Legs: Leg

III without outstanding scales. Tarsus II with streak of light scales on anterior sur-

face of apical segments, streak varying from inconspicuous, with creamy-tanish scales

from apical part of 3-II to base of 5-II, to conspicuous, with white scales from apex

of 2-II to apex of 5-II. Tarsus III with white scales on apical 0.1-0.4 of 4-III and all

of 5-III. Wing: Upper calypter with short row of setae basally. Abdomen: Sternites

II-VII entirely light scaled.

MALE (fig. 10). Essentially as in female except for sexual characters. Head: Fla-

gellum sparsely plumose, the middle segments with about 16 bristles. Legs: Anterior

claw I enlarged, with submedian crestlike swelling; anterior claw II enlarged, simple.

MALE GENITALIA (fig. 10). Segment IX: Tergite lobe about as long as wide.

Setae of lobe 6 or 7, moderately to strongly developed, flattened apically, with long

filamentous apical prolongation. Sternite V-shaped. Sidepiece: Stout. Densest re-

gion of basal tergal setae proximad of level of apex of basal mesal lobe. Basal sterno-

mesal setae 4, moderately long, angularly bent at apex; strongly deveolped medial

sternomesal setae absent. Basal Mesal Lobe: Body conical, with long narrow dorsal

strip; setae directed dorsad, very strongly curled at apex. Clasper: Moderately long;

without basal constriction; uniformly slender beyond base; without setae. Spiniform

straight, slender. Phallosome: Aedeagus weakly pigmented; distal lateral margin not

concave in dorsal aspect; distal end of submedian tergal arm nearly parallel with mate;

apical tergal arm without dorsal preapical teeth or sclerotized apical process. Procti-

ger: Paraproct with 4 moderately large, dorsally curved apical teeth and with moder-

ately sclerotized crenulate apical lamella mesad of apical teeth that does not project

beyond teeth. Cercal sclerite present, elongate.

PUPA, LARVA. Unknown.

SYSTEMATICS. Johnbelkinia leucopus combines characteristics of ulopus and

longipes. From the apparently more closely related but allopatric Jongipes it differs

by the following characters: in the adult most conspicuously by: (1) presence of dark

scales in basal 0.6-0.9 of tarsus 4-III, (2) absence of outstanding scales on leg III, (3)

strongly darkened integument on mep and meron, (4) crestlike submedian swelling

on anterior claw I and enlarged anterior claw II of male and (5) light scaled sternites;

and in the male genitalia by: (1) shorter tergite IX lobe, (2) shape of clasper, which is

without a basal constriction and is uniformly slender beyond base, (3) absence of

dorsal preapical teeth or sclerotized apical process on apical tergal arm of aedeagus

and (4) absence of strongly developed medial sternomesal setae. From ulopus, with

which it is sympartic, leucopus is distinguished as follows: in the adult by: (1) pres-

Zavortink: Reclassification of Trichoprosopon 4]

ence of setae on upper calypter of wing and (2) duller brown scutal scales; and in the

male genitalia by: (1) shape of basal mesal lobe, which has a conical body and a long

narrow dorsal strip, (2) dorsally directed, strongly curled setae of basal mesal lobe,

(3) location of dense patch of basal tergal setae proximad of level of apex of basal

mesal lobe and (4) stouter sidepiece.

Osterhout collected two females and one male of Johnbelkinia in Bocas del Toro,

Panama, in 1903. All three specimens were determined as Jeucopus by Dyar and

Knab, and the genitalia of the male were illustrated repeatedly (Howard, Dyar and

Knab 1913:fig. 47; Dyar 1928:fig. 66; Lane 1953:fig. 849; Cova Garcia, Sutil and

Rausseo 1966a:fig. 124). However, only one of the females is actually Jeucopus; the

other one and the male are ulopus.

I have not noted any significant variations in the few specimens available for study.

BIONOMICS. Very little is known about this species. Kumm, Komp and Ruiz

(1940:397) reported taking adults in daytime captures in the forest in Costa Rica.

The single male collected for the ““Mosquitoes of Middle America” project was taken

resting in the forest between 0700 and 1230 hours on 17 April 1963; the two fe-

males collected for the project were attracted to man in a forest between 1100 and

1400 hours on 21 April 1963 at a height of 12m above the ground.

The reports of captures of leucopus at human bait in the Caribbean versant of Nic-

aragua and western Panama by Galindo and Trapido (1957:146) and Trapido and

Galindo (1957:122, 124, 125, 128, 131, 134, 135, 138) refer in part or whole to

ulopus and are summarized under that species.

DISTRIBUTION. Johnbelkinia leucopus is apparently a rare, relictual species re-

stricted to the Caribbean Coast of Central America from central Nicaragua to west-

ern Panama and the Pacific Coast of Central America in southern Costa Rica. Most

reports of this species in Central America actually pertain to ulopus; I do not know

upon what species most reports from South America are based. Material examined:

16 specimens; 1 male, 15 females.

COSTA RICA. Limon: Liverpool, H. W. Kumm, 2F [BM]; 1F [USNM]. Puntarenas: San Vi-

to (6km S$), 19-30 Apr 1967, D. F. Veirs, 4F [UCLA].

NICARAGUA. Zelaya: Bluefields, W. F. Thornton, 4F (including holotype) [USNM]. Rio

Curinguas (60mi up), 27 Nov 1941, 1F [UCLA].

PANAMA. Bocas del Toro: Chiriqui Grande, Mata de Cacao, 17 Apr 1963, A. Quinonez (PA

220), 1M [UCLA]. Punta de Pena, Chiriquicito, 21 Apr 1963, A. Quinonez (PA 247), 2F [UCLA].

Locality unspecified, 25 Sept 1903, P. Osterhout, 1F [USNM].

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Mision Estud. Patol. Reg. Argent. 24:27-32.

Martini, Erich C. W.

1914. Some new American mosquitoes. Insecutor Inscitiae Mens. 2:65-76.

1931. Ueber einige siidamerikanische Culiciden. Rev. Entomol. 1:199-219.

Morales, Alberto and H. Vidales

1962. Distribucion de mosquitos selvaticos en San Vicenti de Chucuri, Colombia.

Lozania (Acta Zool. Colomb.) 13:1-16.

Zavortink: Reclassification of Trichoprosopon 47

Morales-Ayala, Francisco

1971. A list of the mosquitoes of Peru (Diptera, Culicidae). Mosq. Syst. Newsl.

3:138-145.

Patino Camargo, Luis

1937. Notas sobre fiebre amarilla en Colombia. Colomb., Univ. Nac., Bogota,

Fac. Med., Rev. 6:211-281.

Porter, John E.

1967. A check list of the mosquitoes of the Greater Antilles and the Bahama and

Virgin Islands. Mosq. News 27:35-41.

Robineau-Desvoidy, Jean-Baptiste

1827. Essai sur la tribu des culicides. Soc. Hist. Natur. Paris, Mem. 3:390-413.

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1970. Mosquito Studies (Diptera, Culicidae). XX. The Terrens Group of Aedes

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Senevet, Georges and E. Abonnenc

1939. Les moustiques de la Guyane Francaise.—III. Les sabethines. Inst. Pasteur

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Shannon, Raymond C. |

1931. The environment and behavior of some Brazilian mosquitoes. Entomol.

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48 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

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Zimsen, Ella

1964. The type material of I. C. Fabricius. Copenh., Munksgaard. 656p.

FIGURES

1. Distribution of Johnbelkinia ulopus

2. Distribution of Johnbelkinia leucopus and J. longipes

3. Johnbelkinia ulopus; male and female heads and claws; male tarsus 5-I; female

thorax, wing, legs, genitalia

4. Johnbelkinia ulopus; male genitalia, pupa

5. Johnbelkinia ulopus; fourth instar larva

6. Johnbelkinia ulopus; third instar larva

7. Johnbelkinia ulopus; male genitalia; second instar larval head, siphon

8. Johnbelkinia longipes; male genitalia, male claws, pupa

9. Johnbelkinia longipes; fourth instar larva

0. Johnbelkinia longipes; male genitalia

Johnbelkinia leucopus; male genitalia, male claws

ye

BELKINIA.

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MIDDLE AMERICA

KILOMETERS

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100 1000

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Fig.4

JOHNBELKINIA

NI 52

Zelaya

Nicaragua

cercal sclerite

NI 56, NIC 116

Zelaya

Nicaragua

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fy

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os

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JOHNBELKINIA

longipes

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French Guiana

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preapical teeth

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JOHNBELKINIA

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Maxilla

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Fig. 10

FORE

og submedian tergal arm

S leucopus

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Bocas del Toro

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Zavortink: Reclassification of Trichoprosopon

CONSPECTUS OF PRELIMINARY RECLASSIFICATION OF

COMPOSITE GENUS TRICHOPROSOPON

Genus Johnbelkinia

culicivora longipes

edwardsiana trichopus

leucopus ulopus

Genus 7richoprosopon

andinum obscurum

brevipes pallidiventer

castroi simile

compressum soaresi

digitatum splendens

evansae subsplendens

lampropus townsendi

lanei trichorryes

mogilasium vonplesseni

nivipes wilsoni

Genus Shannoniana

fluviatilis moralesi

longipalpis schedocyclia

Genus Runchomyia

Subgenus Runchomyia

cerqueirai rapax

frontosa reversa

humbolati theobaldi

Subgenus Ctenogoeldia

dicellaphora magna

homotina walcotti

Subgenus Isostomyia

espini perturbans

lunata shropshirei

paranensis

Genus Wyeomyia

luederwaldti

Nomina Dubia

cotopaxensis hyperleuca

60 Contrib. Amer. Ent. Inst., vol. 17, no. 1, 1979

INDEX TO SCIENTIFIC NAMES

Invalid taxa are in italics; valid taxa treated in detail by present paper are in bold-

face. Major page references are in boldface; page reference to key is shown by suffix

“*k”’; figure numbers are in italics.

Aedes, 28

andinum (Trichoprosopon), 6, 11, 13, 59

argyropus (Maorigoeldia), 8

Binotia, 5

brevipes (Trichoprosopon), 5, 6, 11, 59

castroi (Trichoprosopon), 6, 11, 59

cerqueirai (Runchomyia), 3, 6, 12, 59

compressum (Trichoprosopon), 4, 11, 59

conservator (Culex), 5

cotopaxensis (nomen dubium), 6, 12-13, 59

Ctenogoeldia, 5,6, 12, 15k, 17k, 59

culicivora (Johnbelkinia), 3, 4, 11, 22, 25, 29,

33, 39 :

dicellaphora (Runchomyia), 3, 5, 12, 59

digitatum (Trichoprosopon), 4, 5, 11, 13, 59

edwardsiana (Johnbelkinia), 6, 11, 22, 35, 38,

59

espini (Runchomyia), 3, 5, 12, 59

evansae (Trichoprosopon), 6, 11, 59

fluviatilis (Shannoniana), 3, 4, 6, 11, 59

frontosa (Runchomyia), 3, 4, 12, 59

genurostris (Malaya), 9

Goeldia, 4, 5,6

homotina (Runchomyia), 3, 4, 12, 59

humboldti (Runchomyia), 6, 12, 59

Hyloconops, 4, 5, 6

hyperleuca (nomen dubium), 5, 12, 59

Isogoeldia, 5, 6

Isostomyia, 5, 6, 12, 15k, 17k, 59

Isostomyia of authors, 5

Joblotia, 4, 5

Johnbelkinia, 11, 14, 15k, 16k, 17k, 18-23, 41,

59

lampropus (Trichoprosopon), 3, 5, 11, 59

lanei (Trichoprosopon), 5, 11, 59

Lesticocampa, 4,5

leucopus (Johnbelkinia), 3, 4, 11, 21, 22, 23k,

24k, 27, 28, 30, 37, 39-41, 59; figs. 2, 10

leucopus of authors, 22, 29, 30, 41

Limamyia, 6, 11

Limatus, 12

longipalpis (Shannoniana), 4, 11, 59

longipes (Johnbelkinia), 3, 4,5, 11, 18, 21, 22,

23k, 24k, 24, 25, 27, 28, 29, 35-39, 40, 59;

figs. 2, 8-10

longipes of authors, 21, 22, 29, 30, 38

luederwaldti (Wyeomyia), 5, 12, 59

lunata (Runchomyia), 3, 4, 6, 10, 12, 59

lunata of authors, 4

Lutzia, 14

Lynchiaria, 5

magna (Runchomyia), 4, 10, 12, 13, 14, 29, 59

Malaya, 12

Maorigoeldia, 6

mogilasium (Trichoprosopon), 3, 4, 11, 59

moralesi (Shannoniana), 3, 5, 12, 59

nivipes (Trichoprosopon), 3, 4,5, 11, 59

obscurum (Trichoprosopon), 6, 11, 59

Orthopodomyia, 28

pallidiventer (Trichoprosopon), 3, 4, 6, 11, 13,

59

paranensis (Runchomyia), 5, 12, 59

personata (Wyeomyia), 12

perturbans (Runchomyia), 3, 4,5, 10, 12, 59

perturbans of authors, 5

Phoniomyia, 12

Psorophora, 14

Rachisoura, 11, 14, 21

rapax (Runchomyia), 3, 4, 12, 59

reversa (Runchomyia), 6, 12, 59

Rhunchomyia, 6

Rhynchomyia, 5

Runchomyia, 4, 5, 8, 10, 11, 12, 14, 15k, 16k,

F7K: 27 0

Runchomyia of authors, 6, 11

Sabethes, 12

Sabethini, 2, 6, 10

schedocyclia (Shannoniana), 3, 4, 12, 59

Shannoniana, 6, 7, 8, 9, 10, 11-12, 14, 15k, 16k,

fk ,21; 59

shropshirei (Runchomyia), 3, 5, 12, 59

simile (Trichoprosopon), 6, 11, 59

soaresi (Trichoprosopon), 6, 11, 59

solomonis (Malaya), 8, 10, 22

Zavortink: Reclassification of Trichoprosopon 61

splendens (Trichoprosopon), 4, 11, 59 35, 36, 59

squamipennis (Aedeomyia), 9 trichorryes (Trichoprosopon), 3, 4, 11, 59

subsplendens (Trichoprosopon), 3, 5,11, 59 Tripteroides, 6, 7

theobaldi (Runchomyia), 6, 12, 59 ulopus (Johnbelkinia), 3,4, 9, 11, 12, 13, 21,

Topomyia, 12 22, 23, 23k, 24k, 24-35, 37, 38, 40, 41, 59;

townsendi (Trichoprosopon), 3, 6, 11, 59 figs. 1, 3-7

Toxorhynchites, 14 vonplesseni (Trichoprosopon), 3, 4, 6, 11, 59

Trichoprosopon, 4, 5, 6, 10, 11, 12, 14, 15k, Vonplessenia, 6, 11

16k, 1 7K, 21,59 walcotti (Runchomyia), 6, 12, 59

Trichoprosopon of authors, 2,4,5,6,7,10,11, wilsoni (Trichoprosopon), 3, 5,11, 59

21,59 Wyeomyia, 5, 10, 12, 14, 22, 23, 29, 59

trichopus (Johnbelkinia), 3, 5, 11, 21, 22, 29,

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Contributions

of the

American Entomological Institute

Volume 17, Number 2, 1980

MOSQUITO STUDIES (Diptera, Culicidae)

XXXVI. Subgenera Aedinus, Tinolestes and Anoedioporpa of Culex

By

O.G. W. Berlin and John N. Belkin

CONTENTS

INTRODUCTION

MATERIAL AND METHODS .

KEYS TO SUBGENERA OF CULEX IN THE AMERICAS

SUBGENUS AEDINUS

Introduction :

Taxonomic treatment .

General considerations :

Keys to species

&

2

2.

4.

Culex (Aedinus) amazonensis .

Culex (Aedinus) clastrieri

Culex (Aedinus) guyanensis

Culex (Aedinus) accelerans .

SUBGENUS TINOLESTES .

Introduction :

ae treatment. .

. Culex (Tinolestes) aianie

ee ANOEDIOPORPA

Introduction

Taxonomic treatment .

General considerations.

Keys to species

- Conservator Group .

Culex ( A pehonooe) conservator .

Culex (Anoedioporpa) canaanensis .

Culex (Anoedioporpa) damascenoi .

Culex (Anoedioporpa) browni

Culex (Anoedioporpa) bamborum .

. Culex (Anoedioporpa) belemensis .

. Culex (Anoedioporpa) chaguanco

. Culex (Anoedioporpa) originator

. Culex (Anoedioporpa) quasioriginator

. Culex (Anoedioporpa) luteopleurus

ine

Culex (Anoedioporpa) corrigani .

Restrictor Group .

ee

Culex ( Annetadoria) restrictor

REFERENCES CITED

FIGURES

TABLE OF DISTRIBUTIONS | '

CONSPECTUS OF TAXONOMIC CHANGES

INDEX TO SCIENTIFIC NAMES

cot

ne

403

MOSQUITO STUDIES (Diptera, Culicidae)

XXXVI. SUBGENERA AEDINUS, TINOLESTES AND

ANOEDIOPORPA OF CULEX!

By

O.G.W. Berlin and John N. Belkin?

-_ INTRODUCTION

The subgenera Aedinus Lutz 1904, Tinolestes Coquillett 1906 and Anoedioporpa

Dyar 1923 of Culex, revised in the present paper, were all lumped along with Micra-

edes Coquillett 1906 into the subgenus Aedinus Bourroul 1904 by Stone, Knight and

Starcke (1959:281-282) on the basis of a short palpus in the males. As this feature

had been shown to be of small importance (Edwards 1932:188) and had evolved in-

dependently in several unrelated phylads, it was a taxonomic puzzle until Belkin

(1968b: 11-12, 51) explicitly unraveled it. Belkin’s recommendations were followed

by Knight and Stone (1977) and are accepted here as indicated in the following list.

Aedinus is credited to Lutz in Bourroul 1904 with the type species A. amazonensis

Lutz 1905; it takes precedence over Eubonnea Dyar 1919 and includes in addition to

amazonensis 3 other species. Tinolestes Coquillett 1906 is elevated to subgeneric

rank with latisquama Coquillett 1906 as its type and the only included species.

Anoedioporpa Dyar 1923 is considered as a distinct subgenus of Culex with conserva-

tor Dyar & Knab 1906 as its type (with all its current synonyms) and also includes

bamborum Rozeboom & Komp 1948, belemensis Duret & Damasceno 1955, browni

Komp 1936, canaanensis Lane & Whitman 1943, corrigani Dyar & Knab 1907, origi-

nator Gordon & Evans 1922, luteopleurus Theobald 1903 and 3 recently described

species: chaguanco Casal, Garcia & Fernandez 1968, damascenoi Duret 1969 and

quasioriginator Duret 1972. Culex restrictor Dyar & Knab 1906, considered to be a

member of Microculex by Stone, Knight and Starcke (1959:28) and by Knight and

Stone (1977:269), has been included in the subgenus Anoedioporpa in a separate

group on the basis of correlated features of adults and immature stages. Micraedes

Coquillett 1906 is a distinct subgenus as recognized by Berlin (1970). Culex cauch-

ensis Floch & Abonnenc 1945 and C. mojuensis Duret & Damasceno 1955 have been

transferred to the subgenus Melanoconion, which contains primarily species with a

long palpus in the males.

As stated above, the species with a short palpus in the males are not all closely re-

lated and therefore cannot be assembled into a single genus or even subgenus. Ac-

cording to Edwards (1932:188) a study of the male genitalia and other characters

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

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

Contract DA-49-193-MD-2478.

* Department of Biology, University of California, Los Angeles, CA 90024.

2 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

show not only that the Oriental and Neotropical forms belong to different sections

of the genus, but that even among the Neotropical forms with a short palpus in the

males, there are several groups distinct in origin. In the Neotropics, this character

state has developed independently in several groups of Culex such as Micraedes (Bi-

sulcatus Group), Aedinus, Tinolestes, Anoedioporpa (Conservator Group), Belkino-

myia and a few species of Melanoconion and probably other subgenera. Micraedes

has been studied recently by Berlin (1970) and Belkinomyia by Adames and Galindo

(1973). The present paper is an attempt to study in some detail the 3 principal other

groups of species of Culex with a short palpus in the males: Aedinus, Tinolestes and

Anoedioporpa.

We are indebted to the following individuals: Alan Stone, formerly associated with

U. S. National Museum of Natural History (USNM), for the loan of types and other

material; Thomas H. G. Aitken, Yale Arbovirus Research Unit, New Haven, CT, for

valuable information on amazonensis and accelerans; J. Clastrier, Museum d’ Histoire

Naturelle, Paris, France for the loan of the type of guyanensis; Peter F. Mattingly of

the British Museum of Natural History for studying the type of luteopleurus; William

A. Powder for valuable assistance in the preparation of material; Nobuko Kitamura,

L. Margaret Kowalczyk and Nancy Martsh for the preparation of preliminary and fi-

nal drawings; and Sandra J. Heinemann for valuable general assistance and for the

preparation of the text copy for lithoprinting.

MATERIAL AND METHODS

This study is based largely on extensive collections of adults and associated imma-

ture stages accumulated at the University of California, Los Angeles for the project

‘“‘Mosquitoes of Middle America’ (Belkin, Schick et al. 1965) and also on material

from U. S. National Museum of Natural History (USNM) where most of the types

are deposited. For this revision 4802 specimens were studied, including 739 males,

702 females, 1011 pupae, 2350 larvae, with 777 individual rearings (198 larval, 482

pupal and 97 incomplete).

The method of presentation, terminology and abbreviations used in the descrip-

tions of taxa in general follow Belkin (1962). No attempt was made to locate hairs

(?) 16,17,18,19-C on the head capsule and cervical area of the fourth instar larvae,

although these have been noted by others in some groups (Belkin 1962:87,94,236,

249, figs. 36,140,156 etc.; 1968a:12, fig. 6; Hochman and Reinert 1974). The exact

nature of some of these at least has not been determined as yet.

For every species, all the available stages are illustrated. Each illustration in based

on topotypic specimens whenever available but the modal value of each character,

such as hair branching, is based on 5 to 10 specimens when available. In the illustra-

tions of adults, only pertinent details, such as male genitalia, are given. The type spe-

cies of all the 3 subgenera and Groups are illustrated in detail.

In the distribution lists, the arrangements of the countries, their major subdivisions

and the individual localities are listed alphabetically. All material we have examined

as well as published records that we consider reliable are given under distribution.

Political subdivisions and locality names in the distribution lists conform to the re-

commended usage in the Official Standard Names Gazetteers of the United States

Board on Geographic Names.

The abbreviations used for type depositories as well as depositories of material list-

ed in this paper are as given in “‘A catalog of the mosquitoes of the World” 1977

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 3

except for UCLA (Dept. of Biology, University of California, Los Angeles, CA 9002-

4). All the latter material will eventually be deposited in USNM.

KEYS TO SUBGENERA OF CULEX IN THE AMERICAS

ADULTS

i Acrostichal bristles developed on discof mesonotum . . . ye

Acrostichal bristles not developed except at extreme anterior end pra meso-

notum and/or near prescuteliag Snace. 2 eR

2(1). Usually 6 or more lower mep bristles; all femora ere vein C with al-

ternate brown and yellow scaled areas. . . oe 2 ea

Usually | or 2 lower mep bristles, rarely 3 or more; fear not conspicu-

ously speckled; vein C without alternate brown and yellow scaled areas

ot). Abdomen usually with at least indication of apicolateral or apical pale ter-

gal markings on some segments ... . » 1 . Neoculex

Abdomen without apicolateral or apical transverse pale tergal markings; if

pale markings present, basal or basolateral .

4(3). Pleuron with distinct scale patchesonstp . . - oe Caler

Pleuron at most with a few scales along stp bristles . Ge eee rs,

5(4). Tarsi with basal light scales . . . . . . . . . +. . in part Microculex

Tags, completely Gat, a

6(5). Palpus about 0.35 of proboscis in both sexes, very slender; lower mep with-

out bristles (Bisulcatus Group) . . . . . in part Micraedes

Palpus less than 0.25 of proboscis in fralc: and subequal to proboscis in

WIMIGS 2 ee

7(6). Pleural integument uniformly pale to yellowish (Restrictor Group)

. in part Anoedioporpa

Pleural inteoument mostly pale to yellowish but brown on ppn, lower stp

and lower mep (Schicki and Erethyzonfer Groups) .__ in part Micraedes

8(1). Abdomen with iridescent basolateral tergal markings . . . . . Carrollia

Abdomen completely dark or with noniridescent pale tergal markings .9

9(8). Vertex of head with at least a line of broad decumbent scales along orbital

margin, usually all decumbent scales on vertex broad . .Melanoconion

Vertex scales on head narrow, even along orbital lines. . . . . . . 10

10(9). Propleural bristles (ppl) very strongly developed, at least 20 (20-25); mid-

dle mep with a patch of scales and short bristles: . . . . Tinolestes

Propleural bristles (ppl) poorly developed, fewer than 10, usually about 5;

middle mep without patch of scales or short bristles . . . . .. 11

L1(10). : Veine Re and Ry with btoad geales only 5 2 yk tes 4?

Wee Tk, al he with narrow sCalee) a ee aR

12(11). Decumbent scales on vertex and sides of head whitish; stp with a narrow

patch of translucent scales along bristles. . . . ee Aedinus

Decumbent scales on vertex and sides of head dark; stp at most with a few

ROaIOs TlOMe Dristleg 2 ee ee Bae Belkinomyia

4

13(11).

4(3).

5(4).

6(5).

7(6).

8(7).

9(8).

10(7).

Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Upper stp with distinct patch of scales . . . . . . in part Microculex

Upper stp without patch of scales (Conservator Group) .

es at a tae Wea . in part Anoedioporpa

MALE GENITALIA

Paraproct with a dense clump of sharp spicules in addition to blunt teeth

ee ee ge ore . Culex

Paraproct with only a single row Gf bhint teeth ©

Lobe. on sidepiece poorly developed §. ... 2 sk eee es ei

bote Of sderece irene de COR | 8

Phallosome columnar, blunt and without a basal hook on each side

Neoculex

Phallosome not ‘columnar and with a 1 basal hook ¢ on lateral Bee on each

Se a CG e

Clasper short, stout, about ie 2 ees of ee i Bane of lateral

plate with distinct denticles along itslength .. . . . Belkinomyia

Clasper moderately long, at least 0.5 length of ces basal hook of lat-

eral plate usually without denticles. . ... TS CS Gan a A

Paraproct with 22-24 apical teeth; clasper with an external hyaline crest

at about middle; IX tergite lobe flat, bearing erect bristles on tubercles

: Tinolestes

Paraproct with baer than 14 apical teeth: élasper without an external

crest; [IX tergite lobe variable in shape, bearing weaker setae . . . .6

Lobe of sidepiece submedian in position, undivided, stem bearing usually

FP appengare. : . . . Aedinus

Lobe of sidepiece cubavicat } in Benen, - east faintly divided. Bien of

proximal division Hearne 2 appendaees: . 8. sO ed

Distal division of subapical lobe poorly developed, or faintly ee

from proximal division . . Hes)

Distal division of subapical lobe dicanedy separated aad subedual; in length

tO PIOMINID CIVIEION CO ee Se a ee a)

Apex of clasper usually enlarged, variously shaped ea) or proximal divi-

sion with an accessory lobe; distal division when present usually with |

lanceolate seta . . . Carrollia

Apex of clasper without enlargement: proximal division of lobe without

an accessory lobe; distal division usually bearing more than | seta . .9

Lobes of tergite [X distinct, well separated; interlobar space wide, its

width greater than basal width of one lobe . . . . . . . Microculex

Lobes of tergite IX distinct, closer; interlobar space narrow, its width less

than basal wiedthcol One tobe 26. woo ow care... Miereedes

Lobes of tergite [X enlarged, variously shaped; interlobar space narrow .

Hoe . Melanoconion

Lobes of tergite IX smaller. usually moundike: interlobar space wide.

. Anoedioporpa

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 5

PUPAE

CS Abdominal tergite VIII with caudal lobe overlying lateral part of tergite [X

ne:

Abdominal tergite VIII ‘eith caudal lobe not overlying lateral part of dere

Oe tc ean a ee

201). Paddle margins with Be hairlike spicules (Sehiok and Eo ie

Groups) .. UP Ne ne . . . in part Micraedes

Paddle margins smooth pines ie a hilar

3(2), Hair 5-C extremely long, at least 3.0 of 4-C; hair LIX; absent Sausiiéetus

GsTOUP). Le . . . in part Micraedes

Hair 5-C shorter, sabsqtials in length to 4- C: hair 1-IX present but small .

. Anoedioporpa

4(1). Meatus of trumpet with narrow slit from eee part of pinna

: . Melanoconion

Meatus of trumpet without slit foo ni proximal part CUO ste ee aS

5(4). Paddle hairs (1,2-P) both absent; paddle with a distinct pigmented spot

distally along midrib. . . . . Aedinus

At least 1 paddle hair present; paddle aioe: a distal plemented spot <”..:6

6(5). Hair 9-VIII well removed cephalad from caudolateral angle and always

within 0:7 from base 2). gr, dee

Hair 9-VIII at or immediately adjacent: fo. caudolamiat nite Bib orders Ie Ue

7(6). Hair 2-III laterad of 1-III; hair 9-VII short, distinctly smaller than 3-VII .

. . Belkinomyia

Hair 2- 0 ious of 1-Ill: hair 9-VII longer. Rema loyear than 3-VII .8

8(7). Hair: 2-Vi1 laterag Or TeV ee ra rE es nr ge

Hair 2-V1 mesad of T4Vi7 2 fo rs ee oa a aa ake

9(8). Hair 11-C double; 9-VIII double, shorter than pee of tergite VIII

Tinolestes

Hair l ie single: 9 VIII with more < dan 3 branches, ‘at least subequal to

length of terete Vie. oe es, pee tl. Migrecuiex

10(8). Only 1 paddle hair (1-P) present; naka 2-VII mesad Va 1-VII . . Carrollia

Both gas hairs (1 bis distinct; 2-VII eas or laterad in position

ie

11(10). Metanotal hair 10-C cade aches mn with 3,4 i aches i 5 he

Metanotal hair 10-C usually with fewer than 4 branches . . . Neoculex

FOURTH INSTAR LARVAE

1, Labrum produced in front; mouthbrushes thickened, inserted in compact

groups laterally .. Mon ranger ane By 8 7)

Labrum not produced in front: ‘mouthbrushes normal: UG ee MN Gist a Lee

2(1). Hair 2-C strongly developed, always mesad of 1-C; comb scales usually in a

single row, if in a small patch, then ae with long subventral tufts

within pecten’’. . . . . Carrollia

Hair 2-C weakly developed, indistinct c or absent wher present laterad of

[4c subventra: tults Usually Not Within pecten 20

6(5).

7(6).

&(7).

9(8).

10(9).

Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Ventral brush (4-K) with4 pairsofhairs. .. .. . . . . Microculex

Ventral Drish (44X) with 5 Gr morc pats Ol haits 2.

Caudolateral border of anal saddle with long slender spines. . Micraedes

Caudolateral border of anal saddle without spines or with only small in-

Conspicuous pied

Prothoracic hair 3-P of the same order of magnitude and thickness as hair

1-P, usually at least 0.67 aslong . . Culex

Baie hair 3-P much thinner and ote that hak L-P, asually less

than 0.5 of its length Pe ee

Ventral brush with 1 or more hairs proximad of ae siphon without any

subdorsal tufts distinctly above subventral tufts. . . . . Neoculex

Ventral brush without any detached hairs proximad of grid; siphon with 1

or more pairs of subdorsal tufts distinctly above subventral tufts . .7

Siphonal hair 2-S very strongly developed, recurved and usually with a re-

curved tooth on shaft; subdorsal tufts on siphon conspicuous .

. Melanoconion

Siphonal hair 2- S ee poorly developed but of the same type;

subdorsal tufts on siphon less conspicuous .

Saddle hair 2-X long, single . . . . .Anoedioporpa

Saddle hair 2-X with at least 1 Additiodal subbasa beach : ‘ 9

Saddle with an attached acus ventrad; hairs 1,2-VIII on sclerotized plates;

ocular bulge prominent .. . , 4 4. Aedinus

Saddle without any indication of an acus; hairs 1 a VIII on simple alveoli;

ocular buive not prominent. ... ... te

Head capsule with imbrications visible at 200X; hair 2-X mith 3 additional

short to moderately long subbasal branches. . . . Tinolestes

Head capsule smooth at 200X; hair 2-X with only 1 additional short sub-

basal bfaica a a. Belkinomyia

SUBGENUS AEDINUS Lutz 1904

INTRODUCTION

Aedinus was proposed as a distinct genus in 1904 but its authorship was a very in-

volved problem until Belkin (1968b:48-51) solved it. According to Belkin, Lutz was

the author of all the nominal taxa in Bourroul’s thesis except mariae, and therefore

he attributed the generic taxon Aedinus to Lutz in Bourroul 1904 (Lutz 1904a: 12;

1904b:4) and eliminated Aedinus Bourroul 1904 of Stone, Knight and Starcke

(1959-281).

As for the type species, Stone, Knight and Starcke gave for their Aedinus Bourroul

1904 the haplotype Aedeomyia americana Neveu-Lemaire which was the only species

referred to Aedinus in the paper where this genus was described (Lutz 1904b:4). But

in “‘Catalogo”’ (Lutz 1904a:12) in Bourroul’s thesis, americana was questionably re-

ferred to Aedinus and only A. amazonensis Lutz (n. e.) was included in Aedinus Lutz

(n. gen.). According to Belkin (1968b:51): “If the designation of americana, which

is anomen dubium, as type species is accepted, Aedinus also becomes anomen dub-

ium. In my opinion, a better solution is to consider Aedinus without included spe-

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 7

cies in 1904 since both amazonensis and nigricorpus were nomina nuda in 1904 (nei-

ther can be regarded as described by indication by a new generic description since 2

names are involved) and americana is referred to it questionably, and to designate

Aedinus amazonensis Lutz, 1905 as the type species as it was the only included spe-

cies in the first subsequent publication of the genus. This restores Aedinus Lutz,

1904 as a valid subgenus in Culex in the sense of Lutz, 1904 and 1905, and as it has

been used before Stone, Knight and Starcke (1959), replacing its subjective junior

synonym Eubonnea Dyar, 1919.”

In the present study, Aedinus Lutz 1904 is treated as a distinct subgenus of Culex

with amazonensis Lutz 1905 as its type species. In addition, accelerans Root 1927

and 2 recently described species, clastrieri Casal and Garcia 1968 and guyanensis

Clastrier 1970, are included in this subgenus.

TAXONOMIC TREATMENT

Figs. 1-7

1904. Aedinus Lutz 1904a:12. TYPE SPECIES: Aedinus amazonensis Lutz 1905:103-104, Ma-

naus, Amazonas, Brazil. Determination by Belkin 1968b:51.

1919. Eubonnea Dyar 1919:150. *TYPE SPECIES: Culex (Eubonnea) tapena Dyar 1919:150,

Paramaribo, Suriname; only included species. Synonymy with Aedinus by Belkin (1968b:

ty).

Culex (Aedinus) of Dyar (1923b:187,190); Bonne and Bonne-Wepster (1925:279); Edwards (1932:

220); Anduze (1941b:17); Lane (1949:255; 1953:385); Foote (1954:4); Horsfall (1955:547);

Galindo and Blanton (1955:68,69-70); Fauran (1961:43); Belkin (1968b:11,51); Stone (1970:

164); Knight and Stone (1977:194-195, in part).

Culex (Eubonnea) of Stone, Knight and Starcke (1959:280); Stone (1961:47); Belkin (1962:179);

Stone (1963:135); Forattini (1965:31,32,34,35,187,192); Cova Garcia, Sutil and Rausseo

(1966a:27; 1966b:352); Aitken, Spence et al. (1969:213).

Aedinus of Lutz (1904b:4; 1905: 103-104); Blanchard (1905:620,633); Peryassu (1908:36,51,

253); Theobald (1910:487); Surcouf and Gonzalez-Rincones (1911:219,223).

FEMALES (fig. 2). Usually small inornate species, dark brown to blackish and

with unbanded legs. Head: Eyes not distinctly separated above antennae. Decum-

bent scales narrow and linear dorsally, predominantly white; broader on sides and

venter. Erect scales on occiput moderately long, forked apically, extending to sides

of vertex. Orbital and interorbital bristles strong; upper orbitals 5 (5,6) pairs, heavier,

longer and more widely spaced than lower. Clypeus bare, dark brown. Proboscis dis-

tinctly longer than forefemur, entirely dark scaled, with a few basal bristles. Palpus

short, about 0.2 of proboscis length; 4-segmented; segments 1 and 2 ankylosed, with-

out scales; segment 4 about 2.0 of segment 3, both dark scaled. Antenna slightly

longer than proboscis; torus brown, with a few short, dark brown setae mesally; fla-

gellar segments 2-13 with 6 moderately long bristles in basal whorls. Thorax: Integu-

ment dark brown. Mesonotum with narrow, curved, auburn to dark scales except

along a pair of narrow inner dorsocentral “‘bare lines’ extending from anterior mar-

gin to 0.75 of its length. Acrostichal bristles absent. Bristles on anterior promontory

distinct; dorsocentrals, prescutellars and supraalars always present, variously devel-

oped; 2,3 posterior fossal and 1 parascutellar always developed. Antealar area above

paratergite with scattered dark scales. Median scutellar lobe with 6 long and 6 short

marginal bristles and a large patch of dark narrow scales; lateral lobe with about 4

long and 3 short marginal bristles and a small patch of dark narrow scales. Parater-

gite bare. Pleuron dark brown to blackish. Bristles present on apn, ppn, ppl, stp, pra

8 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

and upper mep; propleural area with about 10-12 short bristles; lower mep with sin-

gle, strong bristle; metameron bare. Pleural scaling restricted to ppn and stp; upper

ppn with a few dark scales; stp with 2,3 rows of flat, translucent, moderately broad

scales along bristles. Legs: Dark scaled. Claws simple on all legs. Wing: Veins en-

tirely dark scaled; all scales squamous. Haltere: Stem brown, knob entirely scaled.

Abdomen: Laterotergite with many moderately long bristles.and a few scattered

scales. Tergites II-VII predominantly with dark brown scales. Sternites with dull

white or dark scales.

FEMALE GENITALIA (fig. 2). Only amazonensis, the type species, studied. Seg-

ment VIII partially retracted into segment VII, apex visible, numerous bristles and

scales present; tergite VIII about 0.5 of sternite, both with shallow emargination api-

cally. Tergite IX well developed, lobes distinct, bearing about 9 short setae; sternite

IX narrow laterally, membranous in the middle. Postgenital plate about 0.7-0.8

length of cercus, slightly emarginate apically; each side with 2 apical and 9,10 scat-

tered setae. Cowl moderately developed, irregularly shaped, bowed laterally to join

tergite [X. Sigma membranous, finely spiculose, continued medially as a membran-

ous insula with a group of 9,10 setae. Cerci moderately long, compressed, approxi-

mate, each with many short to moderately long setae; area between cerci slightly bi-

lobed. Spermathecae 3, one a little larger than others.

MALES (fig. 2). Coloration similar to females; sexual dimorphism of head append-

ages marked. Proboscis longer than forefemur. Palpus porrect, about 0.2 of probos-

cis length; apparently 4-segmented; segments 1 and 2 ankylosed, without scales; seg-

ment 4 subequal in length to 3, partly ankylosed, dark scaled. Antenna subequal in

length to proboscis; whorls of flagellar segments 1-12 strongly developed, with at

least 24 long bristles; segments 12 and 13 elongate, 13 slightly longer than 12; torus

not swollen. Claws of foreleg and midleg enlarged, unequal; larger claw on foreleg

with a submedian tooth, that of midleg without tooth, both claws with basal spicules.

Hind claws as in females.

MALE GENITALIA (figs. 3,5,6). Segment IX: Lobes of tergite [IX moderately

long, distinctly separated, digitiform, bearing a few very short, weak, apical setae.

Sidepiece: Roughly oval; length about 2.0 greatest width; with longer bristles on ter-

gal and lateral sides, sternal surface with shorter setae; scales on lateral and/or sternal

surfaces. Lobe submedian in position, bearing 1 or 2 appendages. Clasper: Compar-

atively simple or enlarged subapically; spiniform subapical. Phallosome: Lateral

plate of aedeagus with a broadly sclerotized “‘basal hook,” a caudally directed blunt

apical process and 2 short sternal processes. Proctiger: Basolateral sclerotization

moderately developed, produced into a conical lobe below tergite IX; apex of para-

proct with a crown of 5-9 teeth; cercal setae indistinct.

PUPAE (figs. 3,6). Cephalothorax: Middorsal ridge moderate. Integument with a

characteristic pattern of pigmentation. All hairs present, variously developed. Hair

5-C moderately long, weakly developed, subequal in length to 4-C; hair 6-C smaller

than and cephalad of 7-C; hairs 8,9-C widely separated, double or triple, 8-C slightly

and 9-C distinctly caudad of trumpet base. Trumpet: Not placed on tubercle; mod-

erately long, index 7.0-8.5; pigmentation strong; tracheoid distinct, moderately long;

apex distinctly flared; pinna moderately long. Metanotum: Hair 10-C shorter than

11-C, multiple, weakly branched. Abdomen: Integument with a distinct pattern of

pigmentation. Hair 3-I single; 1-II with at least 15 weak branches, faintly resembling

float hair (1-I); hair 2-II] distinctly mesad of 1-III, submarginal; hair 5-IV-VI moder-

ately long, branched, terminating before the apex of succeeding segment; 6-II-VI dou-

ble; 2-VI,VII always laterad of hair 1; hair 9-VII, VIII single, short, distinctly shorter

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 9

than corresponding tergite length; 4-VIII at least double; 1-[X small, single. Lobes

on posterior margin of sternum mesad of 9-VIII indistinct. Tergite VIII never over-

lapping base of tergite [X. Paddle: Lightly pigmented except for a distal pigmented

patch along midrib; longer than wide, apex smoothly rounded; midrib strongly dif-

ferentiated; paddle margin without spicules; both paddle hairs (1,2-P) absent.

FOURTH INSTAR LARVAE (figs. 4,7). Head: Head capsule faintly imbricate,

without lateral expansion on each side caudad of antenna. Labrum well differenti-

ated dorsally. Ocular lobes distinct. Mouthbrushes with numerous filaments. Collar

moderately develped, narrow. Posterior tentorial pit a short distance from caudal

border. Maxillary suture complete, extending a short distance caudolaterad of pit.

Anterior border of labial plate truncate. Aulaeum with distinct filamentous spicules.

Central tooth of mental plate not shouldered. Hair 0-C small, removed laterad of 1-

C; hair 1-C strong, straight; 2,3-C not developed; 4-6-C closely grouped together and

caudad of level of 7-C; hair 5-C single or double, 6-C single, both shorter than anten-

na; 7-C multiple, with strong, barbed branches; 8-C at least triple (3-5); hair 11-C

with 3-7 weak branches; 13-C closer to 11-C than to 12-C; hairs 14,15-C not moved

anteriad; basal maxillary hair (bmh) small, single; 16,17-C not developed. Antenna:

Hair 1-A long, multiple, about 0.60-0.65 from base; other hairs short to moderately

long, single. Thorax: Roughly oval in outline, wider than long. Integument gla-

brous. Hairs 1-3-P on a distinct tubercle; 3-P moderately long, single or double; 14-P

single or double; O-P, 1,13,14-M, 1,3,13-T with many weak branches, not stellate.

Abdomen: Integument glabrous; segments darkly pigmented except IV. Hair 2 nev-

er moved far cephalad of 1, always within posterior half of segment. Segment VIII:

Comb in a patch of 3,4 irregular rows. Hair 1-VIII on a small tubercle and 2-VIII on

an elongated sclerotized plate. Siphon: Integument glabrous. Acus long, attached.

Pecten with denticles on ventral border; 5,6 pairs of subventral (1,1a-S) and 2 pairs

of subdorsal (2a-S) hairs; proximal subdorsal hair distinctly basad of proximal sub-

ventral hair; 2-S distinct, slightly curved with a subbasal branch. Anal Segment: Sad-

dle complete; integument of saddle glabrous; acus present, small, attached; caudal

margin without spines; lateral hair (1-X) short to very short, multiple, submarginal.

Hair 2-X long, with 2 subbasal branches; 3-X long, single; ventral brush (4-X) with 6

pairs of branched hairs on grid. Gills sausage-shaped, ventral slightly longer than dorsal.

GENERAL CONSIDERATIONS

SYSTEMATICS. Adults of Aedinus can be separated from those of other Ameri-

can subgenera of Culex by the combination of the following characters: (1) short

palpus in both sexes, (2) presence of moderately broad scales on R, and R3 as on

other veins, and (3) absence of acrostichal bristles. In the male genitalia, Aedinus

can be distinguished from other subgenera by the combination of: (1) shape of the

IX tergite lobe which is digitiform and bears short weak setae, (2) nature and posi-

tion of subapical lobe of sidepiece, (3) shape of clasper, and (4) details of phallosome

complex. In the pupal stage it differs from others by the combination of: (1) distinct

pattern of pigmentation on cephalothorax and abdomen, (2) moderately long trum-

pet with an apical flare, (3) nature and branching of hair 9-VII,VIII, and (4) pigmen-

ted spot on the paddle. In the larval stage, Aedinus can be distinguished from other

subgenera by the combination of: (1) width of head greater than length, (2) faint im-

brications on head capsule, (3) distinct ocular bulge, (4) 6 pairs of hairs in ventral

brush, (5) hair 2-X with 1 or 2 additional subbasal branches, (6) hairs 1,2-VIII on dis-

tinct sclerotized plates, and (7) attached acus on anal saddle. The last character dis-

10 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

tinctly separates Aedinus from all other subgenera of Culex.

All recognized species of Aedinus are difficult or nearly impossible to differentiate

as preserved adults. The description and recognition of all species have been primari-

ly based on male genitalia which provide excellent diagnostic features. The imma-

ture stages of clastrieri and guyanensis are unknown, but when found, will probably

be more similar to amazonensis than to accelerans. In the immature stages, amazo-

nensis displays clearcut differences from accelerans; therefore, characters in the lar-

val and pupal stages may also help in differentiating the various species within the

subgenus.

BIONOMICS AND MEDICAL IMPORTANCE. Species of Aedinus breed primari-

ly in swamp margins and root caves in swamp interiors in virgin forests and occasion-

ally in ground pools in partially forested areas. Though blooded females have been

collected, it is not known whether females are attracted to man. Arboviral studies

on amazonensis and accelerans show that they harbor a spectrum of viruses (Aitken,

Spence et al. 1969:212-213). While females of accelerans inoculated into mice yield-

ed 3 viruses (VEE, Caraparu and Nepuyo), those of amazonensis have been found to

be infected with 7 different arboviruses (VEE, Caraparu, Catu, Guama, Wyeomyia,

Bushbush and Aruac). It appears, therefore, that they are involved in the transmis-

sion of these viruses in the different animals in the forests. Additional information

can be found in the bionomics sections of the individual species accounts.

DISTRIBUTION (fig. 1). The subgenus Aedinus is distributed primarily in north-

ern South America and penetrates into Panama. Apparently all species are restricted

to swampy coastal areas and penetrate deep into the mainland only along the mouth

of the Amazon River. Of the 4 described species of the subgenus, amazonensis and

accelerans appear to have a wider distribution in terms of range and number of indi-

viduals. On the other hand, clastrieri and guyanensis have a restricted distribution.

The subgenus is confined to the mainland of the neotropics. The northern limit

appears to be Panama, and no species of Aedinus has been reported north of this

area; the southern limit is Porto das Caixas, Rio de Janeiro, Brazil, the type locality

of accelerans.

AFFINITIES. Aedinus is a distinct subgenus of Culex and several subgenera have

been erroneously synonymized with it (Stone, Knight and Starcke 1959:281).

Though adults of Aedinus show resemblance to those of Tinolestes and Belkinomyia,

the subgenus is distinct in the male genitalia, immature stages and biology (breeding

sites). The following combination of characters in the male genitalia and immature

stages of Aedinus may be diagnostic for the subgenus; in the male genitalia by: (1)

position and details of the lobe of sidepiece, (2) details of clasper and phallosome

complex, and (3) shape of IX tergite lobes; in the pupae by: (1) characteristic integu-

mental pigmentation, (2) nature and branching of hair 9-VII, VIII, (3) pigmented

spot in the paddle, and (4) absence of both paddle hairs (1,2-P); in the larvae by: (1)

faint imbrications on head capsule, (2) hairs 1,2-VIII on distinct sclerotized plates,

(3) distinct attached acus on saddle, and (4) 6 pairs of hairs on ventral brush (4-X).

In spite of some slight overlap with Tinolestes and Belkinomyia, Aedinus is a distinct

group and should be retained as a subgenus of Culex.

2(1).

AY),

3(2).

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 11

KEYS TO SPECIES OF AEDINUS

ADULTS

Abdominal sternites white scaled. . ...°.°.. ¢°. 3...” 1. amazonensis

Abdomina! stermiites dark scaled. 2 2 a 8

Picuron pale browi - a ee 2 clase

CT eo . 3. guyanensis

Pleuron dark, with a smoky bluish gray iridescence. . . . . 4. accelerans

MALE GENITALIA

Clasper simple, not enlarged subapically; — on subapical lobe of

sidepiece distinctly longer than stem . . . . 4. accelerans

Clasper distinctly enlarged subapically; appendage on subapical lobe of side-

piece shorter than siem- 4. ee Py ei ee

Expanded portion of clasper sharply attenuating distally; dorsal surface of

sidepiece laterad of subapical lobe with a few bristles . . J. amazonensis

Expanded portion of clasper gradually tapering distally; dorsal surface of

sidepiece laterad of subapical lobe with a dense patch of bristles . . .3

Stem of subapical lobe of sidepiece with 2 setae, subapical and subbasal;

appendage on subapical lobe moderately long, pointed, about 0.7 of stem

. 2. Clastrieri

Stem of subapical lobe of sidepiece with only mall subasical scta: append-

age on subapical lobe short, blunt apically, less than 0.5 of stem

. 3. guyanensis

PUPAE

(2. clastrieri and 3. guyanensis unknown)

Pinna moderately long, about 0.35 length of trumpet; tergite VIII elon

lightly pigmented; metanotal pigmentation uniformly light .

. . IL, amazonensis

Pinna shorter, less. than 0. 25 length of trumpet: tergite VIII with distinct

median oie lateral pigmented areas extending entire length of tergite;

metanotum pigmented at middie. . ...°:-... .°...... 2, aceelerans

LARVAE

(2. clastrieri and 3. guyanensis unknown)

Hair 5-C double; 2-X usually with only | additional, short subbasal branch;

siphon with 5 Pairs of subventral hairs. 4... . . I. amazonensis

Hair 5-C single; 2-X with 2 additional subbasal ranches: siphon with 6 pairs

OUSUDVENITA Tate ob ee eee? ageelerans

es Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

1. Culex (Aedinus) amazonensis (Lutz)

Figs. 1,2,3

1905. Aedinus amazonensis Lutz 1905:103-104. TYPE: Syntypes male, female, Manaus, Ama-

zonas, Brazil, date and collector not specified [LU].

1919. Culex (Eubonnea) tapena Dyar 1919:150. *TYPE: Holotype male, Paramaribo, Suriname,

5 Jan 1919, J. Bonne-Wepster [USNM, 11623]. Synonymy with amazonensis by Dyar

(1923b:189).

1922. Culex (Carrollia) paraplesia Dyar 1922b:192-193. *TYPE: Holotype male, Puerto Nino,

Colombia, 21 Feb 1922, F.A. Miller [USNM, 25762]. Synonymy with amazonensis by

Dyar (1923b:189).

1923. Culex hildebrandi Evans 1923b:377-380. *TYPE: Holotype male, Manaus, Amazonas,

Brazil, 1922, A.A. Clark [BM]. Synonymy with amazonensis by Dyar (1923b:189).

Culex (Aedinus) amazonensis of Howard, Dyar and Knab (1915:217); Dyar (1923b:190; 1924:

183); Bonne and Bonne-Wepster (1925:184,279-281); Edwards (1932:220); Lane (1939:81;

1953:378,385-386); Anduze (1941b:17; 1947:358); Floch and Fauran (1954:2-3); Galindo and

Blanton (1955:70); Castro, Garcia and Bressanello (1959:552); Cerqueira (1961:31); Fauran

(1961b:43); Belkin (1968b:11,15,51); Belkin, Schick and Heinemann (1971:27); Knight and

Stone (1977:195); Heinemann and Belkin (1978a:183; 1978b:393 407, 437,456; 1978c:523;

1979:80,94,107).

Culex (Eubonnea) amazonensis of Stone, Knight and Starcke (1959:280-281); Stone (1961:47;

1963:135); Forattini (1965:26,35,187); Cova Garcia, Sutil and Rausseo (1966a:27; 1966b:101,

352); Aitken, Spence et al. (1969:210,212-213); Xavier and Mattos (1975:245, as Eubonnea).

Culex amazonensis of Fauran (1961a:8-11).

Aedinus amazonensis of Blanchard (1905:633); Peryassu(1908:51,253-2 54); Theobald (1910:487);

Surcouf and Gonzalez-Rincones (1911:223-224).

Culex (Eubonnea) tapena of Bonne-Wepster and Bonne (1923:126); Stone and Knight (1957:59).

FEMALE (fig. 2). Wing: 2.25 mm. Proboscis: 1.4mm. Forefemur: 1.1 mm. Ab-

domen: 1.6-1.7 mm. As described for the subgenus with the following additional

features. Head: Decumbent scales on vertex narrow, white, broader on sides and

venter. Erect scales dark brown. Palpus short, dark scaled. Thorax: Pleuron light

brown to gray. Propleural areas with 9-12 short bristles. Legs: Coxae with brown

scales on external surface; trochanters with ventral brown scales. Forefemur and

midfemur with auburn scales anteriorly, scales on posterior side creamy basally, rest

auburn; scales on hindfemur mostly creamy ventrally, auburn dorsally. Tibiae and

tarsi of all legs with auburn scales. Wing: Scales on all veins dark, squamous and

moderately broad. Haltere: Stem pale brown, knob with auburn scales. Abdomen:

Tergites auburn, scales on lateral areas paler; sternites with whitish to creamy scales.

MALE (fig. 2). Wing: 2.4 mm. Proboscis: 1.7 mm. Forefemur: 1.25-1.30 mm.

Similar to female in general features. Palpus short, about 0.2 length of proboscis.

MALE GENITALIA (fig. 3). Diagnostic characters as in the key. Segment IX:

Tergite lobes moundlike, distinctly separated, each bearing a subbasal digitiform ap-

pendage with 6-10 short weak apical setae. Sidepiece: Ovate, length about 2.0 of

greatest width; lateral surface with longer bristles and sternal side with shorter setae;

scales present on basolateral and sternal surfaces; 4-6 broad saberlike setae and about

5,6 rounded bristles distad and 5,6 rounded setae laterad of lobe. Lobe: Submedian

in position, stem distinct, undivided; apex bearing a short pointed appendage; a short

subbasal seta on stem. Clasper: Long, about 0.7 length of sidepiece, expanded sub-

apically, sharply attenuating distally; spiniform short, simple. Lateral Plate: Basal

hook broadly sclerotized and strongly curved; apical process broadly rounded and

denticulate; sternal part split into 2 short processes. Proctiger: Cercal sclerite

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 13

short, triangular; cercal setae absent. Paraproct with 8,9 blunt apical teeth.

PUPA (fig. 3). Abdomen: 2.4-2.5 mm. Trumpet: 0.55-0.60 mm; index 8.0-8.5.

Paddle: 0.65-0.70 mm. As described for the subgenus with the following additional

features. Diagnostic characters as in the key. Cephalothorax: Integument with the

characteristic pattern of pigmentation; middorsal ridge area, lateral and posterior

parts of wing case, and middle portions of leg cases moderately pigmented, rest light-

er. Hair 4C usually double; 5-C usually 4-branched (2-6) and subequal in length to

4-C; hair 7-C usually triple (2-4); hairs 8,9-C double. Trumpet: Moderately pigment-

ed, apex distinctly flared; pinna moderately long, about 0.30-0.35 length of trumpet;

tracheoid distinct, strongly pigmented. Metanotum: Uniformly moderately pig-

mented. Hair 10-C multiple (5-9), weakly branched; 1 1-C single; 12-C usually triple

(2-4). Abdomen: Except for tergite V, uniformly lightly pigmented, progressively

lighter caudad; tergite V moderately pigmented and distinctly contrasting with other

tergites. Hair 1-IJ multiple (22-31), weakly branched, faintly resembling float hair

(1-I); hair 1-II-VI multiple, usually with 3-5 branches; 2-III mesad and 2-IV-VII later-

ad of hair 1; hair 9-VII,VIII single; 4-VIII usually triple. Paddle: Lightly pigmented;

pigmented spot apically at level of midrib; apex smoothly rounded. Male genital

lobe extending to 0.4 and female genital lobe to 0.2 of paddle.

FOURTH INSTAR LARVA (fig. 4). Head: 0.6 mm. Siphon: about 1.5 mm.

Anal Saddle: 0.3 mm. As described for the subgenus with the following additional

features. Diagnostic characters as in the key. Head: Width about 1.5-1.6 of length.

Mental plate well developed, with 7,8 lateral teeth. Hair 4-C usually double (2-4),

anteriad of 5-C; hair 5-C double, barbed; 7-C with 5-7 strong barbed branches, ex-

tending to base of antenna; 1 1-C with 3-7 weak branches. Antenna: Integument yel-

lowish; spiculate to base of 1-A; hair 1-A with 22-29 barbed branches. Thorax: Inte-

gument glabrous. Hairs unbranched or with branches of varied length, not stellate.

Abdomen: Integument glabrous. Hairs 1,2,4,11,13-I, 1,2,5,9,13-II, 1,2,5,7-9, 13-III-

VI, 1,2,5,8,10,13-VII unbranched, or with branches of varied length and thickness,

never stellate; 6-I,II long, double, branches subequal; 7-I long, single; 7-II short, mul-

tiple; 6-IJI-V moderately long, at least triple, branches subequal; 6-VI shorter, usually

triple (2,3), branches subequal. Segment VIII: Comb scales spatulate, 30-39 in num-

ber, apex fringed. Hari 2-VIII double; 5-VIII usually 4-branched. Siphon: Integu-

ment yellowish, with a narrow basal dark ring; long, index about 14-15. Pecten with

17 (16-18) teeth, extending to basal 0.25; each tooth with 7,8 denticles on ventral

border; only 5 pairs of subventral (1,1a-S) hairs present. Anal Segment: Integument

yellowish. Hair 1-X short, weakly branched; 2-X long, usually with 1, rarely 2, short

to moderately long subbasal branches. Gills about length of saddle, sausage-shaped,

ventral slightly longer than dorsal.

SYSTEMATICS. Culex amazonensis can be distinguished from the remaining

known members of Aedinus by the following combination of attributes; in the male

genitalia: (1) expanded portion of clasper sharply attenuating distally and (2) dorsal

surface of sidepiece laterad of lobe with only a few bristles; in the pupa: (1) moder-

ately long pinna of the trumpet and (2) tergite VIII uniformly lightly pigmented;

and in the larva: (1) hair 5-C double, (2) 2-X with only 1 additional, short, subbasal

branch, and (3) 5 pairs of subventral hairs (1,1a-S) on siphon.

We have seen adults of amazonensis from Panama, Colombia, Ecuador, Venezuela,

Guyana, Suriname, French Guiana and Brazil. The immature stages are known only

from Trinidad, Brazil, Suriname and Panama. There is no appreciable variation in

the male genitalia of these different populations. We have made no attempt to check

carefully the identity of all the other current synonyms of amazonensis listed in

14 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Stone, Knight and Starcke (1959:280-281) and above, but from their descriptions

and limited material at hand, tapena Dyar 1919, paraplesia Dyar 1922 and hilde-

brandi Evans 1923 appear to be conspecific with amazonensis.

BIONOMICS. The immature stages of amazonensis have been found usually in

swamp margins and root caves in the swamp interior, and occasionally in ground

pools, at low elevations in forested areas. Females are apparently zoophilic, but it is

not known whether they are attracted to man. According to Galindo and Blanton

(1955:70), this species is the commonest Culex of the Pacific coastal swamps of Pan-

ama. The males exhibit a very strong positive phototropism and are picked up in

large numbers in light traps set near the breeding places. Studies on feeding patterns

in Panama show that amazonensis prefers mammals, with 87.9% of the blood meal

from this group. Most specimens (about 65%) fed on cricetid rodents, probably

hamsters which were used as sentinel animals, or cotton rats which were common in

the Tocumen swamps along the Pacific coast of Panama (Tempelis and Galindo

1975:205-209).

Culex amazonensis appears to be an important species in harboring a spectrum of

arboviruses. Studies by Aitken, Spence et al. (1969:207-215) in Trinidad show that

during a 10-year survey of arthropods for natural viral infections, 7 different viruses

(VEE, Caraparu, Catu, Guamo, Wyeomyia, Bushbush and Aruac) have been isolated

from amazonensis. While no viruses were detected during 1953-58 surveys, 7 viruses

were isolated in 1959-63 when inoculated into mice. This shows that this species is

potentially becoming important in harboring viruses, but it is not known whether it

can transmit the viruses in nature by the bites of its females.

DISTRIBUTION (fig. 1). Culex amazonensis is known from the Pacific and A tlan-

tic coasts of Panama, from Colombia, Ecuador, Venezuela, Guayan, Suriname,

French Guiana, and Atlantic lowlands and coastal areas of Brazil.

Material Examined: 404 specimens; 81 males, 104 females, 67 pupae, 152 larvae;

54 individual rearings (19 larval, 23 pupal, 12 incomplete).

BRAZIL. Para: Belem, 2M, det. A. Toda [FH; E-822, E-232]; Bosque Rodrigues Alves, 11 Apr

1941, W.H.W. Komp, 3M (KO 29-8) [UCLA] ; IPEAN, Catu Forest, 20 m, 25 July 1969, B.T. and

B.G. Aitken, 1 pM (BRA 6-14), 1 L (6), 1 lpF (BRA 7-13), 1 lp (7-11), 1 L(7) [UCLA] ; IPEAN,

Reserva de Aura, 1-10 m, 7 Aug 1969, B.T. and B.G. Aitken 1 lpM (BRA 22-14), 1 lpF (22-18), 2

pF (22-25, -44), 4 IP (22-24,-27,-38,-40), 8 L (22), 1 P (BRA 24); IPEAN, Reserva de Aura, 1-10m,

8 Aug 1969, B.T. and B.G. Aitken, 2 pF (BRA 28-11,-13), 1L (28) [UCLA] ; Utinga (Xavier and

Mattos (1975:247), 1M, det. A. Toda [FH; 15964]. Sao Paulo: Juquia, det. P.C.Antunes, Coutin-

ho and J. Lane [FH; 1019, 1020, 1022, 3019, 5930, 5931, 5932].

COLOMBIA. Antioquia: Murindo, 1924, L.H.Dunn, 2M [USNM]. Boyaca: Puerto Boyaca,

Calderon, 170 m, 25 Nov 1970, C.J. Marinkelle, 1M (COM 496) with genitalia slide (701229-11)

[UCLA]. Caqueta: Tres Esquinas, Los Alicangaros, 190 m, 8 Aug 1970, C.J. Marinkelle, 12F

(COM 494) [UCLA]. Cordoba: Monteria, Turipana, 10 m, 5-6 Oct 1969, W.A. Page, 5M, 9F

(COL 422) [UCLA]. Meta: Puerto Porfia, 160 m, 9 Sept 1971, C.J. Marinkelle, 1F (COM 595)

[UCLA]. Villavicencio, Finca Porvenir, 450 m, 22 July 1971, C.J. Marinkelle, 1M (COM 530)

[UCLA]. Santander: Lebrija, 900 m, 10 Apr 1966, C.J. Marinkelle, 4F (COM 64A); 12 Apr 1966,

C.J. Marinkelle, 9F (COM 64C) [UCLA]. Locality unspecified: F.A. Miller, 1F [USNM].

ECUADOR. Los Rios: Babahoyo, Levi-Castillo, 1M, 5F [USNM]|. Napo: Coca, Isla Pompeya,

250 m, 19-26 May 1965, L.E. Pena G., 10F (ECU 19) [UCLA]. Cuyabeno, Tarapoa, 300 m, 25

May 1974, D.J. Pletsch, IMgen (ECU 228/74-307) [UCLA]. Locality unspecified: May 1974, D.

J, Pletsch, 1. (ECU 215) [UCLA].

FRENCH GUIANA. Guyane: Cayenne, Mont Cabassou, 30 m, 31 Jan 1965, T.H.G. Aitken, R.

Martinez and A. Guerra, 1M, 1F (FG 15); foret de Cabassou, 4 Sept 1967, J. Clastrier, 1M (FGC

3247-19); foret de Cabassou, 10 Jan 1968, E.N. DeFreitas, 2M (FGC 3299-45,-76,-97,-121), 3Mgen

(3299-166,-181,-194); foret de Cabassou, 21 Feb 1968, J. Clastrier, 1M (FGC 3305-3); foret de

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 15

Cabassou, 21 Aug 1968, J. Clastrier, 1M (FGC 452-9); foret de Cabassou, 11 Sept 1968, J. Clastrier,

2M (FGC 467-16,-26), 2Mgen (467-70,-75) [UCLA]. Degrad des Cannes, Gravier, Pascaut (Fauran

1961b:43). Kaw, 5 m, 6 Mar 1969, J. Clastrier, 1F (FGC 3996) [UCLA]. Remire, Pripris Cabas-

sou near Port Beauregard, 5 m, 13 Mar 1967, R.X. Schick and J. Frederick, 1 IP (FG 126-42), 1F,

2L (126) [UCLA].

GUYANA. Demerara: Hyde Park, 1 Aug 1941, L.E. Rozeboom, 1M, 1F (BGR 5) [UCLA].

PANAMA. Bocas del Toro: Almirante, 10 m, 29 Apr 1963, A. Quinonez, 2M (PA 288); 14 or

15 Apr 1964, A. Quinonez, 1F (PA 666) [UCLA]. Canal Zone: Mojinga Swamp, 3 Aug 1932, 1M;

Aug 1932, 6Mgen [USNM]. Darien: El Real, 8 Aug 1952, 1M (GML 01325) [USNM]. Panama:

Pacora, 16 Oct 1946, W.H.W. Komp, 1M [USNM]. Tocumen, 1 Oct 1946, W.H.W. Komp, 1M; 10

Oct 1946, W.H.W. Komp, 2M [USNM];5 m, 11 Sept 1963, A. Quinonez, 1 lpM (PA 553-14) [UC-

LA]. No data: 1Mgen [USNM].

SURINAME. Para: Onverwacht, 10 m, 7 Apr 1967, H. deKruif and Nur, 1 pM (SUR 226-100),

4L (226), 1 lpF (SUR 227-20) [UCLA]. Suriname: Paramaribo, Ma Retraite, 5 m, 22-27 Aug

1963, P. Bolwerk, 2M (SUR 17) [UCLA]. ,

TRINIDAD. Nariva: Bush Bush Forest, Nariva Swamp, near sea level, 15 Sept 1959, T.H.G.

Aitken, 2M; 27 Nov 1963, M. Takahashi, 1F; 4 Dec 1963, M. Takahashi, 1M [USNM]; 27 Dec

1963, TRVL, 1M (TR 22) with genitalia slide (660531-6), 2F, SP, 8L (TR 22); 16 Jan 1964, T.H.

G. Aitken, 1M (TR 32) with genitalia slide (660420-3); 18 Feb 1964, TRVL, 4L (TR 74), 1 lpM

(TR 75-137), 1 IpM (TR 76-138), 1 IP (76-139), 3L (76), 4L (TR 80); 17 Mar 1964, TRVL, 3L

(TR 208); 23 Mar 1964, TRVL, 1L (TR 236); 31 Mar 1964, TRVL, 2L (TR 243), 1L (TR 244), 1

L (TR 245), 3L (TR 246); 20 Apr 1964, TRVL, 3 pF (TR 347-126-128), 1L (347), 3L (TR 348),

1 IP (TR 349-160), 4L (349), 1 pM (TR 350-126), 2L (350), 1 pM (TR 351-127), 2 pF (351-121,

-126), 2 pM (TR 352-127,-128), 1 lpF (TR 353-137), 1 pM (353-125), 4 pF (353-122-124,-128), 1

pM (TR 354-130), 1 pF (354-129); 28 Apr 1964, TRVL, 1 lpM (TR 358-193), 1L (TR 359), 1 IP

(TR 360-116), 1L (TR 361), 3L (TR 362), 4L (TR 363); 19 May 1964, TRVL, 1 lpF (TR 415-

115), 1 IP (415-120), 2L (415); 26 May 1964, TRVL, 1L (TR 421), 1L (TR 422), 2L (TR 423); 2

June 1964, TRVL, 1 IP (TR 433-130), 3L (433), 1 lp (TR 434-138), 1L (434), 1 IP (TR 435-133),

2 IpF (TR 436-139,-140), 2L (TR 438), 1 lpF (TR 439-140), 1 IpF (TR 440-186); 9 June 1964,

TRVL, 1 lpF (TR 447-160), 1 lpF (TR 448-182); 31 July 1964, TRVL, 2L (TR 580), 4L (TR

581), 7L (TR 582), 6L (TR 583); 29 Mar 1965, TRVL, 2L (TR 1065), 1L (TR 1067), 1L (TR

1070), 1L (TR 1071) [UCLA]. St. Andrew: Coryal, 50 m, 18-19 June 1964, A. Guerra, 3F (TR

499) [UCLA]. Esperanza Estate, 6-7 Oct 1969, T.H.G. Aitken, 1M [USNM]. La Fortune Estate,

20 Nov 1959, T.H.G. Aitken, 1M [USNM]. Rio Grande Forest, 4 Oct 1958, T.H.G. Aitken, 1M

[USNM]. Turure Forest, 30 m, Oct 1966, F. Guerra, 1F (TR 1618) [UCLA]. Valencia Road 6

milepost, 30 m, 1 Oct 1964, A. Guerra, 1 IlpM (TR 743-105), 1 lpF (743-105), 3M, 2F, 5P, 2L

[UCLA]. St. George: Agua Santa, 30m, 15 July 1965, A. Guerra, 3L (TR 1253); 29 July 1965,

A. Guerra, | IpF (TR 1292-20), 1 pF (1292-101), 1P, 1L (1292); 12 Aug 1965, A. Guerra, 10L

(TR 1317) [UCLA].

VENEZUELA. Monagas: Caripito, July 1937, P. J. Anduze, 1Mgen [USNM]. Zulia: Catatum-

bo River, L.H. Dunn, 2M, 1Mgen,11 F [USNM]. Locality unspecified: 11 Feb 1973, CDC, 1F

(VZ 423) [UCLA]. No data: 1Mgen [USNM]; 1M, det. P.J. Anduze [FH; 10107].

2. Culex (Aedinus) clastrieri Casal & Garcia

Figs, 1.5

1968. Culex (Eubonnea) clastrieri Casal and Garcia 1968:119-120. *TYPE: Holotype male,

Belem, Para, Brazil, 29 Nov 1959, Eber and Evangelista [INM] .

Culex (Aedinus) clastrieri of Stone (1970: 164); Belkin, Schick and Heinemann (1971:27); Knight

and Stone (1977:195); Heinemann and Belkin (1978b:407).

Culex (Eubonnea) clastrieri of Xavier and Mattos (1975:247, as Eubonnea).

FEMALE. Unknown but probably very similar to amazonensis in coloration.

MALE. Wing: 2-3 mm. Proboscis: 1.6 mm. Forefemur: 1.35 mm. Very similar

to amazonensis in general features and coloration, differing in the following. Erect

16 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

scales on vertex brown. Pleural integument light brown. Abdominal tergites com-

pletely brown; sternites dark scaled.

MALE GENITALIA (fig. 5). Diagnostic characters as in key. Segment IX: Ter-

gite lobes distinctly separated, digitform, each with 8-10 short, weak apical setae.

Sidepiece: Ovate, length about 2.0 of greatest width; lateral surface with longer bris-

tles, sternal side with shorter setae; scales present on basolateral and sternal surfaces;

3 broad saberlike setae and 3 rounded bristles distad, and a dense patch of rounded

bristles (25-30) laterad of lobe. Lobe: Submedian in position, stem distinct, undi-

vided; appendage moderately long, pointed, about 0.7 of stem; stem with a subbasal

and a subapical seta. Clasper: Long, about 0.65 length of sidepiece; expanded sub-

apically, gradually tapering distally; expanded region bearing external ridges contin-

ued proximally and distally as fine striations; spiniform short, simple. Lateral Plate:

Basal hook broadly sclerotized and strongly curved; apical process broadly rounded

and denticulate; sternal part split into 2 short processes. Proctiger: Cercal sclerite

short, triangular; cercal setae absent. Paraproct with 6 blunt apical teeth.

SYSTEMATICS. Culex clastrieri is known only by the male and can be distin-

guished from other species of Aedinus in the male genitalia by the combination of:

(1) expanded portion of clasper gradually tapering distally, (2) dorsal surface of side-

piece laterad of lobe with a dense patch of bristles, (3) stem of lobe with a subbasal

and a subapical seta, and (4) moderately long, pointed rod on stem of lobe.

We have seen only 3 males of clastrieri from French Guiana, and they appear to be

similar to the description of the type.

BIONOMICS. The immature stages are unknown but will probably be found in

swamp margins and root caves in swamp interiors. The 3 males from French Guiana

were caught overnight (1700-1000 h) in a Chamberlain light trap in a partially cut

forest. |

DISTRIBUTION (fig. 1). C. clastrieri has been reported from coastal areas of Pa-

ra, Brazil and French Guiana.

Material examined: 4 specimens; 4 males; no individual rearings.

BRAZIL. Para: Belem, 29 Nov 1969, Eber and Evangelista, 1M [INM, holotype].

FRENCH GUIANA. Guyane: Cayenne, Raban, 5 m, 2-3 Feb 1965, T.H.G. Aitken, R. Martinez

and A. Guerra, 3M (FG 43) [UCLA].

3. Culex (Aedinus) guyanensis Clastrier

Figs. 1,5

1970. Culex (Eubonnea) guyanensis Clastrier 1970:115-118. TYPE: Holotype male ({FGC]

3425-12), foret de Cabassou, Cayenne, French Guiana, 24 May 1968, J.Clastrier [MNHP].

Culex (Aedinus) guyanensis of Knight and Stone (1977:195).

Culex (Eubonnea) guyanensis of Fauran arid Pajot (1974:103).

Culex (Aedinus) clastrieri of Heinemann and Belkin (1978b:437).

FEMALE. Unknown, but probably very similar to amazonensis in coloration.

MALE. Wing: 2.6 mm. Proboscis: 1.67 mm. Forefemur: 1.5 mm. Very similar

to amazonensis in general features and coloration, differing in the following. Pleural

integument brown. Abdominal tergites completely brown; sternites dark scaled.

MALE GENITALIA (fig. 5). Not seen; based on the description of Clastrier (1970:

116-118). Diagnostic characters as in key. Segment IX: Tergite lobes distinctly sep-

arated, digitiform, each with 9,10 short, weak, apical setae. Sidepiece: Ovate, length

about 2.0 of greatest width; lateral surface with longer bristles, sternal side with

shorter setae; scales present on basolateral and sternal surfaces; 2 broad saberlike se-

tae and 4 rounded bristles distad, and a dense patch of about 25 bristles laterad of

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 17

lobe. Lobe: Submedian in position, stem distinct, undivided; appendage short,

blunt apically, about 0.5 of stem; stem with a subapical seta. Clasper: Long, about

0.7 length of sidepiece; expanded subapically, gradually tapering distally; expanded

region bearing external ridges, continued proximally and distally as fine striations;

spiniform short, simple. Lateral Plate: Basal hook broadly sclerotized and strongly

curved; apical process broadly rounded and denticulate; sternal part split into 2

short processes. Proctiger: Cercal sclerite and paraproct apparently as in amazonen-

sis.

SYSTEMATICS. Culex guyanensis is known only by the holotype male from Cay-

enne, French Guiana and is separated from other members of Aedinus in the male

genitalia by the combination of: (1) expanded portion of clasper gradually tapering

distally, (2) dorsal surface of sidepiece laterad of lobe with a dense patch of bristles,

(3) stem of lobe with only 1 subapical seta, and (4) short, blunt appendage on stem

of lobe.

The holotype male of guyanensis appears to be very similar to clastrieri in general

features and coloration. |

BIONOMICS. The immature stages of guyanensis are unknown but will probably

be found in swamps. The holotype male was caught in a light trap, and therefore

like other species of Aedinus it appears to be nocturnal.

DISTRIBUTION (fig. 1). Culex guyanensis is known only from its type locality

in French Guiana.

Material examined: | specimen; | pinned male (holotype; genitalia not seen); no

individual rearings.

FRENCH GUIANA. Guyane: Cayenne, foret de Cabassou, 24 May 1968, J. Clastrier, 1M (hol-

otype, [FGC] 3425-12) [MNHP].

4, Culex (Aedinus) accelerans Root

Figs. 1,6,7

1927. Culex (Aedinus) accelerans Root 1927:581. *TYPE: Holotype male, Porto das Caixas,

Rio de Janeiro, Brazil, 15 Apr 1925, F.M. Root [USNM].

Culex (Aedinus) accelerans of Edwards (1932:220); Lane (1939:80; 1953:386); Galindo and Blan-

ton (1955:69-70); Stone and Knight (1957:58); Fauran (1961b:43); Belkin (1968b:11); Belkin,

Schick and Heinemann (1971:27); Knight and Stone (1977:194); Heinemann and Belkin (1978a:

183; 1978b:407,437,445; 1978c:537).

Culex (Eubonnea) accelerans of Stone, Knight and Starcke (1959:280); Stone (1961 :47); Forattini

(1965:36,186,187); Aitken, Spence et al. (1969:210,212-213).

FEMALE. Wing: 2.6-2.7 mm. Proboscis: 1.4-1.5 mm. Forefemur: 1.4mm. Ab-

domen: 2.0-2.1 mm. Similar to amazonensis in general features and coloration, dif-

fering in the following. Erect scales on vertex brown. Pleural integument dark with

a smoky bluish gray iridescence. Abdominal tergites completely dark; sternites dark

scaled.

MALE. Wing: 2.50-2.55 mm. Proboscis: 1.50-1.55 mm. Forefemur: 1.4 mm.

Similar to female in general features. Palpus short, about 0.2 length of proboscis.

MALE GENITALIA (fig. 6). Diagnostic characters as in key. Segment IX: Ter-

gite lobes moundlike, distinctly separated, each bearing a subbasal digitiform append-

age with 5,6 short, weak apical setae. Sidepiece: Ovate, length about 2.0 of greatest

width; tergal and lateral surfaces with longer bristles, sternal side with shorter setae;

scales on lateral surface. Lobe: Submedian in position, stem distinct, undivided;

apex with 2 closely appressed pointed sabers; 1 subbasal seta on stem of lobe.

18 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Clasper: Long, about 0.7 length of sidepiece; uniformly stout up to 0.7 and attenu-

ate distally; a distinct subapical seta on distal 0.8; spiniform short, simple. Lateral

Plate: Basal hook broadly sclerotized and strongly curved; apical process broadly

rounded and denticulate; sternal part split into 2 short processes, Cercal sclerite

short, triangular; cercal setae absent. Paraproct with 5,6 blunt apical teeth.

PUPA (fig. 6). Abdomen: 2.3-2.7 mm. Trumpet: 0.55-0.60 mm; index 7.0-7.5.

Paddle: 0.65-0.75 mm. As described for the subgenus with the following additional

features. Diagnostic characters as in key. Cephalothorax: Integument with charac-

teristic pattern of pigmentation; middorsal ridge areas, lateral and posterior parts of

wing case, basal areas of trumpet and leg cases moderately pigmented, rest lighter.

Hair 4-C usually double; 5-C usually 5-branched (4-7) and subequal in length to 4+C;

hair 7-C usually triple (2-4); hairs 8,9-C usually double (2-3). Trumpet: Moderately

pigmented, apical flare indistinct; pinna short, about 0.2 length of trumpet; trache-

oid distinct; central area darker. Metanotum: Hair 10-C multiple (5-11), weakly

branched; 11-C single; 12-C usually 4-branched (2-4). Abdomen: Except for tergites

V,VL, VIII uniformly lightly pigmented; tergites V,VI and median and lateral areas of

VIII moderately pigmented. Hair 1-II multiple (16-28), weakly branched, indistinct-

ly resembling float hair (1-I); hair 1-IIJ-VI multiple, usually with 3-8 branches; 2-III

mesad and 2-IV-VII laterad of hair 1; hair 9-VII,VIII single; 4-VIII usually double (2-

4). Paddle: Lightly pigmented; pigmented spot apically at level of midrib; apex

smoothly rounded. Male genital lobe extending to 0.37, female to 0.17-0.18 of pad-

dle.

FOURTH INSTAR LARVA (fig. 7). Head: 0.6 mm. Siphon: about 1.5 mm.

Anal Saddle: 0.3 mm. As described for the subgenus, with the following additional

features. Diagnostic characters as in key. Head: Width about 1.5 of length. Mental

plate with 8 (7-9) lateral teeth. Hair 4-C usually double (2,3), cephalad of 5-C; hair

5-C single; 7-C with 5-7 strong barbed branches, extending to base of antenna; 11-C

with 4-7 weak branches. Antenna: Integument yellowish, spiculate to base of 1-A;

hair 1-A with 23-25 barbed branches. Thorax: Integument glabrous. Hairs un-

branched or with branches of varied length, not stellate. Abdomen: Integument gla-

brous. Hairs 1.2.4,11,134, 1,2,5,9, (3-H, 1,2.5,7,9,13411-VL 1,2,5,8,10,13-VII un-

branched or with branches of varied length and thickness, never stellate; 6-I,II long,

double, branches subequal; 1-I long, single; 7-II short, multiple; 6-III-V moderately

long, at least triple, branches subequal; 6-VI shorter, usually with 4 subequal branch-

es (3-5). Segment VIII: Comb scales spatulate, 21-24 in number, apexes fringed.

Hair 2-VIII usually double (1,2); 5-VIII double (2,3). Siphon: Integument yellow-

ish, with a narrow basal dark ring; long, index about 14.0. Pecten with 15 (14-16)

teeth, extending to basal 0.3; each tooth with 1 large and 1,2 small denticles on ven-

tral border; with 6 pairs of subventral (1,1a-S) hairs. Anal Segment: Integument yel-

lowish. Hair 1-X short, weakly branched; 2-X long, with 2 short to moderately long

subbasal branches. Gills about length of saddle, sausage-shaped, dorsal subequal to

ventral.

SYSTEMATICS. Culex accelerans can be separated from the other members of

the subgenus in the male genitalia by: (1) simple clasper without subapical enlarge-

ment, (2) absence of specialized setae on sidepiece, and (3) 2 appressed appendages

on lobe; in the pupa by: (1) short pinna and (2) pigmented area on tergite VIII; and

in the larva by: (1) hair 5-C single, (2) 2-X with 2 additional short, subbasal branches,

and (3) 6 pairs of subventral hairs (1,1a-S) on siphon.

We have examined adults of accelerans from Panama, Colombia, Trinidad, Guyana

and French Guiana, and immatures from all the above countries except Colombia.

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 19

All populations appear to be similar in male genitalic structures and characters in im-

mature stages.

BIONOMICS. Immature stages of accelerans have been found largely along swamp

margins and once in a canal in Guyana. Biting studies in Trinidad with chickens as

bait 55 feet above the ground showed that this species is a night biter. It is not

known whether it is attracted to man.

Culex accelerans appears to be another species of Aedinus harboring arboviruses.

During a 10-year survey of arthropods for natural viral infection by the Trinidad Re-

gional Virus Laboratory, 3 viruses (VEE, Caraparu and Nepuyo) were isolated from

accelerans. While no viruses were detected during surveys in 1953-1958, 3 viruses

were isolated in the period 1959-1963 when specimens were inoculated into mice

(Aitken, Spence et al. 1969:210). It is not known, however, whether females of ac-

celerans can transmit the viruses in nature by their bites.

DISTRIBUTION (fig. 1). Culex accelerans is known from the Pacific coast of Pan-

ama east of Canal Zone, from Colombia, Trinidad, Guyana, French Guiana and the

coastal lowlands of Brazil. It is probably present in Venezuela and Suriname.

Material examined: 152 specimens; 33 males, 16 females, 22 pupae, 81 larvae; 24

individual rearings (13 larval, 4 pupal, 7 incomplete).

BRAZIL. Rio de Janeiro: Porto das Caixas, 15 Apr 1925, F.M. Root, 1M [USNM, holotype].

COLOMBIA. Boyaca: Puerto Boyaca, Lago de Palagua, 145 m, 24 Aug 1971, C. J. Marinkelle,

1M (COM 590) [UCLA]. Locality unspecified: Kerr, 1Mgen (512) [USNM].

FRENCH GUIANA. Guyane: Cayenne, Mont Cabassou, 5 m, 31 Jan-1 Feb 1965, T.H.G. Ait-

ken, R. Martinez and A. Guerra, 1F (FG 17), 6M (FG 18); Mont Cabassou, 30 m, 3-4 Feb 1965, T.

H.G. Aitken, R. Martinez and A. Guerra, 2M (FG 46); foret de Cabassou, 10 Jan 1968, E. N. De

Freitas, 1Mgen (FGC 3299-12); foret de Cabassou, 19 Jan 1968, E. N. DeFreitas, 2Mgen (FGC 33

01-45 ,-48); foret de Cabassou, 30 July 1968, J. Clastrier, 1Mgen (FGC 437-14); foret de Cabassou,

11 Sept 1968, J. Clastrier, 2Mgen (FGC 467-3,-10); foret de Cabassou, 19 Sept 1968, J. Clastrier,

1Mgen (FGC 473-7); Montagne Tigre (Fauran 1961b:43); Raban, 5 m, 3 Feb 1965, T. H. G. Aitken,

R. Martinez and A. Guerra, 1 lpM (FG 37-17), 2L (37); Raban, 2-3 Feb 1965, T. H. G. Aitken, R.

Martinez and A. Guerra, 1M (FG 43) [UCLA]. Matoury, foret de Cogneau, 5 m, 3 May 1968, J.

Clastrier, 1Mgen (FGC 3416-51) [UCLA]. Remire, Pripris Cabassou near Pont Beauregard, 5 m,

13 Mar 1967, R.X. Schick and J. Frederick, 2 lpM (FG 126-40,-41), 1 IP (126-43), 1 1 (126-42), 2

M (126) [UCLA]. Inini: Degrad Edmond, Riviere Comte (Fauran 1961b:43).

PANAMA. Darien: El Real, 21 July 1952, MTA, 1M (01290) [USNM]. Panama: Tocumen,

5m, 9 Oct 1963, A. Quinonez, 2 IpF (PA 558-102,-103), 2 IP (558-105,-108), 1L (558) [UCLA];

Galindo and Blanton (1955:70).

TRINIDAD. Nariva: Nariva Swamp, Bush Bush Forest, near sea level, 21 Apr 1964, TRVL, 5L

(TR 340), 1 pF (TR 341-133), 2L (341), 4L (TR 342), 1 IpF (TR 343-136), 4L (343), 1 1P (TR

344-127), 3L (TR 345), 4L (TR 346); 5 May 1964, TRVL, 1 lp (TR 376-132), 3L (TR 377); 9

June 1964, TRVL, 2 IpM (TR 445-159,-164), 1 IP (TR 446-1), 1 IP (TR 447-150), 2 IpF (TR 448-

158,-159); 7 July 1964, TRVL, 19L (TR 544); 31 July 1964, TRVL, 1L (TR 581), 1L (TR 582);

11 Sept 1964, TRVL, 2L (TR 688); 16 Oct 1964, TRVL, 2 IpM (TR 772-107,-108), 1 IpF (772-

109), 9L (772); Sept 1965, TRVL, 6F (TR 1438) [UCLA]; 1-15 Sept 1959, T.H.G. Aitken, 1M:

2-22 Sept 1959, T.H.G. Aitken, 1M [USNM]. St. Andrew: Rio Grande Forest, 2-9 Oct 1958, T.

H.G. Aitken, 1M, 1Mgen, 1F [USNM]. St. Patrick: Bonasse, Lillette Swamp, 29 July 1966, A.

Guerra, 4L (TR 1568) [UCLA].

20 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

SUBGENUS TINOLESTES Coquillett 1906

INTRODUCTION

Tinolestes was proposed as a distinct genus by Coquillett (1906a: 185) with the

new species Jatisquama Coquillett 1906 from Puerto Limon, Costa Rica as its type.

Howard, Dyar and Knab (1915:303-305) synonymized Tinolestes with Culex, but

Dyar (1918:102) elevated it to subgeneric rank within Culex. In 1928, Dyar placed

latisquama in the subgenus Melanoconion, and as a consequence Tinolestes became a

synonym of Melanoconion. Subsequently Lane (1953:387) restored Tinolestes to

subgeneric rank, but he synonymized the subgenera Micraedes Coquillett, Isostomyia

Coquillett and Anoedioporpa Dyar with it. In 1959, Tinolestes along with Micraedes

and Anoedioporpa were lumped into the subgenus Aedinus Bourroul by Stone,

Knight and Starcke on the basis of a short palpus in the male. But according to Bel-

kin (1968b:11), this character occurs independently in several unrelated phylads,

and therefore he elevated Tinolestes Coquillett 1906 again to subgeneric rank within

Culex with only one included species. In the present study, Tinolestes is treated as a

distinct subgenus of Culex with latisquama as its type and only species.

TAXONOMIC TREATMENT

Figs. 8-11

1906. Tinolestes Coquillett 1906a:185. *TYPE SPECIES: Tinolestes latisquama Coquillett

1906:185, Puerto Limon, Costa Rica; only included species.

Culex (Tinolestes) of Dyar (1918:92,102; 1923b:187); Bonne and Bonne-Wepster (1925:278); Bel-

kin (1968b:11); Knight and Stone (1977:272).

Culex (Tinolestes) in part of Lane (1953:387); Galindo and Blanton (1955:70); Fauran (1961:48).

Culex (Aedinus) in part of Stone, Knight and Starcke (1959:281); Stone (1961:47; 1963:135;

1967:218); Belkin (1962:179); Belkin, Schick and Heinemann (1965:13); Forattini (1965:31,

32,34,35,185,193); Bram (1967:18).

Culex (Melanoconion) in part of Dyar (1928:336-337); Edwards (1932:212,213); Cok (1935'3);

Lane (1939:66); Rozeboom and Komp (1950:78,92,106); Belkin and Hogue (1959:421).

Tinolestes of Coquillett (1906b:17,24; 1910:615); Theobald (1910:627).

FEMALE (fig. 9). Small, inornate species, dark brown to black and with unband-

ed legs. Head: Eyes not distinctly separated above antennae. Decumbent scales

narrow, linear dorsally, broader on sides and venter, predominantly dark, a few scales

whitish laterally. Erect scales on occiput moderately long, dark, forked apically and

extending to sides of vertex. Orbital and interorbital bristles strong; upper orbitals 6,

7 pairs, heavier, longer and more widely spaced than lower. Clypeus bare, dark

brown. Proboscis distinctly longer than forefemur, entirely dark scaled, with a few

basal bristles. Palpus short, about 0.2 length of proboscis, 4-segmented; segments |

and 2 ankylosed, without scales; segment 4 about 2.0 length of segment 3, both dark

scaled. Antenna slightly longer than proboscis; torus dark brown, with a few short

setae mesally; flagellar segments 2-13 with 6 moderately long bristles in basal whorls.

Thorax: Integument dark brown. Mesonotum with narrow, curved, dark brown

scales except along a pair of narrow inner dorsocentral “bare lines” extending from

anterior margin to 0.75 of its length. Acrostichal bristles absent. Bristles on anterior

promontory distinct; dorsocentrals, prescutellars and supraalars present; 3,4 posterior

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa A

fossal and 1 parascutellar developed. Antealar area above paratergite with scattered

dark scales. Median scutellar lobe with 5,6 long and 6 short marginal bristles, and a

large patch of narrow dark scales; lateral lobe with 4 long and 3 short marginal bris-

tles, and a small patch of narrow dark scales. Paratergite bare. Pleuron brown. Bris-

tles present on apn, ppn, ppl, stp, pra and upper mep; propleural area with at least

20-25 bristles; middle mep with a patch of short bristles; metameron with 3,4 short

setae. Pleural scaling restricted to apn, ppn, stp and middle mep; stp with a large

patch of flat, translucent scales covering the entire posterior half. Legs: Dark scaled.

Claws simple on all legs. Wing: Veins entirely dark scaled; all scales squamous, mod-

erately broad. Haltere: Stem pale brown; knob entirely dark scaled. Abdomen:

Laterotergite with many short to moderately long bristles. Tergites II-VII with light

basolateral patches, rest dark scaled. Sternites predominantly creamy, dark scales

restricted to apical 0.3.

FEMALE GENITALIA (fig. 9). Segment VIII partially retracted into segment

VII, apex visible, numerous bristles and scales present; tergite VIII about 0.5 of ster-

nite; apex of sternite distinctly emarginate in the middle. Tergite IX broad, sclero-

tized; lobes faintly visible, each bearing 15,16 moderately long setae; sternite IX nar-

row laterally, membranous in the middle. Postgenital plate deeply emarginate in the

middle, producing 2 conical lobes each bearing 3 apical and 2,3 subapical setae.

Cowl moderately developed, finely setose, bowed laterally to join tergite [X and arti-

culating with sigma. Insula well developed, poorly sclerotized, with a group of 13

moderately long setae, continued laterally as a narrow sigma. Cerci short, compres-

sed, approximate, each with many short to moderately long setae; area between cerci

bilobed. Spermathecae 3, one a little larger than others.

MALE (fig. 9). Coloration similar to females; sexual dimorphism of head append-

ages marked. Proboscis longer than forefemur. Palpus porrect, slender, about 0.45-

0.48 of proboscis length; 5-segmented; segments | and 2 ankylosed, without scales;

segments 3 and 4 elongated, partly ankylosed, segment 4 about 1.3 of segment 3,

both dark scaled; segment 5 short, about 0.25 of 4 and dark scaled. Antenna sube-

qual in length to proboscis; whorls of flagellar segments 1-12 of antenna strongly de-

veloped, with at least 20 long bristles; segments 12 and 13 elongate, 13 about 1.4

longer than 12; torus not swollen. Claws of foreleg and midleg enlarged, unequal;

larger claw of baecta with a submedian tooth, that of midleg without oe both

claws with basal spicules; hind claws as in female.

MALE GENITALIA (fig. 10). Segment IX: Lobes of tergite [X distinct, heavily

sclerotized, flattened, bearing long, erect bristles. Sidepiece: Roughly triangular in

outline; length about 2.0 basal width; with longer bristles on tergal and lateral sur-

faces, sternal side with shorter setae; a few scales on basolateral surface; an elongate

patch of short to moderately long bristles on tergal surface basolaterad of subapical

lobe; caudal and basomesal areas without bristles. Lobe distinct, subapical in posi-

tion, undivided. Clasper: Simple, stout, bearing an external crest; spiniform simple,

apical. Phallosome: Lateral plate of aedeagus with a broadly sclerotized “basal

hook,” a caudally directed sinuous apical process and a sternal spine. Proctiger: Ba-

solateral sclerotization well developed, digitiform; apex of paraproct with a crown of

22-24 teeth; cercal setae distinct.

PUPA (fig. 10). Cephalothorax: Middorsal ridge moderate. Integument uniform-

ly lightly pigmented. Alk hairs present, variously developed. Hair 5-C moderately

long, weakly developed, subequal in length to 4-C; hair 6-C smaller than and cepha-

lad of 7-C; hairs 8,9-C widely separated, double or triple; 8-C cephalad and 9-C dis-

tinctly caudad of trumpet base. Trumpet: Not placed on tubercle; moderately long,

22 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

index 5.0-5.5; pigmentation moderate; tracheoid distinct, short; apical portion slight-

ly flared; pinna short. Metanotum: Hair 10-C subequal in length to 11-C, multiple,

weakly branched. Abdomen: Integument uniformly lightly pigmented. Hair 3-I

double or triple; 1-II multiple, with at least 20 weak branches, faintly resembling

float hair (1-1); hair 2-III] submarginal, always mesad of 1-III; hair 5-IV,V long, single,

extending beyond apex of succeeding segments; 5-VI shorter, barely extending to

middle of succeeding segment; 6-II-VI usually single; 2-VI, VII always laterad of hair

1; hair 9-VII, VIII double, 9-VIII distinctly shorter than tergite VIII; hair 4-VIII dou-

ble; 1-[X small, single. Lobes on posterior margin of sternum mesad of 9-VIII indis-

tinct. Tergite VIII never overlapping base of tergite IX. Paddle: Uniformly lightly

pigmented; longer than wide, apex smoothly rounded; midrib strongly differentiated;

paddle margin without spicules; both paddle hairs (1,2-P) distinct, single.

FOURTH INSTAR LARVA (fig. 11). Head: Head capsule with faint imbrica-

tions; without lateral expansion on each side caudad of antenna. Labrum well differ-

entiated dorsally. Ocular lobes indistinct. Mouthbrushes with numerous filaments.

Collar moderately developed, narrow. Posterior tentorial pit a short distance from

caudal border. Maxillary suture complete, but not extending caudolaterad to collar.

Anterior border of labial plate truncate. Aulaeum with distinct filamentous spicules.

Central tooth of mental plate not shouldered. Hair O-C small, removed laterad of 1-

C; hair 1-C strong, straight; 2,3-C apparently not developed; 4-6-C closely grouped

together and caudad of level of 7-C; hair 5-C usually triple, 6-C single, both longer

than antenna; 7-C multiple, with strong barbed branches; 8-C usually triple (2,3);

hair 11-C with at least 5 (5-8) moderately long branches; 13-C closer to 11-C than

12-C; hairs 14,15-C slightly moved cephalad; basal maxillary hair (bmh) small, single;

16,17-C not developed. Antenna: Hair 1-A large, multiple, about 0.75 from base;

other hairs short to moderately long, single. Thorax: Roughly oval in outline, slight-

ly wider than long. Integument glabrous. Hairs 1-3-P on a distinct tubercle; 3-P with

6-10 weak branches; 14-P double; hairs O-P, 1,13,14-M, 1,3,13-T with many weak

branches, not stellate. Abdomen: Integument glabrous. Pigmentation uniform.

Hair 2 never moved far cephalad of 1, always within posterior half of segment. Seg-

ment VIII: Comb in a patch of 4,5 irregular rows of scales. Hairs 1,2-VIII without

basal tubercles or sclerotized plates. Siphon: Pecten teeth simple; 4 pairs of subven-

tral (1,la-S) and 2 pairs of subdorsal (2a-S) hairs on siphon; both subdorsals distad

of subventral hairs; 2-S distinct, slightly curved, without a submedian branch. Anal

Segment: Saddle complete; acus not developed; caudal margin without spines; later-

al hair (1-X) short, multiple, submarginal. Hair 2-X long with 3 short to moderately

long subbasal branches; 3-X long, single; ventral brush (4-X) with 6 pairs of branched

hairs on a grid. Gills sausage-shaped, ventral slightly shorter than dorsal.

I, Culex (Tinolestes) latisquama (Coquillett)

Figs. 8,9,10,11

1906. Tinolestes latisquama Coquillett 1906a:185. *TYPE: Lectotype male (344c), Puerto

Limon, Costa Rica, 25 Sept 1905, F. Knab (USNM, 8298; designation by Rozeboom and

Komp 1950:92).

Culex (Tinolestes) latisquama of Dyar (1918:90,102; 1922a:95-96; 1923a:177; 1923b:188; 1925:

158,161,168); Bonne and Bonne-Wepster (1925:184,188,278); Lane (1953 :387-389); Duret

and Damasceno (1955:395); Galindo and Blanton (1955:70); Forattini (1958:175); Fauran

(1961b:48); Belkin (1968b:11); Knight and Stone (1977:272); Heinemann and Belkin (1977a:

284; 1977b:443,453; 1978a:185; 1978c:497).

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa pie.

Culex (Aedinus) latisquama of Stone, Knight and Starcke (1959:282); Belkin, Schick and Heine-

mann (1965:13); Forattini (1965 :34).

Culex (Melanoconion) latisquama of Dyar (1928:336-337); Edwards (1932:212,213); Lane (1939:

66); Rozeboom and Komp (1950:78,92,106); Belkin and Hogue (1959:421).

Culex latisquama of Dyar and Knab (1906:208,222); Howard, Dyar and Knab (1915:217,222,229,

303-305); Kumm, Komp and Ruiz (1940:387); Forattini, Rabello and Cotrim (1970:43).

Tinolestes latisquama of Coquillett (1910:615); Theobald (1910:627); Stone and Knight (1957:

52),

FEMALE (fig. 9). Wing: 2.70-2.75 mm. Proboscis: 1.65-1.70 mm. Forefemur:

1.4mm. Abdomen: 1.6-1.7 mm. As described for the subgenus with the following

additional features. Head: Decumbent and erect scales dark. Palpus short, dark

scaled. Thorax: Pleuron brown, darker on stp, psp and mep; bristles on pp/ strongly

developed, at least 20 in number; scales on stp and middle mep moderately broad

and translucent. Legs: Coxae with dark scales on external surfaces; trochanters with

ventral dark scales. Forefemur and midfemur dark scaled anteriorly, posterior side

creamy basally, rest dark; hindfemur creamy ventrally and dark scaled dorsally. Tib-

iae and tarsi of all legs dark. Wings: Scales on all veins dark, squamous and moder-

ately broad. Haltere: Stem pale brown; knob entirely dark scaled. Abdomen: Ba-

solateral light patches on tergites moderately broad, rest dark scaled; sternites

creamy basally, dark scaled distally.

MALE (fig. 9). Wing: 2.55-2.60 mm. Proboscis: 2.0 mm. Forefemur: 1.50-1.55

mm. Similar to female in general features. Palpus moderately long, about 0.45-0.48

length of proboscis. Antenna subequal in length to proboscis.

MALE GENITALIA (fig. 10). Segment IX: Tergite lobes distinctly separated by

a triangular bar; each lobe flattened, wider than long, bearing 20-22 long erect bris-

tles inserted on distinct tubercles. Sidepiece: Slightly more than 2.0 as long as wide,

with a conically tapered tip; longer bristles on tergal and lateral surfaces, sternal side

with shorter setae; a few scales on basolateral surface; a dense elongated patch of 30-

35 moderately long bristles on tergal surface basolaterad of subapical lobe. Subapi-

cal Lobe: Bearing a smaller and a larger rod with cup shaped expanded tips, and a

larger inner and a smaller outer seta laterad of rods; 4,5 short to moderately long

simple setae distad of rods. Clasper: About 0.5 of sidepiece length, stout, slightly

swollen in middle, bearing an external crest; | inner and | outer seta subapically;

spiniform apical. Lateral Plate: Basal hook broadly sclerotized; apical process

strongly sclerotized, sinuous, with a pointed recurved tip; sternal spine slightly re-

curved apically, pointing laterad from dorsal aspect. Proctiger: Cercal sclerite dis-

tinct, moderately long, digitiform, pointing caudad; cercal setae 2 in number. Para-

proct with a crown of 22-24 apical blunt teeth.

PUPA (fig. 10). Abdomen: 2.50-2.55 mm. Trumpet: 0.35 mm; index about 5.0-

5.5. Paddle: 0.65-0.70 mm. Cephalothorax: Hair 4-C laterad of 5-C; hair 6-C single,

7-C double, approximate; 8-C usually triple (2,3), cephalad of 9-C. Trumpet: Mod-

erately long, brownish; contrasting with rest of cephalothorax; apex slightly flared.

Pinna about 0.20-0.25 of total length. Metanotum: Hair 10-C multiple (7-11), clos-

er to 11-C than to 12-C; hair 11-C double; 12-C usually triple (2,3). Abdomen: In-

tegument lightly pigmented. Hair 1-I,I] multiple, branches weak; 5-IV,V long, single,

extending beyond apex of succeeding segments; 5-VI short, barely extending to mid-

dle of succeeding segment; 2-III-V mesad and 2-VI,VII laterad of hair 1; 9-II-VI cau-

dolaterad of hair 6; hair 9-VII, VIII moderately long, double; 9-VIII shorter than

length of tergite VIII. Paddle: Longer than wide. Midrib distinct; external buttress

slightly developed; apex smoothly rounded, margin without spicules; both paddle

24 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

hairs (1,2-P) present, 2-P longer than 1-P. Male genital lobe extending to 0.47, female

to 0.28 of paddle.

FOURTH INSTAR LARVA (fig. 11). Head: 0.75 mm. Siphon: 1.4-1.6 mm; in-

dex about 11.0-12.0. Anal Saddle: 0.35 mm. Head: Width about 1.2 of length;

head capsule indistinctly imbricate. Hair 4-C short, double, anteriad of 5-C; hair 5-C

long, usually triple (3,4); hair 6-C long, single; 7-C long, branched (4,5), extending to

base of 1-A; hair 11-C multiple (5,5-8), weakly branched; 13-C multiple, closer to 11-

C than to 12-C. Mental plate well developed, with 10 (9-11) lateral teeth. Antenna:

Strongly pigmented, spiculose to base of 1-A; hair 1-A long, with 16-19 barbed

branches. Thorax: Integument glabrous. Hairs unbranched or with branches of var-

ied length and thickness, not stellate. Abdomen: Integument glabrous, Hairs 1 ,2,4,

11,13-I, 1,2,5,9,13-H, 1,2,5,7,9,13-II-VI, 1,2,5,8,10-13-VII unbranched or with

branches of varied length and thickness, never stellate. Hairs 6-I,II, 7-I long, double,

branches unequal; 7-II short, multiple (4-6); hair 6-III-V moderately long, double,

subequal; 6-VI long, double, branches subequal. Segment VIII: Comb scales in an

irregular patch of 4,5 rows, 50-69 in number; individual scale spatulate, apex fringed.

Hairs 1-5-VIII double or triple. Siphon: Integument light brown, with wartlike spic-

ules, base with a dark ring. Pecten extending to basal 0.25, teeth 12-15 in number;

individual tooth without denticles. Hair 1-S distinctly caudad of last pecten tooth.

Subventral hairs (1,1a-S) 4 pairs, short, at least double; subdorsal hairs (2a-S) 2 pairs,

short, single, located on distal 0.25 of siphon. Anal Segment: Integument brown

and indistinctly imbricate. Hair 1-X short, weakly branched (4,3-5); hair 2-X with 3

additional short to moderately long basal branches. Gills short, ventral smaller than

dorsal.

SYSTEMATICS. Tinolestes can be readily separated from other subgenera of Cu-

lex in the New World by the following characters: in the adults by (1) strongly devel-

oped propleural bristles, (2) presence of a broad patch of translucent scales covering

almost entire stp, (3) 3,4 short setae on metameron, and (4) distinct patch of scales

and hairs on middle mep. The moderately long palpus of the male, about 0.45-0.48

length of proboscis, distinguishes it from most members of Culex. In the male geni-

talia, Tinolestes differs from other subgenera by the combination of: (1) morpholo-

gy of the clasper, (2) position and structural components of the subapical lobe, (3)

nature of the phallosome, and (4) number of apical teeth on the paraproct. In the

pupa, Tinolestes can be distinguished by the combination of: (1) position of hair 2-

VI, (2) hair 11-C always double, (3) hair 9-VII, VIII always double, and (4) hair 9-

VIII shorter than tergite VIII. In the larvae, Tinolestes can be distinguished from

most of the subgenera except Aedinus and Lutzia by the slightly imbricate head cap-

sule; however, it is separated from Aedinus by the absence of an attached acus on the

anal saddle, and from Lutzia by the structure of the mouthbrushes.

Tinolestes is undoubtedly a distinct subgenus of Culex that was erroneously

lumped under Aedinus (Stone, Knight and Starcke, 1959:281-282). Later, Belkin

(1968b: 11-12) suggested separating the subgenera Eubonnea and Aedinus of Stone,

Knight and Starcke (1959) into Aedinus (=Eubonnea), Anoedioporpa, Micraedes

and Jinolestes. In spite of the similarities of Tinolestes with other subgenera of the

Melanoconion complex, it is distinct in adults, immature stages and in ecology

(breeding sites). Although there is a slight overlap with Aedinus and Belkinomyia,

Tinolestes is a very distinct group and should be retained as a subgenus of Culex.

BIONOMICS. The immature stages of latisquama have been found in crabholes,

and are often catholic in their association with other crabhole breeding mosquitoes.

This species has been found with 6 species of Deinocerites: curiche Adames 1971,

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 25

colombianus Adames 1971, epitedeus (Knab 1907), melanophylum Dyar & Knab

1907, panamensis Adames 1971 and pseudes Dyar & Knab 1909; in addition, it is of-

ten associated with members of the Inflictus Group of Culex (Culex). Rarely it has

been collected in association with Anopheles (Anopheles) eiseni Coquillett 1902 (CR

524, PA 573), Culex (Melanoconion) iolambdis Dyar 1918 (CR 525, PA 615) and

Culex (Lutzia) allostigma (Howard, Dyar & Knab 1915) (CR 525).

Very little is known about the bionomics of the adults. Apparently the adults use

crabholes as resting sites. The presence of a few engorged females in the collection

suggests that they are zoophilic as they show no inclination to bite or even alight up-

on humans (Howard, Dyar and Knab 1915:305).

DISTRIBUTION (fig. 8). Tinolestes is primarily Central American in distribution.

To date it has been collected from the Caribbean side of Honduras, Nicaragua and

Costa Rica, and from the Pacific and Caribbean sides of Panama and Colombia.

The report of latisquama from Suriname (Stone, Knight and Starcke 1959:282) is

erroneous, and the record of it on the basis of 1 male supposedly collected in 1906

by J.B. Vanduzee in Estero, Lee County, Florida, U.S.A. (Stone 1968:101) is prob-

ably due to an erroneous labelling (Belkin 1970:57-58), which occurred several times

during the course of preparation of material for the studies on the ‘“‘Mosquitoes of

North and Central America and the West Indies’ (Howard, Dyar and Knab). Culex

latisquama may have a wider distribution in Central America, but extensive collec-

tions from different localities on both the Caribbean and Pacific sides should be

made to ascertain this.

Material examined: 615 specimens; 99 males, 104 females, 98 pupae, 314 larvae;

91 individual rearings (65 larval, 18 pupal, 8 incomplete).

COLOMBIA. Antioquia: Turbo, Brazo del Coco, near sea level, 29 Aug 1967, A.J. Adames and

A. Quinonez, 1 pM (COA 30-101), 5 IpM (COA 31-10,-11,-14,-15,-19), 7 lpF (31-12,-13,-16,-17,

-20,-22,-23), 2 pM (31-100,-101), 11 L (31), 2 lpF (COA 32-10,-11), 2 IpF (COA 33-11,-12), 1 pM

(33-100), 1 IP (33-10) [UCLA]. Choco: Coredo, El Naranjo, near sea level, 26 Aug 1967, A.J.

Adames and A. Quinonez, 1 pF (COA 17-102);31 Aug 1967, A.J. Adames and A. Quinonez, 1 IpF

(COA 48-17), 2L (48), 2 lpM (COA 50-15,-17), 1 lpF (50-16) [UCLA]. Magdalena: Near mouth

of Rio Buritaca, near sea level, May-June 1970, H.G. Henning, 20M, 22F (COA 100), 4L (COA

101), 1M (COA 102) [UCLA].

COSTA RICA. Limon: Cieneguita, June 1946, D.C.R. Butts, 1M [USNM]. Puerto Limon, 25

Sept 1905, F. Knab, 1M, 1Mgen, 8F (344c, type series) [UCLA, USNM]. Portete, near sea level,

30 Sept 1971, D. Schroeder, 1M, 1F (CR 399); 1-2 Oct 1971, D. Schroeder, 1M (CR 461); 2 Oct

1971, D. Schroeder, 1 IpM (CR 465-11), 3 IpF (465-13-15), 1 pM (465-110), 4 pF (465-100-102,

106); 7 Nov 1971, D. and K. Schroeder, 3 IpM (CR 520-31-33), 5 IpF (520-30,-34,-35,-56,-57), 6L

(520), 1M (CR 523), 1 lpM (CR 524-61), 3 IpF (524-60,-62,-63), 1L (CR 525) [UCLA]. Westfalia,

near sea level, 4 Dec 1962, C.L. Hogue and W.A. Powder, 1L (CR 76) [UCLA].

HONDURAS. Cortes: Puerto Cortes, Rio Mar, near sea level, 15 Aug 1967, A.J. Adames, 1 pM

(HON 84-100) [UCLA]. Colon: Trujillo, Rio Cristales, near sea level, 7 Mar 1945, 1M [USNM].

NICARAGUA. Zelaya: Bluefields, near sea level, 12 Sept 1967, A.J. Adames and A. Herrera, 5

L (NIC 69), 2F (NIC 70), 6L (NIC 71); 25-26 Nov 1971, D. Schroeder, 1F (NIC 101). Punta Ma-

saya, 26 Nov 1971, D. and K. Schroeder, 2 IpM (NIC 109-33,-34), 5L (109); 27 Nov 1971, D. and

K. Schroeder, 2 lpF (NIC 111-20,-32), 2L (111); 27-28 Nov 1971, D. Schroeder, 1F (NIC 123); 28

Nov 1971, D. and K. Schroeder, 4 IpM (NIC 124-10,-11,-15,-16), 4 lpF (124-12-14,-16), 4 pM

(124-100-103), 1P, 30L (124) [UCLA].

PANAMA. Bocas del Toro: Almirante, near sea level, 28 Apr 1963, A. Quinonez, 1L (PA 273),

1 pF (PA 275-101), 1 lp (275-102) [UCLA] ; Feb 3, 1Mgen [USNM]. Canal Zone: Fort Sherman,

12 Nov 1919, J. Zetek, 1M [USNM]. Miraflores, July 1912, J. Zetek, 2F [USNM]. Mojinga

Swamp, Aug 1932, 1Mgen; 12 June 1933, 1Mgen; 17 June 1952, MTH, 1M (01164); 18 Mar, 1M

gen [USNM] ; 10 m, 29 Jul 1972, J. H. Arnell, 1 IlpM (PA 1149-14) [UCLA]. Rodman Naval Sta-

tion, near sea level, 11 Dec 1965, R.X. Schick and A. Quinonez, 1 lpF (PA 888-20), 3 lpM (PA

26 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

889-11-13), 2 lpF (889-14,-16), 2 lp (889-10,-19), 1 IP (889-20), 1P (889); 12 Dec 1965, A. Quin-

onez, 1 IP (PA 891-17), 1P, 3L (891), 1 IP (PA 895-10), 1L (895); 15 Dec 1965, R.X. Schick and

A. Quinonez, 1 lpM (PA 907-11), 1 lpF (907-10), 1 IpM (PA 908-10), 1 pM (908-101), 1P (908), 1

lpM (PA 909-10), 3 lpM (PA 910-11,-12,-14), 2 lpF (910-16,-19), 1 IP (910-15), 3M, 2P, 3L (910)

[UCLA]. Colon: Maria Chiquita, near sea level, 6 Sept 1967, A. J. Adames, 1F (PA 1008) [UCL

A]. Pina, near sea level, 30 Nov 1963, A. Quinonez, 1 lpF (PA 573-101), 1 pM (573-103), 23L

(573), 1M, 2F (PA 574) [UCLA]. Portobelo, near sea level, 9 Dec 1963, A. Quinonez, 4L (PA

598); 11 Dec 1963, A. Quinonez, 1F (PA 604) [UCLA]. Darien: El Real, near sea level, 12 Jan

1964, A. Quinonez, 1L (PA 620); 13 Jan 1964, A. Quinonez, 4L (PA 621) [UCLA]. Jaque, near

sea level, 18 Dec 1963, A. Quinonez, 17L (PA 611); 20 Dec 1963, A. Quinonez, 98L (PA 615)

[UCLA]. Panama: El Libano, near sea level, 27 Aug 1963, A. Quinonez, 10M, 13F (PA 535); 26

Nov 1963, A. Quinonez, 14L (PA 566), 3M, 1F (PA 568) [UCLA]. Locality unspecified: Male,

det. J. Lane [FH, 6577; Forattini, Rabello and Cotrim (1970:43)] .

2U.S.A. Florida: Lee County, Estero, J.B. Vanduzee, 1M (with genitalia slide, no. 68-19) [US

NM]. Probably erroneous record, see p. 25 and below.

NO DATA. A.H. Jennings, 6M,5F [USNM]|. 1Mgen [USNM]. Probably collected in Panama

or Canal Zone as Jennings worked there. It is possible that the male supposedly from Florida

(above) was actually collected by Jennings and misplaced and erroneously labelled.

SUBGENUS ANOEDIOPORPA Dyar 1923

INTRODUCTION

Anoedioporpa was proposed as a distinct subgenus of Culex by Dyar (1923b:

190), with Culex conservator Dyar & Knab 1906 as its type species. Besides con-

servator, Dyat’s Anoedioporpa included originator Gordon & Evans 1922, bifolia-

tus Dyar 1922, corrigani Dyar & Knab 1907, homoeopas Dyar & Ludlow 1921

and restrictor Dyar & Knab 1906. All the other species we are including in this

subgenus were previously assigned to different subgenera of Culex such as Isosto-

myia (Dyar 1918:102; Edwards 1932:217), Melanoconion (Dyar 1925:158; 1928:

336) or Tinolestes (Lane 1953:387). In 1959, Stone, Knight and Starcke included

Tinolestes, Micraedes and Anoedioporpa in the subgenus Aedinus Bourroul 1904,

and as a result all species were lumped into a composite group on the basis of a

short palpus in the males. But as Belkin (1968b:11) indicated, this character has

evolved independently in several unrelated phylads, and he elevated Anoedioporpa

to subgeneric rank. In the present paper, Anoedioporpa is treated as a distinct

subgenus of Culex, with 12 species assigned to it. As well as the species presently

considered to be members of Anoedioporpa (Knight and Stone 1977:195-196), we

are also including restrictor Dyar & Knab 1906, presently considered to belong to

the subgenus Microculex (Knight and Stone 1977:269).

TAXONOMIC TREATMENT

Figs. 12-30

1923. Anoedioporpa Dyar 1923b:190. *TYPE SPECIES: Culex conservator Dyar & Knab

1906:221-222, St. Joseph, Trinidad; designation by Dyar (1923b:190).

Culex (Anoedioporpa) of Bonne and Bonne-Wepster (1925:184,260); Belkin (1968b:11).

Culex (Anoedioporpa) in part of Knight and Stone (1977:195-196).

Culex (Isostomyia) of Dyar (1918:92,102-103); Gordon and Evans (1922:327); Edwards (1932:

217-218); Senevet and Abonnenc (1939:112); Lane (1939:73); Anduze (1941a:16); Floch and

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa Zz]

Abonnenc (1942:9; 1947:6); Lane and Whitman (1943:397); Arnett (1948:189; 1950: 107);

Lane (1949:255); Foote (1954:3,4); Horsfall (1955:547,548).

Culex (Aedinus) in part of Stone, Knight and Starcke (1959:281); Belkin (1962:179); Stone (1961:

47; 1963:135; 1967:218); Forattini (1965:31,32,34,35,185,193); Cova Garcia, Sutil and Raus-

seo (1966a:27; 1966b:342); Bram (1967:18).

Culex (Microculex) in part of Knight and Stone (1977:269).

Culex (Melanoconion) in part of Dyar (1925:158; 1928:336); Komp (1935:3).

Culex (Tinolestes) in part of Lane (1953:387); Duret and Damasceno (1955:393); Galindo and

Blanton (1955:70); Fauran (1961 :43).

FEMALES (fig. 13). Usually small, inornate species, tan to brown and with un-

banded legs. Head: Eyes not distinctly separated above antennae. Decumbent

scales narrow and linear dorsally, white; broader on sides and venter. Erect scales on

occiput long, forked apically, light to moderately brown, extending to sides of ver-

tex. Orbital and interorbital bristles strong; upper orbitals 5 pairs, heavier, longer

and more widely spaced than lower. Clypeus bare, dark brown. Proboscis longer

than forefemur, entirely dark scaled, with a few basal bristles. Palpus short, about

0.16 length of proboscis, 4-segmented; segments 1 and 2 ankylosed, without scales;

segment 4 about 1.5 of segment 3, both dark scaled. Antenna slightly shorter than

proboscis; torus brown, with a few short setae mesally; flagellar segments 2-13 with

6 moderately long bristles in basal whorls. Thorax: Integument tan to brown. Mes-

onotum predominantly with narrow, curved, auburn scales except along a pair of

narrow inner dorsocentral “bare lines’ extending from anterior margin to 0.75 of its

length; anterior margin with narrow white scales. Acrostichal bristles usually absent

on disc, present in restrictor and corrigani. Bristles on anterior promontory distinct;

dorsocentrals, prescutellars and supraalars always present, variously developed; | pos-

terior fossal and | parascutellar always developed. Antealar area above paratergite

with scattered scales. Median scutellar lobe with 5,6 long and 6 short marginal bris-

tles, and a large patch of narrow dark scales; lateral lobes with 3 long and 3 short

marginal bristles, and a very small patch of narrow dark scales. Paratergite bare.

Pleuron yellowish to tan. Bristles present on apn, ppn, ppl, stp, pra and upper mep;

lower mep with or without a strong bristle; metameron bare. Pleural scaling restrict-

ed to ppn and stp; upper ppn with a few light to dark scales; stp with a few scattered,

flat, translucent scales along bristles. Legs: Coxae with white scales on external sur-

face; trochanters with ventral white scales. Forefemur and midfemur dark anteriorly,

posterior side predominantly creamy, dark scales restricted to dorsal side; hindfemur

predominantly creamy on both sides, dark scales only dorsally. Tibiae and tarsi of

all legs with auburn to dark brown scales. Claws simple on all legs. Wing: Veins en-

tirely dark scaled; plume scales on Rs, R, and R;. Haltere: Stem pale; knob entire-

ly dark scaled. Abdomen: Laterotergite with many short to moderately long bris-

tles. Tergites II-VII with creamy basolateral patches, rest dark scaled. Sternites with

creamy scales.

FEMALE GENITALIA (fig. 13). Only conservator, the type species, studied.

Segment VIII partially retracted into segment VII, apex visible, numerous bristles

and scales present; tergite VIII about 0.65 of sternite; apex of sternite slightly emar-

ginate. Tergite IX uniformly narrow, lobes not prominent, with 8 moderately long

setae. Postgenital plate about 0.6 length of cercus, not joined to cowl basally; apex

broadly emarginate, with a patch of 16,17 scattered setae. Cowl moderately devel-

oped, finely setose, bowed laterally to join tergite [X and articulating with sigma.

Cerci short, compressed, widely separated, with many short to moderately long setae;

area between cerci not lobed. Insula poorly developed, with a group of 8-10 short

28 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

setae, continued laterally as finely setose sigma. Spermathecae 3, one a little larger

than others.

MALES (figs. 13,28). Coloration similar to females; sexual dimorphism of head

appendages marked. Proboscis longer than forefemur. Palpus porrect, about 0.16

length of proboscis, but subequal in restrictor; entirely dark scaled; 4-segmented

(Conservator Group) or 5-segmented (Restrictor Group). Antenna subequal in

length to proboscis; whorls of flagellar segments 1-12 strongly developed, with about

20-24 long bristles; segments 12 and 13 elongate, 13 about 1.45 of 12; torus as in fe-

males, Claws of foreleg and midleg enlarged, unequal; larger claw with a submedian

and smaller with a basal tooth; both claws with basal spicules. Hindclaws as in fe-

males.

MALE GENITALIA (figs. 14,29). Segment IX: Lobes of tergite usually mound-

like, small, widely separated, bearing a variable number of short to moderately long

setae. Sidepiece: Roughly conical, length about 2.0 of basal width; tergal surface

with longer bristles and a few basal scales, sternal side with shorter setae. Lobe post-

median in position, with distinct proximal and distal divisions; stem of proximal di-

vision distinct, bearing 2 rods apically; distal division with specialized apical setae,

and with or without leaves on stem. Clasper: Comparatively simple, about 0.6 of

sidepiece; distal 0.3 with many closely set spiculelike transverse ridges externally.

Phallosome: Lateral plate of aedeagus with a broadly sclerotized “‘basal hook,” a

caudally directed apical process and a shorter sternal process. Proctiger: Basolater-

al sclerotization moderately developed, usually produced into a digitiform append-

age below tergite IX; apex of paraproct with a crown of 6-13 teeth; cercal setae vari-

able.

PUPAE (figs. 14,29). Cephalothorax: Middorsal ridge moderate. Integument uni-

formly lightly to moderately pigmented. All hairs present, variously developed. Hair

5-C short to moderately long, weakly developed, subequal in length to 4-C; hair 6-C

smaller than and cephalad of 7-C; hair 7-C double or triple; 8,9-C widely separated,

usually single, 8-C caudolaterad and 9-C caudad of trumpet base. Trumpet: Not

placed on tubercle; moderately long, index 5.0-9.0; strongly pigmented; tracheoid

distinct, short; apex not flared; pinna short. Metanotum: Hairs 10,11-C removed

from 12-C, moderately close together; 10-C usually with fewer than 6 branches; 1 1-

C usually single, rarely double. Abdomen: Without distinct pattern of pigmentation.

Hair 3-I usually single; 1-I] multiple, branches short and weak, not resembling float

hair (1-1); hair 2-II-VII submarginal, situated either laterad or mesad of hair 1; hair

5-IV,V moderately long, single, usually extending to apex of succeeding segment; 5-

VI moderately long, usually single, never extending beyond basal 0.6 of succeeding

segment; 6-II-VI usually single, rarely double; 2-VI,VII laterad of hair 1; hair 9-II-VI

caudolaterad of hair 6; hair 9-VII at least 3-branched, 9-VIII 4-branched and about

0.5 of segment length, both strongly developed; 4-VIII usually double; 1-IX small,

single (absent in restrictor). Lobes on posterior margin of sternum VIII distinct.

Tergite VIII slightly overlapping base of tergite IX. Paddle: Without any pigmented

spot; longer than wide, apex rounded; midrib strongly differentiated; external but-

tress distinct; paddle margin without spicules; both paddle hairs (1,2-P) apparently

not developed.

FOURTH INSTAR LARVAE (figs. 15,30). Head: Width about 1.2 of length.

Head capsule with conspicuous lateral expansion on each side caudad of antenna.

Labrum well differentiated dorsally. Ocular lobes slightly marked. Mouthbrushes

with numerous filaments. Collar moderately developed, narrow. Posterior tentorial

pit a short distance from caudal border. Maxillary suture complete, extending a

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 29

short distance caudolaterad of pit. Anterior border of labial plate truncate. Aulae-

um with distinct filamentous spicules. Mental plate well developed, with 7-9 lateral

teeth; median tooth usually shouldered. Hair O-C small, removed laterad of 1-C; hair

1-C strong, straight; 2,3-C apparently both not developed; 4-C small, usually single

and anteromesad of 5,6-C; hair 5-C multiple, longer than antenna; 6-C anterolaterad

of 5-C, single and distinctly longer than antenna; 7-C multiple, with barbed branches;

8-C usually double, rarely triple; 1 1-C with 2-6 weak branches; 13-C slightly closer to

12-C than to 11-C; 14,15-C moved slightly anteriad; basal maxillary hair (bmh) small,

single; 16,17-C not developed. Antenna: Moderately long, with distinct spinelike

spicules in basal 0.75. Hair 1-A large, multiple and inserted 0.75 from base; other

hairs short to moderately long, single. Thorax: Roughly oval in outline, slightly wi-

der than long. Integument glabrous. Hairs 1-3-P on a distinct tubercle; 1-P long, sin-

gle; 3-P moderately long, single to 5-branched; 14-P single; 0-P, 1,13,14-M, 1,3,13-T

with weak branches, not stellate, restrictor with longer, stronger branches in 13-T.

Abdomen: Integument glabrous. Hairs 1,2,4,11,13-I, 1,2,5,9,13-H, 1,2,5,7,9,13-HI-

VII short to moderately long, varied in branching, never stellate; hair 2 never moved

far cephalad of 1, always within posterior half of segment; 6-I long, at least triple; 7-I

long, single; 6-II long, double or triple; 7-II shorter, multiple; 6-III-V at least moder-

ately long, double or triple; 6-VI single, long. Segment VIII: Comb scales in a patch

of 3-5 irregular rows; individual scale narrow, elongate, broadly rounded apically.

Hair 1-VIII multiple, with short weak branches; 5-VIII moderately long, double or

triple; 1,2-VIII not on tubercle or sclerotized plate except in restrictor. Siphon:

Long, index 13.0-30.0. Integument with minute wartlike spicules. Acus distinct,

attached, with a short blunt ventral and a narrow pointed dorsal projection, except

no dorsal projection in restrictor. Pecten not extending beyond basal 0.25. Hair 1-S

distinctly caudad of last pecten tooth; 4-6 pairs of subventral (1,la-S) and 2 pairs of

subdorsal hairs (2a-S) on siphon; both subdorsal hairs distal in position; hair 2-S dis-

tinct, slightly curved, with or without submedian branch. Anal Segment: Saddle

complete; integument imbricate and faintly spiculose distally; acus not developed;

caudal margin without spines; hair 1-X short to moderately long, branched. Hairs 2,

3-X long, single; ventral brush (4-X) with 5 pairs (6 in restrictor) of branched hairs on

a grid with distinct lateral bar, not attached to saddle; no detached proximal hairs.

Gills varied in length, ventral shorter than or subequal to dorsal.

GENERAL CONSIDERATIONS

TAXONOMIC CHARACTERS. Adults of Anoedioporpa, like those of most other

subgenera of Culex, are very similar in external morphology and ornamentation, and

therefore we have not been able to find many reliable characters for the separation

of the 2 groups. The problem becomes even more difficult at the specific level. The

only reliable characters we found useful in the separation of groups are: (1) presence

or absence of acrostichal bristles on the disc of the mesonotum and (2) length of

male and female palpus. The characters used in the separation of species are rather

tenuous, such as (1) coloration of scales on upper margin of ppn and antealar area

above paratergite and (2) coloration of erect scales on vertex.

The male genitalia, however, show significant and reliable characters at the specific

level such as: (1) nature of the lobes of [X tergite, (2) position and structure of sub-

apical lobe, (3) details of clasper, (4) presence or absence of foliaceous setae on the

subapical lobe, and (5) number and nature of cercal setae and apical spines on the

paraproct.

30 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Like the adults, the immature stages are difficult to diagnose at the specific level,

but reliable characters are present to separate them into groups. In the pupae, the

characters used at the level of groups are: (1) length of 10-C in relation to 11-C, (2)

nature and branching of hair 1-II, (3) length of 6-I,II in relation to 7-I,[I, and (4)

presence or absence of hair 1-[X. The fourth instar larvae show many significant

characters at the group and, sometimes, specific levels. Among the most important

characters used in the separation of groups are: (1) nature of the central tooth of

mental plate, (2) presence or absence of a sclerotized plate on segment VIII, (3)

number of hairs in the ventral brush (4-X), and (4) number of subdorsal hairs (2a-S)

on siphon.

SYSTEMATICS. The following combination of characters of the adults and im-

mature stages will distinguish Anoedioporpa from other subgenera of Culex. In the

adults, the subgenus is characterized by the combination of: (1) only narrow decum-

bent scales on the vertex, (2) short palpus in both sexes (except in restrictor), (3)

general absence of acrostichal bristles on the mesonotum (except in corrigani and re-

strictor), (4) generally yellowish pleural integument, and (5) dark tarsi. In the male

genitalia, Anoedioporpa is distinguished by: (1) shape and position of lobe of IX ter-

gite, (2) development and structure of subapical lobe, (3) details of clasper, and (4)

shape of the lateral plate of phallosome. In the pupae, the subgenus is diagnosed by:

(1) strongly pigmented trumpet, (2) hair 9-VIII long, subequal in length to tergite

VIII, (3) presence of hair 1-IX (absent in restrictor), (4) absence of marginal spicules

on paddle, and (5) absence of both paddle hairs. The most important characters

that separate Anoedioporpa from other subgenera in the larvae are: (1) absence of

head hair 2-C (present in restrictor only), (2) shouldered central tooth on mental

plate (simple in restrictor), (3) 4 pairs of subventral and 2 pairs of subdorsal hairs on

the siphon, (4) apical hook (2-S) of siphon with a small submedian branch, (5) dor-

sal saddle hairs (2,3-X) long, single, and (6) ventral brush usually with 5 pairs of hairs

(6 pairs in restrictor).

We are including in Anoedioporpa Culex restrictor, considered a member of the

subgenus Microculex by Stone, Knight and Starcke (1959:250) and Knight and

Stone (1977:269). On the basis of correlated features of adults and immatures, re-

strictor appears to be closer to Anoedioporpa than to other subgenera. However, re-

strictor is very distinct in certain features, and therefore it is placed in a monotypic

group. It has retained several ancestral attributes not exhibited by members of the

Conservator Group.

In the present revision 12 species are recognized in the subgenus, falling into 2

well defined groups: (1) Conservator Group with 11 species (conservator, canaanen-

sis, damascenoi, browni, bamborum, belemensis, chaguanco, originator, quasiorigina-

tor, luteopleurus and corrigani) and (2) monotypic Restrictor Group.

BIONOMICS. All members of Anoedioporpa are container breeders, and their im-

mature stages have been collected mainly in treeholes and/or bamboo, and occasion-

ally in artificial containers. The adults of all species are primarily sylvan, but some

have been collected in suburban surroundings at low to moderately high elevations.

Field studies in Panama show that immature stages are found near ground level as

well as in the forest canopy (Galindo, Carpenter and Trapido 1951), and apparently

the adults do not show any preference for height. The females of Anoedioporpa are

not reported to bite man and like most of the species of Culex are nocturnal. Some

species exhibit positive phototropism and are collected in large numbers in light traps

(Galindo and Blanton 1955). Females of Culex conservator lay their eggs in rafts on

the surface of water in treeholes (Howard, Dyar and Knab 1915:310), but nothing is

known about the ovipositing behavior of other species.

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 31

DISTRIBUTION (figs. 12,27). Anoedioporpa is primarily Central and South Amer-

ican in its distribution; apparently all species are confined to the mainland with the

exception of conservator and originator. The subgenus has a wide distribution, ex-

tending from Mexico through Central America, Colombia, Venezuela, the Guianas,

northern and eastern parts of Brazil, and northern Argentina. The northern limits of

distribution are found in the states of Jalisco on the Pacific side and San Luis Potosi

on the Atlantic side of Mexico (restrictor). The southern limits of distribution are

found in the departments of Salta and Misiones in northern Argentina (chaguanco).

On the Pacific side of the Andes in South America, Anoedioporpa extends south on-

ly a short distance to the department of Valle del Cauca in Colombia (browni). This

subgenus has not been reported from central or southern Brazil, but this is probably

due to paucity of collections. Further surveys from various critical areas in Brazil

will probably reveal some hitherto unknown forms, and fill some of the lacunae in

our present understanding of the distribution of the various species of Anoedioporpa.

Of all the known species of Anoedioporpa, Culex conservator appears to be the

dominant and most widespread one in terms of range and number of individuals. It

is sympatric with all the other species of the subgenus with the possible exception of

canaanensis and chaguanco. It is the only species occupying almost the entire range

of the distribution of Anoedioporpa. Four species (browni, corrigani, originator, re-

strictor) are known from numerous localities in Central and/or northern South Amer-

ica. Only 2 species are known from any of the islands of the West Indies: conserva-

tor from Trinidad and Tobago, and originator from Trinidad, Grenada and Martin-

ique. All the remaining 7 species are each known from only one or 2 localities in

South America.

KEYS TO SPECIES OF ANOEDIOPORPA

ADULTS

l. Mesonotun: with at least a few acrostichal bristles 4. 625.55. . .2

Mesonotum with no indication of acrostichal bristles . . . .... .3

2(1). Acrostichal bristles extending from anterior promontory to prescutellar

area, strong, distinct; lower mep witha bristle . . . . 12. restrictor

Acrostichal bristles represented by 3,4 shorter setae near prescutellar area;

lower mien Dale 6 oe Ge a oa A, corricant

3(1). Antealar area above paratergite with black scales . . . 10. luteopleurus

Antealar area above paratergite with scattered white scales. . . .. .4

4(3). Frect scales on vertex white tocreamy .... . .. . .J. conservator

Erect sczes On Vertes Diwan ga 8

5(4). Scalesonupperppn brown. . . . . 2. canaanensis; 8. originator

Scales on upper ppn white . . 3. damascenoi: 4. browni; 5. bamborum;

HD eae y 6. belemensis; 7. chaguanco,; 9. quasioriginator

32

4(3).

5(3).

6(1).

7(6).

8(6).

9(8).

10(8).

11(10).

Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

MALE GENITALIA

Inner surface of sidepiece below proximal division of subapical lobe with a

large densely setose area .... , es

Inner surface of sidepiece be:ow Beal Son ae apical lobe at

RIOSE Wilh 2 lew SCattcred. 20tke i hi ek SS) Des

Proximal division of subapical lobe densely setose on nee surface to

basal 0.5 in addition to asetose areaonsidepiece . . 2. canaanensis

Proximal division of subapical lobe not setose on undersurface. . . .3

Setose area of sidepiece below proximal division of subapical lobe flat-

bene oF ee, oe

Setose area of gdeniecs below peoniinal @ivision of subapical fobs | in the

Porn evr Os ee ee

Distal division of subapical lobe with 2 enlarged leaves near middle of

Stel 2k ' . .d. conservator

Distal division of subapical lobe with only : enlarged leaf near middle of

a ee, Cemascenol

Distal division of subapical lobe with a lanceolate seta midway on stem;

setose area of sidepiece below Ace division (boss) without any en-

largec setae 2... . . 4. browni

Distal division of sub anival lobe with an oflsicen eat as on stem;

setose area of sidepiece below proximal division (boss) with a few spec-

feed colnieed Setge eS Domborum

Proctiser Wib a densely setose subapical area. . . . ww ee ye el

Proctiger with a few very short, weak cercalsetac . ........8

Stem of proximal division of subapical lobe with a patch of short setae ex-

tending from base to middle... . . &. originator

Stem of proximal division of subapical lobe Shout any setae

ee a 9. guosioriginator

Distal division of subapical lobe short, wider than Sin without a narrow

stem |. oe

Distal division of subapical lobe longer than wide, with a narrow stem . 10

Distal division of lobe with only specialized setae; cercal sclerite of procti-

ger broad, oe bent mesad; clasper stout, about 0.5 length of

sidepiece |” . . . LL. corrigani

Distal division of lobe, in addition to specialized setae, with 4 apical leaves;

cercal sclerite of proctiger digitiform, bent laterad; clasper narrow, slen-

der, about0,60-10. 65 lenetn di sidepiece . . . cw Ee. restrictor

Distal division of subapical lobe with 5 enlarged leaves; ninth tergite lobes

INCISUINCL «64: . . 10. luteopleurus

Distal division of subapical lobe with { or i) enlarged leaves: ninth tergite

Popes (istic, MGNR he 68 Oa a ee ce ae a LT

Distal division of subapical lobe with 1 enlarged leaf midway on stem

ee 7. chaguanco

Distal dicen ar subapical lobe with 2 onlersed leaves midway on stem

. 6. belemensis

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 33

PUPAE

(6. belemensis, 9. quasioriginator and 10. luteopleurus unknown;

2. canaanensis not included)

Restrictor Group

1. Hair 10-C strongly developed, double, longer than 11-C . . 12. restrictor

Hair 10-C weakly developed, multiple, shorter than or subequal to 11-C .2

Conservator Group

Zt4). Hair 1-VII at least double; 4-VI usually aii nee of 5- VI

; 11. corrigani

Han 1-VII ‘single: AVI cephalolaterad of 5-V1 . eS hea ta Sante a gee

32). Hair 2-II extremely small, branches indistinct at 1OOX . . ... . 4.44

Hair 2-II small, branches distinct av TOOX gs es Pe Ene tee

4(3). Apex of paddle slightly produced; trumpet hohe ae index about

moe eo ie ie ; . . 3. bamborum

Apex of os rounded: trumpet shorter, index Jess than - 0 on

. 3. damascenoi

StS): Hair 5-II] Ae tate (4-7) CI eh I aa es) he CGM NOPVaLar

Flair STI Sinsie Or GOUUIe 8 ee ee ee re

6(5). Hair 1-Vi usaally doubic., 62 oe ae 7 eeeanco

Hair i-VIisingle .3 ooo a a ae OW ©. Crigineror

LARVAE

(3. damascenoi, 6. belemensis, 9. quasioriginator and 10. luteopleurus

unknown; 2. canaanensis not included)

Restrictor Group

t; Ventral brush (4-X) with 6 pairs of hairs; central tooth of mental plate not

shouldered; hair 2-VIII on an oval sclerotized plate. . . 12. restrictor

Ventral brush (4-X) with 5 pairs of hairs; central tooth of mental plate

shouldered hair 2-VIIl not ona sclerotized plate: a ee

Conservator Group

ath): Hair 1-V short, branched, barely extending to apex of segment .

sae 11. corrigani

Hair 1-V esteem long, single. ciendine Ravend apex of succeeding

SOCTIOIE sari Pica) lett dd On eee renntie Meee ER ici CewN dom aa nt ge

a2), Siphon extremely long, index more than 25.0. . . . . . 5. bamborum

SIN OM SNOTICT ee eae aA De er ee ee eee ae

4(3). Gills long, dorsal at least 2.0 length of saddle . . .... . 4 browni

Gills shorter, dorsal subequal tolength of saddle. oe we el SS

5(4). Siphon with 5 paris of subventral hairs(1,la-S) . . . . . 7. chaguanco

Siphon wih + pais OF supventral Naits(ivieay ee eee 6

34 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

6(5). Hair 5-C usually triple (2-5); hair 3-P single or double; 7-P usually double,

rarely single . . . .d. conservator

Hair 5-C at least 4- branched (4-7): nae 3- P P usualy tripe (2-5); hair 7-P

usually single, rarely double ... . . &. originator

Conservator Group

FEMALES. Usually small, inornate species. Head: Erect scales forked apically,

creamy to brown, Proboscis longer than forefemur, entirely dark scaled. Palpus

short, about 0.15 length of proboscis, 4-segmented, segment 4 about 2.0 of segment

3. Antenna shorter than proboscis. Thorax: Mesonotal scales tan to brown. Acro-

stichal bristles usually absent (2,3 short setae present near prescutellar space in corri-

gani). Pleural integument yellowish to greenish; scaling restricted to upper ppn and

stp; lower mep usually with 1 strong bristle. Abdomen: Tergites IJ-VII with basolat-

eral light patches, predominantly dark scaled; sternites predominantly creamy, dark

scaled distally.

MALES. Similar to female in coloration. Palpus short, about 0.15 length of pro-

boscis, 4-segmented; segment 4 about 1.6 of segment 3.

MALE GENITALIA. Segment IX: Tergite lobe moundlike or indistinct, with var-

ied number of short to moderately long setae. Sidepiece: Roughly conical, length

1.8-2.0 of greatest width. Lobe situated at 0.7, directed caudomesad, divided into

proximal and distal divisions; proximal division with 2 distinct rods; distal division

with varied number of setae; leaf present or absent on stem. Clasper: About 0.6-0.7

length of sidepiece, curved inwards, distal portion usually with transverse ridges on

external margin. Phallosome: Lateral plate with apical process broadly rounded or

truncate from dorsal aspect; sternal spine short, pointed. Proctiger: Paraproct with

6-13 apical teeth; cercal setae variable in number.

PUPAE. Cephalothorax: Hair 5-C short, weakly developed, subequal in length to

4-C; hair 10-C weakly developed, at least double, distinctly shorter than 11-C. Abdo-

men: Hair 1-I] small, weakly branched, not resembling float hair (1-I); hair 1-III-VII

moderately long, varied in branching; 6-I,II moderately long, subequal to 7-II, III; hair

5-IV,V long, single, either barely reaching or extending beyond apex of succeeding

segment; 2-II-VI short, subequal in length to 9-II-VI; hair 9-VII strong, at least 2-

branched (2-5); hair 9-VIII with 3-8 strong branches, slightly shorter than segment

VIII; hair 1-[X present, small. Posterior margin of sternum VIII mesad of hair 9-VIII

distinctly lobed. Paddle: Elongate, lightly pigmented; hairs 1,2-P absent.

FOURTH INSTAR LARVAE. Head: Hair 5-C at least 3-branched; 6-C single; 7-C

with 6-10 and 11-C with 3,4 branches. Central tooth of mental plate distinctly

shouldered. Antenna: Hair 1-A inserted about 0.75 from base. Thorax: Hairs 5,6-P

single, 7-P single or double, all strongly developed. Abdomen: Hair 6-III-V moder-

ately long, but less than 0.5 length of 6-I,I]; hair 9-II-VI short, single and spinelike.

Segment VIII: Hair 2-VIII not on a sclerotized plate. Siphon: Subventral hairs (1,

la-S) 4,5 pairs; subdorsal hairs (2a-S) only 2 pairs, both located in distal half of si-

phon. Anal Segment: Ventral brush (4-X) with 5 pairs of hairs on a grid. Gills var-

ied in length, ventral usually shorter than dorsal.

DISCUSSION. The Conservator Group includes all members of the subgenus ex-

cept restrictor, and is strongly differentiated from the Restrictor Group in the adults

and immature stages. In the adults, both sexes have a short palpus, and are devoid of

acrostichal bristles on the disc with the exception of corrigani. The pupae are readily

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 35

recognized by the weakly developed hair 10-C which is distinctly smaller than or sub-

equal to 11-C. The larvae are immediately separated from all others by the presence

of a shouldered central tooth on the mental plate and 5 pairs of hairs in the ventral

brush.

Of all the included species, conservator appears to be the dominant species with an

extensive distribution from southern Mexico through Central America to central

(Anapolis, Goias) and eastern (Alagoas) Brazil, including the islands of Trinidad and

Tobago. Culex originator has a more restricted distribution in northern South Amer-

ica (Venezuela, the Guianas, NE Brazil), and is the only other species known from

Trinidad; it is the only species reported from any of the islands of the Lesser Antilles

(Grenada and Martinique). Culex corrigani is found only in Central America, along

the Caribbean side from Nicaragua to Panama, and on the Pacific side in Panama.

Culex browni is known from eastern Panama (Darien) to the Pacific side of the An-

des in Colombia (Valle del Cauca) and the Atlantic side of the Andes in Ecuador

(Napo); browni is the only species reported from the Pacific side of the Andes in

South America, and the only species reported from Ecuador. Culex belemensis and

damascenoi are both known only from 2 localities each in French Guiana and north-

eastern Brazil. Culex chaguanco is the most southern species, being known from 2

localities in Argentina (Salta and Misiones). The other 4 species are known only

from their type localities: bamborum from Colombia (Meta), canaanensis from Brazil

(Espirito Santo), and Juteopleurus and quasioriginator both from the state of Para in

Brazil.

1. Culex (Anoedioporpa) conservator Dyar & Knab

Figs, 12,13,14,15

1906. Culex conservator Dyar & Knab 1906:221-222. *TYPE: Lectotype larval skin (13.12)

with associated pupal skin and male, St. Joseph (St. George), Trinidad, 15 June 1905, A.

Busck [USNM; designation of Stone and Knight 1957:46].

1906. Culex divisior Dyar & Knab 1906:222-223. *TYPE: Lectotype larval skin (B15-6) with

associated pupal skin and male, Trinidad, 9 Nov 1905, F.W. Urich [USNM; designation of

Stone and Knight 1957:47-48]. Synonymy with conservator by Dyar 1922:95.

1922. Culex (Isostomyia) bifoliata Dyar 1922a:94-96. *TYPE: Lectotype male (164.4) with

genitalia slide, Miraflores, Canal Zone, Panama, 15 Dec 1921, J.B. Shropshire [USNM, 25

254; designation of Stone and Knight 1957:44]. Synonymy with conservator by Dyar

1928:345.

1923. Culex paganus Evans 1923a:104-106. *TYPE: Lectotype male (D/409) with genitalia

on 3 slides, from village, Estado Aragua, Venezuela, 2 Aug 1922, M. Nunez Tovar [BM-

LIVER; designation of Belkin 1968b:18-19]. NEW SYNONYMY.

Culex (Anoedioporpa) conservator of Bonne and Bonne-Wepster (1925:188,260,261-263); Belkin

(1968b: 11-12); Bertram (1971:745,751); Xavier and Mattos (1975:246); Knight and Stone

(1977:195); Heinemann and Belkin (1977a:283; 1977b:415,427,442; 1977c:528; 1978a:183,

194; 1978b:393 437; 1979:94).

Culex (Isostomyia) conservator of Dyar (1918:103; 1922:95,96); Gordon and Evans (1922:325,

326,327); Edwards (1932:218); Komp (1936:327); Chagas, da Cunha et al. (1938:194); Lane

(1939:73-74); Anduze (1941:16; 1943:196; 1947:359); Lane and Whitman (1943:397,398);

Rozeboom and Komp (1948:430); Arnett (1948:189-190; 1950:107,111,112); Galindo, Car-

penter and Trapido (1951:102,104,105,107,108,110,111,112,113,126); Horsfall (1955:548).

Culex (Aedinus) conservator of Dyar (1923b:189); Bonne-Wepster and Bonne (1923:125); Stone,

Knight and Starcke (1959:281, in part); Belkin, Schick and Heinemann (1965:70); Cova Garcia,

Sutil and Rausseo (1966a:28,107, in part; 1966b:40-41, 83-84, 342, in part).

Culex (Melanoconion) conservator of Dyar (1928:345-346); Lima (1930:255); Shannon (193 1:8,

23); Komp (1935:10); Chagas, da Cunha et al. (1937:385,387,388,389).

36 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Culex (Tinolestes) conservator of Lane (1953:391-392); Duret and Damasceno (1955:394,395);

Galindo and Blanton (1955:70); Fauran (1961b:48); Cerqueira (1961:131).

Culex (Anoedioporpa) bifoliatus of Dyar (1923a:177); Bonne and Bonne-Wepster (1925:260,263).

Culex (Isostomyia) bifoliata of Gordon and Evans (1922:327); Evans (1923:105,106); Stone and

Knight (1957:44).

Culex (Aedinus) bifoliatus of Dyar (1923b:189).

Culex (Melanoconion) bifoliatus of Dyar (1925:158,161,169).

Culex (Anoedioporpa) paganus of Bonne and Bonne-Wepster (1925:260,265); Belkin (1968:12,18,

19); Xavier and Mattos (1975:246).

Culex (Isostomyia) paganus of Lane and Whitman (1943:397).

Culex (Aedinus) paganus of Stone, Knight and Starcke (1959:282); Belkin, Schick and Heinemann

(1965:75); Cova Garcia, Sutil and Rausseo (1966a:28; 1966b:41,84,343).

Culex (Tinolestes) paganus of Lane (1953:394-395); Duret and Damasceno (1955 :394,395 397,

407,408); Fauran (1961b:48).

Culex conservator of Dyar (1906:18); Urich (1913:529); Howard, Dyar and Knab (1915:222,229,

308-310); Soper, Penna et al. (1933:574); Kumm and Novis (1937:511); Kumm, Komp and

Ruiz (1940:403); Galindo, Carpenter and Trapido (1955:159-161); Stone and Knight (1957:

46); Mattos and Xavier (1965:281); Forattini, Rabello and Cotrim (1970:37).

Culex paganus of Forattini, Rabello and Cotrim (1970:468).

FEMALE (fig. 13). Wing: 2.4-2.5 mm. Proboscis: 1.8-1.9 mm. Forefemur: 1.5-

1.7mm. Abdomen: 1.60-1.65 mm. As described for the subgenus and group, with

the following additional features. Head: Decumbent scales on vertex narrow,

creamy; sides and venter with broad white scales. Erect scales white to creamy. Pal-

pus short, about 0.16 length of proboscis. Thorax: Scales on mesonotum predomi-

nantly narrow, auburn except for white scales on anterior promontory and along an-

terior part of lateral prescutal area. Antealar area above paratergite with moderately

broad white scales. Pleural integument usually yellowish. Upper margin of ppn with

narrow white scales. Lower mep with a strong bristle. Abdomen: Scales on tergites

predominantly brown except for basolateral light areas. Sternites with creamy to

whitish scales.

FEMALE GENITALIA (fig. 13). As described and figured for Conservator Group.

MALE (fig. 13). Wing: 2.4-2.5 mm. Proboscis: 2.0 mm. Forefemur: 1.6 mm.

Similar to female in general coloration. Palpus about 0.15 length of proboscis.

MALE GENITALIA (fig. 14). As figured; diagnostic characters as in the key. Seg-

ment IX: Tergite lobes slight, widely separated, each with 4-6 short weak setae.

Sidepiece: Roughly conical; length about 2.0 of greatest width; lateral surface with

a few longer bristles and scales; sternal surface with shorter setae; tergal surface later-

ad of proximal division of subapical lobe with a triangular, densely setose area.

Lobe: Proximal division with a moderately long stem bearing 2 rods with hooked

tips, upper rod inserted at apex, lower slightly basad. Distal division distinct, with 1

subbasal (sometimes basal) and 1 submedian leaf on stem; apex with a long filament

with recurved expanded tip on upper surface, 2 broad filaments with expanded apex-

es on upper angle, between these a shorter lanceolate seta. Clasper: About 0.60-0.65

length of sidepiece, sharply bent mesad at distal 0.4; distal 0.3 of outer surface with

many closely set transverse ridges; 1 submedian and 1 subapical seta on inner surface.

Lateral Plate: Basal hook sclerotized, strongly curved, bearing a few denticles; apical

process distinct, moderately long and expanded widely at tip; sternal spine short,

pointed. Proctiger: Apex of cercal sclerite narrow, digitiform, directed caudomesad;

cercal setae 1 or 2. Paraproct with a row of 6,7 blunt apical teeth.

PUPA (fig. 14). Abdomen: 2.3-2.5 mm. Trumpet: 0.35-0.40 mm; index about

5.0-5.5. Paddle: 0.50-0.55 mm. As figured; diagnostic characters as in the key.

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 37

Cephalothorax: Integument lightly pigmented, wing case slightly darker. Hairs 1,3-

C single; 2-C usually double (2,3); hairs 4,5,7-C double; 6-C single; 8,9-C single; 10-C

at least 3-branched (3-6). Trumpet: Very strongly pigmented and distinctly con-

trasting with cephalothorax. Abdomen: Integument lightly pigmented. Hair 1-II

short, weakly branched, easily distinguishable at 100X; hair 7-II single; 4-VI short,

weakly branched, cephalolaterad of 5-VI; hair 1-VI,VII usually single; 9-VII with 4,5

branches; 9-VIII usually 6-branched (3-7). Paddle: Lightly pigmented, longer than

wide, about 2.0 of segment VIII. Male genital lobe extending to 0.4 and female to

0.25 of paddle.

FOURTH INSTAR LARVA (fig. 15). Head: 0.80-0.85 mm. Siphon: 1.6-1.7 mm;

index 15.0-18.0. Anal Saddle: 0.3 mm. As figured; diagnostic characters as in key.

Head: Width about 1.2 of length. Hair 4-C usually single; 5-C usually with 3 branch-

es (2-5); hair 9-C usually with 6 branches (4-8); hair 14-C single; 15-C usually with 3

branches (3,4), short, not extending to base of mental plate. Mental plate with a

strong medium shouldered tooth and 6,7 distinct teeth on either side. Antenna:

Length about 0.5 of head, distinctly spiculose to hair 1-A; all hairs single except 1-A

(20,16-24). Thorax: Hair 0-P short, weak, with dendritic branches; 1-3-P long, sin-

gle; 4-P double; 5-7-P single; 14-P small, single; 4-M double. Abdomen: Hairs 3,6-I

usually with 3 branches (3,4); 7-I single; 6-II moderately long, usually triple; 6-III-V

shorter than segment length, double; 1-V long, single. Segment VIII: Comb scales in

a patch of 4,5 irregular rows, about 53-57 in number; individual scale with spatulate

fringed apex. Siphon: Integument moderately pigmented, with a darker basal ring.

Subventral hairs (1,1a-S) 4 pairs, progressively smaller distad; subdorsal hairs (2a-S)

2 pairs, short, located in distal third of siphon. Pecten extending barely to proximal

0.25; individual tooth long, pointed apically, without lateral denticles. Anal Seg-

ment: Integument moderately pigmented, slightly imbricate. Hair 1-X short, weakly

branched; 2,3-X long, single. Gills short; dorsal 0.65-0.90 length of saddle; ventral

slightly shorter than dorsal.

SYSTEMATICS. Culex conservator, the type species of Anoedioporpa, can be

distinguished from other members of the subgenus in the adults by the presence of

white to creamy erect scales on vertex; in the male genitalia by the combination of:

(1) presence of a subbasal and a submedian leaf on distial division of subapical lobe

and (2) large, triangular, densely setose area on the inner surface of sidepiece below

subapical lobe; in the pupa by the combination of: (1) hair 1-VII single, (2) shorter

trumpet with index of 5.0-5.5, and (3) 5-III at least triple (3-7); in the larva by the

combination of: (1) hair 5-C usually triple (2-5), (2) hair 3-P single or double, and (3)

hair 7-P usually double, rarely single.

Culex conservator is the most abundant and widespread species of Anoedioporpa

and constitutes the dominant species of the subgenus in terms of range and number

of individuals. It is sympatric with all species except canaanensis and chaguanco.

This dominance indicates that conservator may possibly be one of the more modern

species of Anoedioporpa in relation to the complex distribution pattern and geologi-

cal history of the area occupied by this species. However, it may represent the an-

cestral stock from which several segregates have developed in the Conservator Group.

Very little variation in adult morphology is apparent throughout the known wide

distribution of conservator, and no significant population differentiation in the im-

mature stages has been detected in the relatively large sample examined.

We have examined the types of divisior Dyar & Knab 1906 and bifoliata Dyar

1922, and they are undoubtedly conspecific with conservator; hence their synonymy

(Dyar 1922:95; 1928:345) is justified. We are synonymizing paganus Evans 1923

38 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

with conservator. Although we have not studied the types in detail, the description

of paganus matches in all pertinent details the diagnostic features of conservator. On

the other hand, surukumensis Anduze 1941 which has been synonymized with con-

servator in the past appears to be conspecific with originator (see).

BIONOMICS. The immature stages of Culex conservator have been found in tree-

holes and bamboo in sylvan areas. They are frequently found breeding in open tree-

holes in suburban areas away from forests. This species is particularly addicted to

bamboo traps and was found almost equally as often in traps situated near the

ground as in the,forest canopy (Galindo, Carpenter and Trapido 1951:127). Ap-

parently the eggs are laid in a raft floating on the surface of the water (Howard, Dyar

and Knab 1915:310). The females are innocuous and do not bite humans (Shannon

1931: 23),

DISTRIBUTION (fig. 12). Culex conservator is the most widely distributed spe-

cies in the subgenus, extending from southern Mexico (Tabasco) to at least northern

Brazil (Amazonas, Para). It is reported to occur as far south as central (Anapolis in

Goias) and eastern (Alagoas, Ceara, Piaui) Brazil. It is found throughout Central

America ( all countries except El Salvador and Nicaragua), in the islands of Trinidad

and Tobago, and in parts of northern South America, including the Guianas, Colom-

bia (Meta) and Venezuela (Aragua, Carabobo, Monagas). It has been collected from

near sea level to 740 m above sea level.

Material examined: 1636 specimens; 236 males, 207 females, 315 pupae, 878 lar-

vae; 227 individual rearings (142 larval, 62 pupal, 23 incomplete).

BELIZE. Cayo: Augustine, Mountain Pine Ridge, 500-1000 m, 10 Aug 1967, P. Williams, 1M

(BH 490) [UCLA].

BRAZIL. Alagoas: Maceio (Chagas, da Cunha et al. 1937:388). Amazonas: Rio Paruary, det.

J. Lane [FH, 5933-35]. Maues, Mgen, det. J. Lane [FH, 6123; Forattini, Rabello and Cotrim 1970:

37]. Ceara: Crato (Chagas, da Cunha et al. 1937:387). Goias: Anapolis (Mattos and Xavier 1965:

281). Para: Belem, Conceicao do Araguaia; and Utinga (as paganus) [INER, Xavier and Mattos

1975:246]. Curralinho, Rio Camucu, 1936, H.W. Kumm, 1M (K74) [USNM] ; Rio Massaranduba

[INER, Xavier and Mattos 1975:246]. Ilha de Marajo (Kumm and Novis 1938:511). Piaui: Tere-

sina (Thorezina) (Chagas, da Cunha et al. 1937:385). Locality unspecified: H.W. Kumm, |Mgen

(9043) [USNM].

COLOMBIA. Meta: Villavicencio, 1944, M. Bates, 3M (CV 61,61-1,61-8), 1F (61-1); Bosque

Ocoa, 19 Aug 1948, M. Bates, 1Mgen (CV 1009-101) [UCLA].

COSTA RICA. Puntarenas: Golfito, H.W. Kumm, 1Mgen (509) [USNM]. Osa Peninsula, Rin-

con, near sea level, 29 June 1963, C.L. Hogue, 1 IpM (CR 130-201), 3 lpF (131-203,-205,-206), 3

pM (130-601-603), 3M, 7F, 9P, 83L (130), 2 lpM (CR 131-201,-204), 5M, 8F, 12P, 168L (131), 1

M (CR 138) [UCLA].

FRENCH GUIANA. Guyane: Organabo, 10 m, 16 Feb 1969, J. Clastrier, 2L (FGC 3914) [UC

LA].

GUATEMALA. Izabal: Quirigua ruins, 70 m, 4 Aug 1964, T. and J. Zavortink and W. Almen-

gor, 1 lpM (GUA 90-101), 1 IpF (90-100) [UCLA].

HONDURAS. Atlantida: Lancetilla, 50m, 19 Aug 1964, A. Quinonez, | lpF (HON 54-20), 2

pF (54-101,-203), 2P (54), 2 pM (HON 55-100,-101), 2 pF (55-102,-103), 2P, 2L (55) [UCLA].

MEXICO. Tabasco: Comalcalco, 10 m, 12 July 1970, K. and D. Schroeder, 1 IP (MEX 553-10),

1 pM (MEX 554-101), 2 pF (554-100,-102), 12L (554), 1 IpF (MEX 555-20), 6L (555) [UCLA].

PANAMA. Bocas del Toro: Almirante, 10 m, 27 Apr 1963, A. Quinonez, 1 lpF (PA 259-104),

4L (259); 14,15 Apr 64, A. Quinonez, 1M (PA 666) [UCLA]. Canal Zone: Arraijan, 9 Oct 1950,

S.J. Carpenter, 2L; 20 Nov 1950, S.J. Carpenter, 8L; 30 Oct 1950, S.J. Carpenter, 1L [UCLA].

Balboa, 28 Dec 1921, J.B. Shropshire, 2F [USNM] ; 18 Feb 1943, 4F (KO 37-3) [UCLA]. Barro

Colorado Is., 25-170 m, 7 May 1943, W.H.W. Komp, 1M (KO 37-23), 3F (KO 37-18, 37-30, 41-43);

12 May 1943, 1M (KO 41-12); 31 May 1943, 1F (KO 37-31); 20 Aug 1944, K. Frick, 3M (ASM 80-

1, 90-1); 15 May 1945, 6L (52-89, 53-11); 26 July 1946, 1M, 1F (KO 31-32); 4 Dec 1965, A.

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 39

Quinonez, 1 lpM (PA 861-12), 1 lpF (861-30) [UCLA] ; Miller Tower area, 15 May 1945, 1M, 1F

[USNM] ; Pearson Trail, 7 May 1943, 1M (KO 41-17), 2F (KO 41-26,-39); S.M. Trail, 7 May 1943,

1F (KO 41-37) [UCLA]. Chiva Chiva, 30 m, 10 Nov 1965, A. Quinonez, 3 IpM (PA 764-10-12), 2

IpF (764-13,-14), 2 pF (764-100,-101), 1P (764); 11 Nov 1965, A. Quinonez, 1 lpM (PA 768-30),

1F (768), 4 lpM (PA 769-11,-13,-15,-16), 3 lpF (769-10,-12,-14), 1F, 1P, 17L (769) [UCLA].

Chepo Road, 22 Oct 1939, 6M [USNM]. Corozal, 26 Apr 1942, 4M, 4F; Corozal Lab, 30 Aug

1943, 2M, 3F; Corozal Road, East, 27 Apr 1942,3M, 2F [USNM]. Fort Sherman, 26 Aug 1949,

1 IpF (918-12); 10 Feb 1950, 1 lpF (2645-2); 24 Feb 1950, 1 lpF (2755-7); 26 Oct 1950, S.J. Car-

penter, 16L; Sweet Water Reservoir, 80 m, 10 Nov 1964, A. Quinonez, 1 lpF (PA 732-20), 4L

(732) [UCLA]. Gatun, 25 Aug 1926, D.P. Curry, 2M, 2F; 25 July 1928, 2L [USNM]. Madden

Forest Preserve, less than 200 m, 29 Nov 1965, 2 IpM (PA 839-11,-15), 3 lpF (839-10,-12,-13), 1 IP

(839-14), 5L (839) [UCLA]. Mandingo, 22 Dec 1921, J.B. Shropshire, 2M (25284), 1F [type ser-

ies of bifoliata, USNM]|. Margarita, 10 m, 8 Oct 1964, A. Quinonez, 1 pM (PA 716) [UCLA].

Miraflores, 28 Dec 1921, J.B. Shropshire, 1F [USNM]. Mojinga Swamp, 29 June 1922, 1Mgen; 2

Mgen [USNM];5 m, 13 Oct 1964, A. Quinonez, 1 pM (PA 724-100) [UCLA]. Nuevo Emperador,

road near, 100 m, 23 Nov 1965, A. Quinonez, 2 IpM (PA 832-10,-11A), 1M, 1P (832) [UCLA].

Old Gaillard Hwy, 31 Oct 1941, 1M; 14 Sept 1941, 1F; 1 Sept 1941, IM, 2F [USNM]. Summit,

25 May 1945, 1L [USNM]. Venado Beach, 21 Oct 1939, 7L [USNM]. Locality unspecified, 31

Dec 1921, J.B. Shropshire, 2M [USNM]. Colon: Corredor de Colon, near sea level, 24 Sept 1964,

A. Quinonez, 11L (PA 713) [UCLA]. Darien: Rio Chucunaque, 16 Feb 1958, GML, 4M (GG 1-

146); mouth of Rio Tuquesa, 10 m, 17 Feb 1958, GML, 4M, 2F (GG 1-156) [UCLA]. El Real,

Piriaque, near sea level, 13 Jan 1964, A. Quinonez, 2L (PA 622) [UCLA]. Jaque, Rio Jaque, near

sea level, 19 Dec 1963, A. Quinonez, 1 lpM (PA 612-107), 2 pF (612-119), 1 IP (612-118), 128L

(612) [UCLA]. Morti, Morti Hydro, 100 m, 29 Nov 1966, O.G.W. Berlin, 1 pM (PA 958-100)

[UCLA]. Paya Camp, 50 m, 3 July 1958, GML, 1 pF (GG 1-115) [UCLA]. Santa Fe, 20 m, 10

Dec 1966, O.G.W. Berlin and M. Mena, | pM (PA 997-101) [UCLA]. Tacaracuna, El Salto, 9 Sept

1958, GML, 1 lpM (GG 118-103), 1 IpF (118-101); Rio Tacarcuna valley, 600 m, 5 July 1963, A.

Quinonez, 2 lpF (PA 437-101,-103), 1 pF (437-102), 1L (437) [UCLA]. Rio Tuira, mouth of Rio

Paya, 50 m, 3 Mar 1958, GML, 2M (GG 68) [UCLA]. Panama: El Libano, near sea level, 26 Nov

1963, A. Quinonez, 5 IlpM (PA 567-110,-115,-117,-119,-120), 4 lpF (567-112,-116,-121,-124), 22L

[UCLA]. Juan Mina, 40 m, 18 Jan 1963, A. Quinonez, 2 lpF (PA 8-101,-102), 1L (8) [UCLA].

La Chorrera, 17 Oct 1944, Adams, 4M, 2F, 6L (ASM 210-2) [UCLA]. Panama Viejo, 17 Nov

1944, R. Arnett and K. Frick, 1F (ASM 290-1) [UCLA]. Pacora, 27 July 1950, S.J. Carpenter, 1

L; 29 Aug 1950, S.J. Carpenter, 5L [UCLA]. Locality unspecified: 2 Nov 1934, L.E. Rozeboom,

1L [USNM]. No data: 1M (KO H-13-12), 4L (KO 5-366,-402,-411) [UCLA] ; M, det. Galindo

[FH, 9012; Forattini, Rabello and Cotrim 1970:37].

SURINAME. Locality unspecified: J. Bonne-Wepster, 2M (BB 310,2a), 2F (BB 311,2e)

[USNM].

TOBAGO. St. George: Caledonia, 290 m, 17 Nov 1965, T.H.G. Aitken, R. Martinez and A.

Guerra, 1 IP (TOB 39-20), 1 lpM (TOB 42-32), 1 IpF (42-31), 5L (42) [UCLA]. St. Patrick: Buc-

coo, 25 m, 19 Nov 1965, R. Martinez and A. Guerra, 7L (TOB 61) [UCLA]. Scarborough, Orange

Hill, 150 m, 27-28 Nov 1965, R. Martinez and A. Guerra, 1M (TOB 126); 30 Nov 1965, R. Marti-

nez and A. Guerra, 1 IpM (TOB 138-12), 2 IpF (138-10,-11), 1 pF (138-101), 1 lpF (TOB 139-10),

2P, 4L (139) [UCLA]. Locality unspecified: July 1905, A. Busck, 1M (690723-2) [USNM].

TRINIDAD. Nariva: Archers Estate, 50m, 5 Nov 1964, A. Guerra, 1 IpM (TR 812-113) [UCL

A]. Nariva Swamp, Bush Bush Forest, near sea level, 26 Feb 1964, TRVL, 1L (TR 100) [UCLA].

Tabaquite, Charuma Forest, 50-150 m, 27 Aug 1964, A. Guerra, 2 pM (TR 637-108,-109), 2L

(637); 8 Oct 1964, A. Guerra, 3L (TR 752) [UCLA]. St. Andrew: Cumaca, 200 m, 15 May 1964,

A. Guerra, 1L (TR 407-127) [UCLA]. Vega de Oropouche, Esperanza Estate, 11 Nov 1960, 1M

[USNM]. St. George: Aripo Valley, 150 m, 25 Feb 1965, F. Powdhar, 3L (TR 1015) [UCLA].

Monos Island, 50 m, 17 May 1964, R.L. Manuel, 1 IpM (TR 410-103), 2 lpF (410-108,-109), 2 pM

(410-184,-200), 1 pF (410-185), 6L (410) [UCLA]. Piarco, Centeno Propagating Station, 10 m,

21 Jan 1965, A. Guerra, 1 pM (TR 958-130), 8L (958) [UCLA]. St. Joseph, 15 June 1905, A.

Busck, 1 IpF (13-3), 5M (13-1,-5,-6,-7,-10), 2Mgen (13-2,-7) [type series, USNM] ; 3M (34-2), 2M,

1F [USNM] ; La Baja Rd., 30 m, 9 Sept 1965, A. Guerra, 1 IpM (TR 1390-10), 1 IpF (1390-11),

40 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

1pM (1390-100) [UCLA]. Verdant Vale, 300 m, 11 Mar 1965, A. Guerra, 8L (TR 1037); 200 m,

29 Jan 1966, A. Guerra, 4L (TR 1448) [UCLA]. U.S. Naval Base, males, det. Heredia [FH, 10926-

29; Forattini, Rabello and Cotrim 1970:37]. Locality unspecified: 9 Nov 1905, F. Urich, 1 lpM

(B15-3), 1 pM (B15-10), 3M (B15-1,-7,-8), 4F (B15-2,-4,-5,-9), 1F [type series of divisior, USNM].

VENEZUELA. Aragua: Cata, near sea level, 15 Aug 1969, J. Pulido, 2 pM (VZ 353-103,-104),

5 pF (353-100-102,-105,-106), 1M, 1P, SL (353), 1 IpM (VZ 354-10), 3L (354); 21 Aug 1969, J.

Clavijo and J. Valencia, 15L (VZ 388) [UCLA]. Choroni, Natl. Rt. 2 between Maracay and Chor-

oni, 550m, 15 Aug 1969, J. Valencia, 2 lpF (VZ 358-20,-21), 3L (358) [UCLA]. Guamitas, 740

m, 15 July 1969, J. Bergland and T. Zavortink, 2 lpF (VZ 203-90,-91) [UCLA]. Maracay, M, F

[FH, 8936-37; Forattini, Rabello and Cotrim 1970:37]; 13 Sept 1926, M. Nunez Tovar, 2M; 10

Oct 1926, M. Nunez Tovar, 1M, 2F; 13 Oct 1926, M. Nunez Tovar, 5F; 14 Nov 1926, M. Nunez

Tovar, 8M, 5F; 3 Dec 1926, M. Nunez Tovar, 5M, 6F; M. Nunez Tovar, 1M [USNM]; L.E. Roze-

boom, 9M, 6F (VZR 256); 6 June 1967, 1F [UCLA]. Maracay, San Jacinto, 9 Sept 1928, L.E.

Rozeboom, 1M (VZR 64); 450 m, 17 July 1969, J. Pulido and J. Valencia, 1 lpM (VZ 238-10), 1

pM (238-100), 2P, 4L (238), 1 IpM (VZ 239-12), 2 lpF (239-10,-15), 2 pM (239-101,-102), 1 pF

(239-100), 3 IP (239-11,-13,-16), 8M, 2F, 13P, 9L (239), 3 lpM (VZ 240-11,-12,-14), 2 IpF (240-

15,-16), 1 pM (240-101), 1 pF (240-100), 3 IP (240-10,-13,-17), 3M, 3F, 9P, 29L (240) [UCLA].

Maracay, Natl. Rt. 2 between Maracay and Choroni, 900 m, 26 July 1969, J. Pulido and J. Valen-

cia, 1 pF (VZ 269-100); 20 km N of Maracay on Natl. Rt. 2, 800 m, 6 Aug 1969, J. Valencia, 1

IpF (VZ 315-30) [UCLA]. Ocumare de la Costa, 10 m, 10 July 1969, J. Valencia and T. Zavor-

tink, 4L (VZ 132) [UCLA]. Ocumare de la Costa, MAC Cacao Dispersion Center, 100 m, 10 July

1969, J. Bergland, J. Pulido, J. Valencia and T. Zavortink, 3 lpM (VZ 139-15-17), 5 lpF (139-10-

14), 2 IP (139-18,-19), 1M, 4F, 4P, 10L (139); 12 July 1969, J. Pulido and J. Valencia, 1 lpM (VZ

177-30), 1 lpF (177-31), 1 lpM (VZ 179-21), 2 lp (179-20,-22), 2P, 4L (179), 2 IpM (VZ 180-20,

-23), 2 IpF (180-21,-24), 1 pF (180-100), 1 lp (180-22), 1L (180), 1 IpF (VZ 184-20); 5 Aug 1969,

J. Valencia, 1 pM (VZ 306-103), 1 pF (306-104), 3L (306), 3 pF (VZ 308-100-102), 1P, 6L (308)

[UCLA]. Ocumare de la Costa, Rio Cumboto, 60 m, 28 July 1969, J. Pulido and J. Valencia, 1L

(VZ 275), 1 pM (VZ 277-100) [UCLA]. Puerto Ocumare, near sea level, 10 July 1969, J. Berg-

land and T. Zavortink, 2 lpM (VZ 133-20,-21), 1L (133) [UCLA]. Rancho Grande, 14 km NW of

of Natl. Rt. 3, 500 m, 12 July 1969, J. Pulido and J. Valencia, 1 lpM (VZ 187-14), 1 IpF (187-13),

1L (187), 1 lpM (VZ 189-12), 1 lpF (189-11), 1 IP (189-10), 2P, 7L (189), 3 lpM (VZ 190-11,-13,

-14), 2 lpF (190-15,-16), 1 pM (190-100), 1 1P (190-10), 3 IpM (VZ 193-13,-15,-16), 5 IpF (193-

10,-11,-14,-17,-21), 1 pM (193-100), 13P, 16L (193) [UCLA]. Rancho Grande, 16 km NW of on

Natl. Rt. 3, 400 m, 5 Aug 1969, J. Valencia, 1 lpF (VZ 305-10), 4 pM (305-100-102,-104), 1 pF

(305-103), 1 lp (305-30), 2M, 1F, 2P, 22L (305) [UCLA]. Turiamo, Sept 1944, 1M (VZK 31)

[UCLA]. Carabobo: Borburata, 5 m, 24 July 1969, J. Pulido and J. Valencia, 4 lpM (VZ 264-10,

-12-14), 2 IpF (264-11,-15), 1 pF (264-100), 2M, 1F, 4P, 7L (264) [UCLA]. Guigue, 500 m, 24

July 1969, J. Pulido and J. Valencia, 1F, 1P (VZ 266), 1 pF (VZ 267-100) [UCLA]. Mariara, 420-

450 m, 19 July 1969, J. Pulido and J. Valencia, 3 lpM (VZ 246-10,-11,-17), 5 IpF (246-12-16), 2F,

2P, 2L (246), 1 IpF (VZ 247-10), 3L (247) [UCLA]. San Joaquin, 420 m, 19 July 1969, J. Pulido

and J. Valencia, 1 lpM (VZ 249-10), 2 lpF (249-11,-14), 1 pM (249-102), 1 pF (249-101), 5 IP

(249-12,-13,-15-17), 1P, 6L (249) [UCLA]. Monagas: Quiriquire, June 1935, 1Mgen [USNM].

NO DATA. 28 Dec 1933, J. Diaz, 1Mgen; 1L [USNM].

2. Culex (Anoedioporpa) canaanensis Lane & Whitman

Figs. 12,26

1943. Culex (Isostomyia) canaanensis Lane & Whitman 1943:398-400. TYPE: Holotype male

(2692), Sao Joao de Petropolis, Vale do Canaa, Espirito Santo, Brazil, Apr or July 1940,

L. Whitman [CPRR].

Culex (Anoedioporpa) canaanensis of Belkin (1968b:12); Belkin, Schick and Heinemann (1971:

27); Xavier (1973: 160); Knight and Stone (1977:195).

Culex (Isostomyia) canaanensis of Rozeboom and Komp (1948:400).

Culex (Aedinus) canaanensis of Stone, Knight and Starcke (1959:281); Stone (1967:218).

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 41

Culex (Tinolestes) canaanensis of Lane (1953:396-398); Duret and Damasceno (1955:394,395).

Culex canaanensis of Forattini, Rabello and Cotrim (1970:37).

FEMALE. Not seen; description based on Lane and Whitman (1943:398-400).

Similar to conservator, differing in the following. Head: Decumbent scales on ver-

tex predominantly whitish. Palpus about 2.0 length of clypeus. Thorax: Mesonotal

scales dark brown. Pleural integument blackish and greenish. Scales on upper mar-

gin of ppn brown.

MALE. Not seen. Similar to female in coloration. Palpus about 2.0 length of cly-

peus.

MALE GENITALIA (fig. 26). Not seen; description and figure based on Lane and

Whitman (1943:398-400). As figured; diagnostic characters as in key. Segment IX:

Tergite lobes small, moundlike, each with 6-10 short setae; interlobar space wide.

Sidepiece: Roughly triangular in outline; length about 2.0 greatest width; tergal sur-

face with longer bristles; sternal surface with shorter setae; tergal surface laterad of

subapical lobe with a densely setose area. Lobe: Proximal division with a distinct

stem, setose on under surface to basal 0.5 and bearing 2 rods with hooked tips, upper

rod inserted at apex, lower slightly basad. Distal division distinct, with 1 large striat-

ed submedian leaf and 1 smaller subapical leaf; apex with 4,5 short setae. Clasper:

About 0.55 length of sidepiece, smoothly curved; | submedian and 1 subapical seta

on inner surface. Lateral plate probably as in rest of Conservator Group. Proctiger:

Paraproct with 6,7 blunt apical teeth.

PUPA. Not seen; description based on Lane (1953:397). Cephalothorax: Integu-

ment lightly pigmented. All hairs short. Trumpet: Strongly sclerotized, index about

6.0. Abdomen: Integument lightly pigmented. Hair 1-II very small, weakly

branched, distinguishable at 100X; hair 5-III moderately long, extending to middle

of succeeding segment; 5-IV,V long, extending to apex of succeeding segment; 5-VI

barely extending to apex of succeeding segment; 9-VII,VIII with 3-5 short branches.

Paddle: Elongate, longer than wide, more than 2.0 of segment VIII.

FOURTH INSTAR LARVA. Not seen; description based on Lane and Whitman

(1943:400) and Lane (1953:397-398). Head: Width slightly greater than length.

Hair 4-C single; 5-C with 3 branches; 9-C with 4 branches. Antenna: Length about

0.5 of head, spiculose to hair 1-A; all hairs single except 1-A. Thorax: Hairs 1,2-P

single; 3-P double; 4-P double. Segment VIII: Comb scales in a patch of several

rows. Siphon: Long, index about 15.0. Integument strongly pigmented. Subven-

tral hairs (1,1a-S) apparently 4 pairs; subdorsal hairs (2a-S) 2 pairs. Pecten barely ex-

tending to proximal 0.2; individual tooth spinelike. Anal Segment: Integument

strongly pigmented. Hair 1-X small, weakly branched; 2,3-X long, single. Gills short;

ventral apparently shorter than dorsal.

SYSTEMATICS. Culex canaanensis is distinguished from other members of the

subgenus in the male genitalia by the combination of: (1) proximal division of sub-

apical lobe setose on under surface to basal 0.5, (2) tergal surface of sidepiece laterad

of subapical lobe with a densely setose area, and (3) distal division with 2 leaves mid-

way on stem. Although the immature stages have been described (Lane and Whitman

1943:398-400; Lane 1953:396-398), their structural details are poorly known. We

have not seen any material of canaanensis, but it appears to be closely related to con-

servator.

The holotype of canaanensis, whose location has been unknown in the past, is

now in the collection at the “‘Centro de Pesquisas Rene Rachou” [CPRR], Instituto

de Endemias Rurais, Fundacao Instituto Oswaldo Cruz, Belo Horizonte, Minas Ger-

ais, Brazil (Xavier 1973:160).

42 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

DISTRIBUTION (fig. 12). Culex canaanensis is known only from the type locali-

ty in Espiritu Santo, southeastern Brazil.

Material examined: none.

BRAZIL. Espirito Santo: Sao Joao de Petropolis, Vale do Canaa, Apr or July 1940, L. Whit-

man, holotype male (2692) [CPRR] ; Vale do Canaa, male paratype [FH, 3843]. Data from origi-

nal description and Forattini, Rabello and Cotrim (1970:37).

3. Culex (Anoedioporpa) damascenoi Duret

Figs. 12,26

1969. Culex (Anoedioporpa) damascenoi Duret 1969:143-145. TYPE: Holotype male, Rio

Preto, Municipio de Joao Goulard, Amazonas, Brazil, 18 July 1964, J.P. Duret [Duret] .

1971. Culex (Aedinus) menui Clastrier 1971:649-653. TYPE: Holotype male with pupal skin

({FGC] 3612-30), Maripasoula, Inini, French Guiana, 24 Aug 1968, J. Clastrier [MNHP].

NEW SYNONYMY.

Culex (Anoedioporpa) damascenoi of Stone (1970: 164); Belkin, Schick and Heinemann (1971:27);

Knight and Stone (1977:196); Heinemann and Belkin (1978b:437).

Culex (Aedinus) menui of Fauran and Pajot (1974: 102).

FEMALE. Unknown.

MALE. Not seen; description based on Duret (1969: 143-145). As described for

the subgenus with the following additional features. Head: Palpus short, approxi-

mately 3.0 length of clypeus. Decumbent scales on vertex narrow, grayish. Erect

scales forked, dark brown. Thorax: Mesonotal integument chestnut gray. Pleural

coloration mostly yellowish. Haltere with a pale stem; scales on knob dark. Abdo-

men: Tergites with dark scales; sternites creamy.

MALE GENITALIA (fig. 26). Not seen; figure and description based on Duret

(1969: 143-145). As figured; diagnostic characters asin key. Segment IX: Tergite

lobes indistinct, each with 4,5 fine setae; interlobar space wide. Sidepiece: Roughly

conical; length about 2.0 greatest width; basotergal and lateral surfaces with scat-

tered long bristles; sternal surface with shorter setae; tergal surface laterad of proxi-

mal division with a large densely setose area. Lobe: Proximal division with a moder-

ately long stem bearing 2 rods with hooked tip, upper rod inserted at apex, lower

slightly basad. Distal division with | large subbasal leaf on stem; apex with a long fil-

ament with recurved expanded tip on upper surface, 3 broad filaments with expand-

ed apexes on upper angle, between these a shorter lanceolate seta. Clasper: About

0.65-0.70 length of sidepiece, slightly bent mesad at distal 0.4; distal 0.3 of outer sur-

face with many closely set spiculelike transverse ridges; 1 submedian and | subapical

seta on inner surface. Lateral Plate: Basal hook sclerotized, strongly curved, bearing

a few scattered denticles; apical process long, expanded distally; sternal spine short,

pointed. Proctiger: Paraproct with a row of 6,7 blunt apical teeth.

PUPA. Not seen; description based on that of menui by Clastrier (1971:65 1-653).

Abdomen: 1.7-1.8 mm. Trumpet: 0.35 mm; index about 6.5-7.0. Paddle: 0.4 mm.

Cephalothorax: Integument lightly pigmented, wing case slightly darker. Hairs 1,3-

C single; 2-C double; 4,5-C double; 8,9-C single; 10-C triple. Trumpet: Strongly pig-

mented and distinctly contrasting with cephalothorax. Abdomen: Integument mod-

erately pigmented, progressively lighter caudad. Hair 1-II] extremely small, branched;

4-VI short, double, cephalolaterad of 5-VI; hair 1-VI,VII single; 9-VIII with 5 branch-

es. Paddle: Lightly pigmented, longer than wide, about 2.0 of segment VIII. Male

genital lobe extending to 0.6 of paddle.

LARVA. Unknown.

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 43

SYSTEMATICS. Culex damascenoi is known only by the holotype male. In the

male genitalia it is distinguished from other members of the subgenus by the combi-

nation of: (1) tergal surface of sidepiece laterad of subapical lobe with a large dense-

ly setose area and (2) distal division of subapical lobe with only | enlarged leaf near

middle of stem. We have not seen the type of damascenoi, but from the description

and illustration it appears to be distinct, although closely related to conservator.

We are also synonymizing Culex menui, described by Clastrier (197 1:649-653),

with damascenoi. Although we have not seen the type material of menui, it is evi-

dent that this species from French Guiana is conspecific with Culex damascenoi

from Brazil, as the original description of menui obviously matches in all pertinent

details the diagnostic characters of damascenoi. Culex menui is described from the

holotype male with the associated pupal skin. The pupal features of damascenoi,

therefore, are based on the description of menui Clastrier 1971.

BIONOMICS. The immature stages of damascenoi are found in treeholes. Noth-

ing is known about the behavior of the females.

DISTRIBUTION (fig. 12). Culex damascenoi is presently known only from the

state of Amazonas in Brazil and the territory of Inini in French Guiana.

Material examined: none.

BRAZIL. Amazonas: Rio Preto, Municipio de Joao Goulard, male [Duret; holotype of dama-

SCEnOL| .

nce GUIANA. Inini: Maripasoula, track to Wacapou near airfield, 100 m, 24 Aug 1968,

J. Clastrier, male (FGC 3612-30) [MNHP, holotype of menui] ; Inini Experiment Station, adult

(Fauran and Pajot 1974:102).

4. Culex (Anoedioporpa) browni Komp

Pigs: 12,16, 17

1936. Culex (Isostomyia) browni Komp 1936:326-328. *TYPE: Holotype male with genitalia,

Gatun, Canal Zone, Panama, 29 Nov 1933, C.G. Brown [USNM].

Culex (Anoedioporpa) browni of Belkin (1968b:12); Knight and Stone (1977:195); Heinemann

and Belkin (1978a:183; 1978c:523).

Culex (Anoedioporpa) sp undetermined of Heinemann and Belkin (1979:107).

Culex browni of Galindo, Carpenter and Trapido (1955:159).

Culex (Isostomyia) browni of Lane (1939:73); Rozeboom and Komp (1948:403); Galindo, Car-

penter and Trapido (1951:102,104,105,108,110,111,126); Stone and Knight (1957:58).

Culex (Aedinus) browni of Stone, Knight and Starcke (1959:281); Belkin, Schick and Heinemann

(1965:55-56).

Culex (Tinolestes) browni of Lane (1953:395-396); Duret and Damasceno (1955:395,400,404);

Galindo and Blanton (1955:70).

FEMALE. Wing: 2.9 mm. Proboscis: 2.1 mm. Forefemur: 1.75-1.80 mm. Abdo-

men: about 2.2 mm. Essentially similar to conservator, differing in the following.

Head: Decumbent scales on vertex predominantly white, a few median ones brown-

ish. Erect scales on occiput brown. Palpus about 0.15 length of proboscis. Thorax:

Scales on mesonotum predominantly brown, except for light scales on anterior prom-

ontory and along anterior part of lateral prescutal area. Pleural integument yellow-

ish to faintly green. Scales on upper margin of ppn white; antealar area above para-

tergite with white scales; lower mep with a bristle.

MALE. Wing: 2.9 mm. Proboscis: 2.25 mm. Forefemur: 1.8 mm. Similar to fe-

male in general coloration. Palpus about 0.15 length of proboscis,

MALE GENITALIA (fig. 16). As figured; diagnostic characters as in key. Seg-

ment IX: Tergite lobes small, moundlike, each with 7,8 short, fine subbasal setae;

44 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

interlobar space wide. Sidepiece: Roughly conical; length about 2.0 of greatest

width; lateral surface with longer bristles and a few scales; sternal surface predomi-

nantly with shorter setae, long bristles restricted to apical half; tergal surface basad

of lobe with a densely setose, swelling boss; scattered short setae basad of boss.

Lobe: Proximal division with a moderately long stem; with enlarged apex bearing 2

rods with hooked tip, upper rod inserted at apex, lower slightly basad. Distal divi-

sion distinct, with a short lanceolate seta midway on stem; 4 subapical lanceolate se-

tae on lower surface; apex with a long filament with a recurved expanded tip on up-

per surface, a pointed leaf and a lanceolate filament on upper angle, between these 2

short foliaceous setae with pointed apexes. Clasper: About 0.65-0.70 length of side-

peice, strongly curved at middle; distal 0.4 of outer surface with many closely set

spiculelike transverse ridges; 2 subapical setae on inner surface. Lateral Plate: Basal

hook sclerotized, strongly curved, bearing a few scattered denticles; apical process

distinct, moderately long, expanded at tip; sternal spine short, pointing laterad.

Proctiger: Cercal sclerite broad basally, apex hooked and pointing basad; cercal setae

absent. Paraproct with a row of 6,7 blunt apical teeth.

PUPA (fig. 16). Abdomen: 2.0-2.2 mm. Trumpet: 0.38-0.43 mm; index about

5.5-6.5. Paddle: 0.55-0.60 mm. As figured; diagnostic characters asin key. Cephalo-

thorax: Integument lightly pigmented. Hairs 1,3-C single; 2-C double; 4-C double;

5-C usually triple; 6-C single; 7-C double; 8,9-C always single; 10-C at least 3-branched

(3-5). Trumpet: Very strongly pigmented and strongly contrasting with cephalo-

thorax. Abdomen: Integument lightly pigmented. Hair 1-II short, weakly branched,

readily distinguishable at 100X; hair 7-II usually single; 4-VI short, weakly branched,

cephalolaterad of 5-VI; hair 1-VI,VII single; 9-VII usually 5-branched (3-5); hair 9-

VIII usually 6-branched (4-7). Paddle: Lightly pigmented, longer than wide, about

2.0 of segment VIII. Male genital lobe extending to 0.50 and female to 0.25 of

paddle.

FOURTH INSTAR LARVA (fig. 17). Head: 0.90-0.95 mm. Siphon: 2.0-2.1 mm;

index 13.0-14.0. Anal Saddle: 0.4 mm. As figured; diagnostic characters as in key.

Head: Width subequal to length. Hair 4-C usually single (1,2); hair 5-C usually with

5 branches (4-6); hair 9-C usually with 6 branches (5-8); hair 14-C single; 15-C usual-

ly double (2,3), short, not extending to base of mental plate. Mental plate with a

strong median shouldered tooth and 8,9 distinct teeth on either side. Antenna:

Length about 0.5 of head, distinctly spiculose to hair 1-A; all hairs single except 1-A

(13,12-14). Thorax: Hair O-P short, weak, with dendritic branches; 1,2-P long, sin-

gle; 3-P moderately long, about 0.4 of 1-P, usually triple (2-4); hair 4-P usually dou-

ble (2,3); hairs 5,6-P long, single; 7-P usually double (1-3); hair 14-P small, single; 4-

M double or triple. Abdomen: Hair 3-I double or triple; 6-I with 4 branches (3,4);

hair 7-I single; 6-II long, longer than segment, usually triple (2-4); hair 6-III-V longer

than segment, double; 1-V long, single. Segment VIII: Comb scales in a patch of 3,4

irregular rows, about 39-50 in number; individual scale with fringed apex. Siphon:

Integument lightly pigmented, darker basally and with a narrow dark basal ring. Sub-

ventral hairs (1,1a-S) 4 pairs, proximal 2 pairs long and single, distal 2 small and

branched; subdorsal hairs (2a-S) 2 pairs, short, located in distal 0.2 of siphon. Pecten

extending barely to proximal 0.2; individual tooth long, pointed apically, with a sub-

basal denticle ventrally. Anal Segment: Integument moderately pigmented, slightly

imbricate. Hair 1-X short, weakly branched; 2,3-X long, single. Gills long; dorsal 2.0-

2.5 length of saddle; ventral shorter, subequal to length of saddle.

SYSTEMATICS. Culex browni can be distinguished from other members of the

Conservator Group by the combination of the following: in the male genitalia by

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 45

(1) densely setose area on tergal surface of sidepiece basad of subapical lobe in form

of a swelling boss and (2) a lanceolate seta midway on stem of distal division of sub-

apical lobe; in the pupa by (1) hair 1-VI,VII single and (2) hair 5-III single or double;

in the larva by (1) hair 1-V long, single and (2) gills long, dorsal pair about 2.0-2.5

length of saddle.

The record of browni from Ecuador is somewhat doubtful, since it is based only

on 2 larvae collected in Napo Province (Amazon drainage). However, the larval fea-

tures agree very well with those of the type population from Panama.

BIONOMICS. The immature stages of browni have been collected in bamboo,

treeholes and occasionally artificial containers including bamboo pots. They are

found in bamboo traps near ground level and in the forest canopy (Galindo, Carpen-

ter and Trapido 1951:126-127). Females of browni are not known to bite man, and

males are only occasionally found in light trap collections (Galindo and Blanton

1935270),

DISTRIBUTION (fig. 12). Culex browni is known to occur in Panama (Canal

Zone, Darien), Colombia (Caribbean, Pacific and Orinoco drainages) and Ecuador

(Amazon drainage). It is the only species of Anoedioporpa known from the Pacific

side of the Andes in South America, being found as far south as the department of

Valle del Cauca in Colombia. It has been collected from near sea level to about 600

m above sea level.

Material examined: 234 specimens; 34 males, 24 females, 71 pupae, 105 larvae;

66 individual rearings (39 larval, 17 pupal, 10 incomplete).

COLOMBIA. Antioquia: Zaragoza, 22 km W, 18 Dec 1970, C.H. Porter, 1 pM (COP 193-10)

[UCLA]. Meta: Villavicencio, Bosque Ocoa, Apiary Rd., 23 Apr 1948, 1 lp (CV 571-2) [UCLA].

Valle del Cauca: Buenaventura, Rio Raposo, R.F. Virus Field Station, 30-40 m, 9 Dec 1964, V.H.

Lee, 1 IpF (COL 35-16), 8L (35); 13 Jan 1965, V.H. Lee, 1 lpM (COL 36-10), 1 IpF (36-11), 4 pM

(COL 37-12-15), 3 IpF (37-10,-18,-19), 1 pM (37-100), 2 IP (37-11,-17), 4L (37), 3 lpM (COL 40-

20,-22,-23), 1 IpF (40-21), 1 IP (40-24); 27 Jan 1965, V.H. Lee, 1 pM (COL 45-100), 2 lpM (COL

48-10,-11); near mouth of Rio Raposo, less than 10 m, 23 Feb 1965, V.H. Lee, 1M, 1P (COL 56);

Rio Raposo, R. F. Virus Field Station, 30-40 m, 19 Jan 1966, V.H. Lee, 1 lpM (COL 146-22), 2

pM (146-100,-102), 2 pF (146-101,-103), 2 pM (COL 147-103,-104), 3 pF (147-100-102), 2 IpM

(COL 148-10,-14), 4 lpF (148-11,-12,-15,-16), 1L (148); 23 Mar 1966, V.H. Lee, 1 IP (COL 149-

12), 15L (149), 3 IpM (COL 150-13-15), 1 IpF (150-11), 1 Ip (150-16), 2 pM (150-101,-102), 2 IP

(150-10,-12), 1M (150-100), 17L (150), 1 IpF (COL 151-12), 1 pF (151-100) [UCLA].

ECUADOR. Napo: Cuyabeno, about 50 km W of, Tarapoa, 300 m, 26 May 1974, D.J. Pletsch,

2L (ECU 212) [UCLA].

PANAMA. Canal Zone: Gatun, 29 Nov 1933, C.G. Brown, M [USNM, holotype]. Darien:

Morti, Morti Hydro, 80 m, 7 Dec 1966, O.G.W. Berlin and R. Hinds, 1 pF (PA 989-100) [UCLA].

Rio Tacarcuna valley, 600 m, 14 June 1963, A. Quinonez, 6 lpM (PA 387-103,-107,-108,-111,-114,

-116), 6 IpF (387-106,-109,-110,-112,-113,-115), 3 lp (387-101,-102,-104), 7L (387); 24 June 1963,

A. Quinonez, 2 p (PA 420-103,-105) [UCLA]. Province not specified: Cerro Sete, 2 Apr 1946, P.

Galindo, 1M, 1F [USNM]. Santa Clara, La Venta, 9 Apr 1943, 1F [USNM].

5. Culex (Anoedioporpa) bamborum Rozeboom & Komp

Figs, 12,168.19

1948. Culex (Isostomyia) bamborum Rozeboom & Komp 1948:399-400. *TYPE: Holotype

male with associated pupal and larval skins and genitalia ({CV] 404), Acacias, Meta, Co-

lombia, 3 June 1947, L.E. Rozeboom [USNM].

Culex (Anoedioporpa) bamborum of Belkin (1968b:12); Knight and Stone (1977:195).

Culex (Aedinus) bamborum of Stone, Knight and Starcke (1959:281); Belkin, Schick and Heine-

mann (1965:10).

46 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Culex (Isostomyia) bamborum of Stone and Knight (1957:58).

Culex (Tinolestes) bamborum of Lane (1953:399-400); Rozeboom and Komp (1955:395,400,407).

FEMALE. Wing: 2.4 mm. Proboscis: 1.7 mm. Forefemur: 1.5 mm. Abdomen:

about 1.6 mm. Essentially similar to conservator, differing in the following. Head:

Decumbent scales on vertex white. Erect scales on occiput light brown. Palpus

about 0.13-0.14 length of proboscis. Thorax: Scales on mesonotum predominantly

brown, except for light ones on anterior promontory and along anterior part of later-

al prescutal area. Pleural integument smoky bluish gray. Scales on ppn white; ante-

alar area above paratergite with white scales; lower mep with a single bristle.

MALE. Wing:2.4 mm. Proboscis: 1.8 mm. Forefemur: 1.5 mm. Similar to fe-

male in general coloration. Palpus about 0.14 length of proboscis.

MALE GENITALIA (fig. 18). As figured; diagnostic characters asin key. Seg-

ment IX: Tergite lobes small, moundlike, each bearing 6-8 fine subbasal setae; inter-

lobar space wide. Sidepiece: Roughly conical; length about 2.0-2.3 of greatest

width; lateral surface with long bristles; sternal surface with shorter setae. Lobe:

Distinct swollen boss proximad of subapical lobe, clothed with fine setae in promi-

nent tubercles, and with 2-4 specialized, enlarged, foliaceous setae near upper angle.

Proximal division with a moderately long stem bearing 2 rods with hooked tips, up-

per rod inserted at apex, lower slightly basad. Distal division with 1 submedian en-

larged leaf and 4 subapical broad filaments; apex with a long filament with a recurved

expanded tip on upper surface, a slender leaf on upper angle, between these a lanceo-

late seta and a pointed seta. Clasper: About 0.50-0.55 length of sidepiece, distal

half strongly curved mesad; distal 0.3 of outer surface with many closely set spicule-

like transverse ridges; 1 submedian and 1 subapical seta on inner surface. Lateral

Plate: Basal hook sclerotized, strongly curved, bearing a few scattered denticles; api-

cal process moderately long, expanded apically and truncate; sternal spine short,

pointed mesad. Proctiger: Cercal sclerite long, attenuated distally, digitiform, and

with a smaller subbasal pointed sternal process; usually one cercal seta. Paraproct

with a row of 9-12 blunt apical teeth and a few fine hairs.

PUPA (fig. 18). Abdomen: 2.2-2.5 mm. Trumpet: 0.45-0.50 mm; index about

8.0-9.0. Paddle: 0.5-0.6 mm. As figured; diagnostic characters asin key. Cephalo-

thorax: Integument moderately pigmented, wing case slightly darker. Hairs 1,3-C

single; 2-C double; 4,5-C double; 6-C single; 7-C double; 8,9-C single; 10-C usually 2-

4-branched. Trumpet: Very strongly pigmented and strongly contrasting with ceph-

alothorax. Abdomen: Integument moderately pigmented, progressively lighter cau-

dad. Hair 1-II extremely small, rarely visible at 100X; hair 7-I] usually single, rarely

double; 4-VI short, weakly branched, cephalolaterad of 5-VI; hair 1-VI,VII single; 9-

VII at least 4-branched (4-6); hair 9-VIII with 6 branches. Paddle: Lightly pigment-

ed, longer than wide, about 2.0 of sigment VIII. Male genital lobe extending to 0.4-

0.5 and female to 0.3 of paddle.

FOURTH INSTAR LARVA (fig. 19). Head: 0.8 mm. Siphon: 1.5-2.1 mm; index

25.0-27.0. Anal Saddle: 0.25 mm. As figured; diagnostic characters as in key.

Head: Width about 1.1 of length. Hair 4-C usually single; 5-C usually with 4 branch-

es (4,5); hair 9-C usually with 5 weak branches (5,6); hair 14-C single; 15-C single or

double, short, not extending to base of mental plate. Mental plate with a strong me-

dian shouldered tooth and 8,9 distinct teeth on either side. Antenna: Length about

0.5 of head, distinctly spiculose to hair 1-A; all hairs single except 1-A (15,12-18).

Thorax: Hair O-P short, weak, with dendritic branches; 1,2-P long, single; 3-P moder-

ately long, about 0.5 of 1-P, double; 4-P long, double; 5,6-P single; 7-P double; 14-P

small, single; 4-M double. Abdomen: Hair 3-I triple; 6-I usually with 3,4 branches;

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 47

7-I single; 6-II long, triple; 6-III-V moderately long, subequal to or slightly longer

than segment; 1-V long, single. Segment VIII: Comb scales in a patch of 3,4 irregu-

lar rows, about 64-69 in number; individual scale slightly spatulate, with fringed

apex. Siphon: Integument lightly pigmented, darker basally and with a narrow dark

basal ring. Subventral hairs (1,la-S) 4 pairs, distal ones small; subdorsal hairs (2a-S)

2 pairs, located in distal 0.2 of siphon. Pecten extending barely to proximal 0.16; in-

dividual tooth long, pointed apically, without lateral denticles. Anal Segment: Inte-

gument moderately to strongly pigmented, lightly imbricate. Hair 1-X short, weakly

branched; 2,3-X long, single. Gills variable, usually long; dorsal 1.5-2.0 length of sad-

dle; ventral subequal to length of saddle.

SYSTEMATICS. Culex bamborum can be distinguished from all other species of

Anoedioporpa in the male genitalia by the combination of: (1) setose area on side-

piece in form of a swollen boss proximad of subapical lobe, (2) 2-4 specialized, en-

larged, foliaceous setae on boss, and (3) enlarged leaf midway on stem of distal divi-

sion of lobe; in the pupa by (1) hair 1-I] extremely small, rarely visible at 100 X and

(2) moderately long trumpet with an index of about 8.0-9.0; and in the larva by ex-

tremely long siphon with index more than 25.0.

BIONOMICS. The immature stages of bamborum have been collected only in un-

cut bamboo internodes perforated by holes bored by insects. Little is known about

the behavior of females.

DISTRIBUTION (fig. 12). Culex bamborum is presently known only from the

department of Meta (Orinoco drainage) in Colombia at moderate elevations (about

400-500 m).

Material examined: 38 specimens; 9 males, 4 females, 7 pupae, 11 larvae; 6 indi-

vidual rearings (2 larval, 4 incomplete).

COLOMBIA. Meta: Acacias, 3 June 1947, L.E. Rozeboom, | lpM ({[CV] 404-1), 1 IpF ({CV]

404-7) [USNM, type series]. Villavicencio area, 1944, M. Bates, 2p, 1M, 1F (CV 65-1), 21(65-1A),

IM, 2F (65), 1 IM (CV 73-1) [UCLA]. Villavicencio, Bosque Ocoa, 22 July 1948, 1Mgen (CV

589-105), 3 lp (CV 590-1,-3,-4); 19 Aug 1948, M. Bates, 3L (CV 1008) [UCLA]. Villavicencio,

Bosque San Jose, 16 June 1944, M. Bates, 2M (CV 102) [UCLA]. Villavicencio, San Martin Rd.,

Chichimene, 11 Aug 1947, 1Mgen (CV 468-2), 1Mgen (CV 469-6) [UCLA].

6. Culex (Anoedioporpa) belemensis Duret & Damasceno

Figs, 12,26

1955. Culex (Tinolestes) belemensis Duret & Damasceno 1955:404-407. TYPE: Holotype male,

Belem, Para, Brazil, 19 Aug 1953, R.G. Damasceno and J.P. Duret [Duret] .

Culex (Anoedioporpa) belemensis of Belkin (1968b:12); Belkin, Schick and Heinemann (1971:27);

Xavier and Mattos (1975:245); Knight and Stone (1977:195).

Culex (Aedinus) belemensis of Stone, Knight and Starcke (1959:281); Fauran and Pajot (1974:

102).

Culex (Tinolestes) belemensis of Fauran (1961b:48).

FEMALE. Unknown, but presumably similar to male in general features.

MALE. Not seen; description based on Duret and Damasceno (1955:404-407).

Essentially similar to conservator, differing in the following. Head: Decumbent

scales on head grayish white. Erect scales on occiput brown. Palpus about 2.0 length

of clypeus. Thorax: Mesonotal scales brown. Pleural integument greenish yellow.

Scales on upper margin of ppn brown.

MALE GENITALIA (fig. 26). Not seen; description and figure based on Duret

and Damasceno (1955:406-407). Diagnostic characters as in key. Segment IX: Ter-

gite lobes distinct, moundlike, each with 8-12 submedian setae; interlobar space wide.

48 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Sidepiece: Roughly oval in outline; length about 1.5 of greatest width; tergal surface

of sidepiece caudad of lobe with scattered short setae. Lobe: Proximal division with

a moderately long stem bearing 2 rods with hooked tip, upper rod inserted at apex,

lower slightly basad. Distal division distinct, with 1 submedian and | subapical leaf;

apex with a long filament with recurved tip and a short seta on upper surface, 3

broad filaments on upper angle, between these a moderately long lanceolate seta.

Clasper: About 0.65 length of sidepiece, smoothly curved mesad; distal 0.3 of outer

surface with many closely set spiculelike transverse ridges; 2 subapical setae on inner

surface. Lateral Plate: Basal hook sclerotized, strongly curved; apical process mod-

erately long, expanded at tip; sternal spine short, slightly hooked. Proctiger: Cercal

setae indistinct. Paraproct with 8,9 blunt apical teeth.

PUPA, LARVA. Unknown.

SYSTEMATICS. Culex belemensis is known only by the holotype male. In the

male genitalia, belemensis is readily separated from all other members of the subge-

nus by the combination of: (1) absence of a densely setose area on the sidepiece

proximad of subapical lobe, (2) 2 distinct leaves on stem of distal divison of lobe,

and (3) moundlike IX tergite lobes. We have not seen the type, but from the original

description it appears to be distinct although closely related to conservator.

BIONOMICS. The immature stages of belemensis are unknown, but like other

members of the subgenus they should be found in treeholes and/or bamboo. The

holotype male was presumably taken in a sweeping collection from a tree trunk.

Nothing is known about the behavior of females.

DISTRIBUTION (fig. 12). Culex belemensis is known only from Belem, Para,

Brazil at low elevations and the interior of French Guiana.

Material examined: none.

BRAZIL. Para: Belem, 19 Aug 1953, R.G. Damasceno and J.P. Duret, 1M (Br.15,E.10) [Duret,

holotype].

FRENCH GUIANA. Inini: Experiment station (Fauran and Pajot 1974:102).

7. Culex (Anoedioporpa) chaguanco Casal, Garcia & Fernandez

Figs. 12,20,21

1968. Culex (Aedinus) chaguanco Casal, Garcia & Fernandez 1968:217-218. TYPE: Holotype

male (GA[=ARG]| 616-16) with associated larval and pupal skins and genitalia slide, 7 km

from Tablillas, nearest town Vespucio, Dep. San Martin, Salta, Argentina, 25 Feb 1967,

O.H. Casal and M. Garcia [INM].

Culex (Anoedioporpa) chaguanco of Stone (1970:164); Knight and Stone (1977:195).

FEMALE. Wing: 3.3 mm. Proboscis: 2.2 mm. Forefemur: 2.1mm. Abdomen:

about 2.2 mm. Similar to conservator, differing in the following. Head: Decumbent

scales on vertex mainly white, median ones brown. Erect scales on occiput dark

brown. Palpus about 0.16 length of proboscis. Thorax: Scales on mesonotum pre-

dominantly brown except for light scales on anterior promontory and anterior part

of lateral prescutal area. Pleural integument yellowish greenish. Scales on upper

margin of ppn brown. Antealar area above paratergite with white scales; lower mep

with a bristle.

MALE. Wing: 3.3 mm. Proboscis: 2.3 mm. Forefemur: 2.1 mm. Essentially sim-

ilar to female in coloration. Palpus about 0.16 length of proboscis.

MALE GENITALIA (fig. 20). As figured; diagnostic characters as in key. Seg-

ment IX: Tergite lobes distinct, moundlike, each with 4,5 subapical setae. Side-

piece: Roughly conical; length about 2.0 of greatest width; lateral surface with mod-

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 49

erately long bristles; sternal surface with shorter setae. Lobe: Proximal division with

long stem bearing 2 narrow rods with hooked tip, upper rod-inserted at apex, lower

slightly basad. Distal division distinct, with a large striated leaf midway on stem, a

smaller bent subapical leaf and 4 apical setae. Clasper: About 0.50 length of side-

piece, stout, slightly curved at middle and enlarged distally; distal 0.3 of outer sur-

face with closely set spiculelike transverse ridges; 2 subapical setae on inner surface.

Lateral Plate: Basal hook sclerotized, strongly curved; apical process long; sternal

spine short, slightly hooked. Proctiger: Apex of cercal sclerite digitiform; cercal se-

tae 1 or 2. Paraproct with a row of 6,7 blunt apical teeth.

PUPA (fig. 20). Abdomen: 2.3-2.4mm. Trumpet: 0.45 mm; index about 6.0.

Paddle: 0.55 mm, As figured; diagnostic characters as in key. Cephalothorax: Inte-

gument moderately pigmented, wing case slightly darker. Hairs 1-3-C usually single;

4-7-C double; 8,9-C usually single; 10-C at least 2-branched (2-4). Trumpet: Very

strongly pigmented and strongly contrasting with cephalothorax. Abdomen: Integu-

ment moderately pigmented, progressively lighter caudad. Hair 1-I] short, weakly

branched, readily distinguishable at 100X; hair 7-II single or double; 4-VI short,

weakly branched, cephalomesad of 5-VI; hair 1-VI usually double; 1-VII single; 9-VII

usually triple (2-4); hair 9-VIII with 5 branches (4-7). Paddle: Lightly pigmented,

longer than wide, about 2.0 of segment VIII. Male genital lobe extending to 0.5 and

female to 0.3 of paddle.

FOURTH INSTAR LARVA (fig. 21). Head: 0.90-0.95 mm. Siphon: 2.1-2.2 mm;

index 18.0-19.0. Anal Saddle: 0.35 mm. As figured; diagnostic characters as in key.

Head: Width about 1.1 of length. Hair 4-C usually single (1,2); hair 5-C usually with

4 branches (4,5); hair 9-C short, with 6,7 weak branches; 14-C single; 15-C usually

with 3 branches (2-4), short, not extending to base of mental plate. Mental plate

with a strong median shouldered tooth and 7 distinct teeth on either side. Antenna:

Length about 0.5 of head, distinctly spiculose to hair 1-A; all hairs single except 1-A

(18,16-21). Thorax: Hair 0-P short, weak, with dendritic branches; 1 ,2-P long, sin-

gle; 3-P long, about 0.5 of 1-P, usually single (1,2); hair 4-P double; 5,6-P single; 7-P

double; 14-P small, single; 4-M double. Abdomen: Hair 3-I usually double (2,3);

hair 6-I usually with 4 branches (3-5); hair 7-I single; 6-II moderately long, subequal

to segment, usually with 3 branches (2-4); hair 6-III-V moderately long, shorter than

segment; 1-V long, single. Segment VIII: Comb scales in a patch of 3,4 irregular

rows, 48-63 in number; individual scale short, with fringed apex. Siphon: Integu-

ment moderately pigmented, darker basally and with a narrow dark ring. Subventral

hairs (1,la-S) 5 pairs, progressively smaller distad; subdorsal hairs (2a-S) 2 pairs, lo-

cated in distal 0.25 of siphon. Pecten extending barely to proximal 0.25; individual

tooth long, pointed apically, without denticles. Anal Segment: Integument strongly

pigmented, slightly imbricate. Hair 1-X short, weakly branched; 2,3-X long, single.

Gills usually short; dorsal subequal to length of saddle; ventral about 0.6 of dorsal.

SYSTEMATICS. Culex chaguanco can be readily separated from other members

of the subgenus in the male genitalia by the combination of: (1) absence of setose

area on sidepiece proximad of subapical lobe, (2) 1 enlarged leaf midway on stem of

proximal division of lobe, and (3) short stout clasper; in the pupa by the combina-

tion of: (1) shorter trumpet with an index of less than 7.0, (2) hair 1-VII single, and

(3) hair 1-VI usually double; and in the larva by the combination of: (1) hair 1-V

long and single, (2) short gills, dorsal pair subequal to length of saddle, and (3) 5

pairs of subventral hairs (1,1a-S) on siphon.

Culex chaguanco was described from northern Argentina, and its distribution ap-

parently represents the southern limit of Anoedioporpa.

50 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

BIONOMICS. The immature stages of chaguanco have been collected in treeholes,

where they were associated with Aedes (Pro.) terrens (Walker 1856) and Aedes (Pro.)

casali Schick 1970. All the known adults were reared from immatures, and nothing

is known about the behavior of females.

DISTRIBUTION (fig. 12). Culex chaguanco is presently known from the prov-

inces of Salta and Misiones in northern Argentina.

Material examined: 42 specimens; 3 males, 8 females, 11 pupae, 20 larvae; 11 indi-

vidual rearings (3 larval, 8 pupal).

ARGENTINA. Misiones: San Pedro, Route 14, Km 327, 13 Feb 1966, O.H. Casal and M. Gar-

cia, 1 lpF (ARG[=GA] 143-11), 11 (143-10), 1L (143) [UCLA]. Salta: Dep. San Martin, nearest

town Vespucio, 7 km from Tablillas, 25 Feb 1967, O.H. Casal and M. Garcia, 1 lpF (ARG[=GA]

616-10, paratype), 1 lpM (616-11, paratype), 1 pF (ARG[=GA] 618-115), 1SL (618); 6 June 1969,

O.H. Casal and M. Garcia, 2 pM (ARG[=GA] 773-104,-112), 5 pF (773-100,-101,-107,-108,-111)

[UCLA]. |

8. Culex (Anoedioporpa) originator Gordon & Evans

Figs, 12.22.23

1922. Culex originator Gordon & Evans 1922:323-327. *TYPE: Lectotype male (13.2/463)

with genitalia on 2 slides, Macapa, Manaos (Manaus), Amazonas, Brazil, 21 Dec 1921

(emerged 1 Jan 1922), R.M. Gordon [BM; designation of Belkin 1968b:18].

1941. Culex (Melanoconion) surukumensis Anduze 1941a:833-834. TYPE: male, Rio Surukum,

Bolivar, Venezuela, Nov 1940, P.J. Anduze, [DPFA]. NEW SYNONYMY.

Culex (Anoedioporpa) originator of Bonne and Bonne-Wepster (1925:260,264); Belkin (1968b:12,

18); Belkin, Schick and Heinemann (1971:27); Xavier and Mattos (1975:246); Belkin and Hei-

nemann (1976:272); Knight and Stone (1977:196); Knight (1978:41); Heinemann and Belkin

(1978b:393 407,437; 1979:80,94).

Culex (Isostomyia) originator of Edwards (1932:218); Senevet and Abonnenc (1939:112-114,130,

133); Lane (1939:74); Floch and Abonnenc (1942:9; 1947:7,8); Lane and Whitman (1943:

397); Rozeboom and Komp (1948:403).

Culex (Aedinus) originator of Dyar (1923b:189); Stone, Knight and Starcke (1959:282); Stone

(1963:135); Forattini (1 965:27).

Culex (Melanoconion) originator of Dyar (1928:346-347); Lima (1930:255); Komp (1935:10).

Culex (Tinolestes) originator of Lane (1953:394); Duret and Damasceno (1955:394,395-397,407);

Fauran (1961b:43-44); Cerqueira (1961:131-132).

Culex originator of Kumm and Novis (1937:501,511); Fauran (1961a:12-13); Aitken, Spence et al.

(1969:210); Forattini, Rabello and Cotrim (1970:45; 1973:468).

Culex (Melanoconion) surukumensis of Anduze (1941b:16).

Culex (Aedinus) surukumensis of Belkin, Schick and Heinemann (1965:75).

Culex (Aedinus) conservator in part of Stone, Knight and Starcke (1959:281); Cova Garcia, Sutil

and Rausseo (1966b:342-343). _

Culex (Anoedioporpa) conservator in part of Knight and Stone (1977:196).

Culex (Isostomyia) conservator of Senevet and Quievreaux (1941:264).

FEMALE. Wing: 2.6 mm. Proboscis: 1.9mm. Forefemur: 1.6 mm. Abdomen:

about 2.0 mm. Essentially similar to conservator, differing in the following. Head:

Decumbent scales on vertex predominantly white, a few median ones brownish.

Erect scales on occiput brown. Palpus about 0.15 length of proboscis. Thorax:

Scales on mesonotum predominantly dark brown, except for light scales on anterior

promontory and along anterior part of lateral prescutal area. Scales on upper margin

of ppn brownish. Lower mep area with a single bristle. Antealar area above parater-

gite with scattered white scales.

MALE. Wing: 2.5 mm. Proboscis: 2.1mm. Forefemur: 1.8 mm. Similar to fe-

male in general coloration. Palpus about 0.15 length of proboscis.

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 51

MALE GENITALIA (fig. 22). As figured; diagnostic characters as in key. Seg-

ment IX: Tergite lobes small, moundlike, each with 5,6 short subapical setae; inter-

lobar space wide. Sidepiece: Roughly oval in outline; length about 2.0 of median

width; tergal surface laterad of distal division of lobe with 2 patches of bristles: an

inner patch of 10-12 moderately long setae and an outer lateral group of 9-12 long

bristles; sternal surface with short setae. Lobe: Proximal division with a long stem,

setose in basal 0.5, bearing 2 distinctly separated rods with hooked tip, upper rod in-

serted at apex, lower basad. Distal division distinct, with a moderately long subbasal

seta on stem; apex bilobed, proximal lobe with a long filament with an expanded re-

curved tip, a foliform seta with expanded tip and a short lanceolate seta, and distal

lobe with 1 longer and 1 shorter foliform seta with expanded apex. Clasper: About

0.70 length of sidepiece, sharply bent mesad at middle; distal 0.3 of outer surface

with spiculelike transverse ridges; 2 submedian setae on inner surface. Lateral Plate:

Basal hook sclerotized, strongly curved, bearing distinct denticles; apical process

moderately long, digitiform, pointing caudolaterad; sternal spine short, pointed.

Proctiger: Cercal sclerite short, broad, apex sharply pointed. Paraproct with 11-13

blunt apical teeth; subapical area densely setose.

PUPA (fig. 22). Abdomen: 2.2-2.4 mm. Trumpet: 0,35-0.40 mm; index about

5.5-6.0. Paddle: 0.55-0.60 mm. As figured; diagnostic characters as in key. Ceph-

alothorax: Integument lightly pigmented. Hairs 1,3-C single; 2,4,5,7-C double; 6-C

single; 8,9-C usually single; 10-C at least double (2-9). Trumpet: Strongly pigment-

ed and strongly contrasting with cephalothorax. Abdomen: Integument lightly pig-

mented. Hair 1-II short, weakly branched, readily distinguishable at 100X; hair 7-I]

usually single; 5-III single or double; 4-VI shorter, weakly branched, cephalolaterad

of 5-VI; hair 1-VI,VII always single; 9-VII with 3,4 branches; 9-VIII usually 5-

branched (4-6). Paddle: Lightly pigmented, longer than wide, about 2.0 of segment

VIII. Male genital lobe extending to 0.50 and female to 0.25 of paddle.

FOURTH INSTAR LARVA (fig. 23). Head: 0.85-0.90 mm. Siphon: 1.3-2.0 mm;

index 15.0-20.0. Anal Saddle: 0.35 mm. As figured; diagnostic characters as in key.

Head: Width about 1.1 of length. Hair 4-C usually double (1,2); hair 5-C usually

with 5 branches (4-7); hair 9-C usually with 7 branches (5-9); hair 14C single; 15-C

usually with 3 branches (2-4). Mental plate with a strong median shouldered tooth

and 7,8 distinct teeth on either side. Antenna: Length about 0.5 of head, distinctly

spiculose to hair 1-A; all hairs single except 1-A (14,10-16). Thorax: Hair O-P short,

weak, with dendritic branches; 1,2-P long, single; 3-P moderately long, about 0.5 of

1-P, usually with 4 branches (3-5); hair 4-P usually triple (2,3); hairs 5-7-P usually

single; 14-P small, single; 4-M usually with 3 branches (3,4). Abdomen: Hair 3-I usu-

ally single (1-3); hair 6-I usually with 4 branches (3,4); hair 7-I single; 6-I] moderately

long, subequal to segment, usually triple (2-4); hair 6-III-V longer than segment; 1-V

long, single. Segment VIII: Comb scales in a patch of 3,4 irregular rows, about 32-

44 in number; individual scale with spatulate fringed apex. Siphon: Integument

moderately pigmented, darker basally and with a narrow dark basal ring. Subventral

hairs (1,la-S) 4 pairs, progressively smaller distad; subdorsal hairs (2a-S) 2 pairs,

short, located in distal third of siphon. Pecten extending barely to proximal 0.2; in-

dividual tooth long, pointed apically, with denticles up to basal 0.5. Anal Segment:

Integument moderately pigmented, slightly imbricate. Hair 1-X short, weakly

branched; 2,3-X long, single. Gills short; dorsal subequal in length to saddle; ventral

about 0.4 of dorsal.

SYSTEMATICS. Culex originator is a distinct species, and is readily recognized

from other members of the subgenus in the male genitalia by the combination of:

52 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

(1) 2 distinct patches of bristles on tergal surface of sidepiece laterad of subapical

lobe, (2) a distinct, moderately long seta midway on stem of distal division of subapi-

cal lobe, (3) details of setae on apex of distal division, (4) sharply bent clasper, and

(5) densely setose area on subapical part of paraproct; in the pupa by: (1) hair 1-VI,

VII single and (2) hair 5-III single or double; and in the larva by the combination of:

(1) hair 1-V long and single, (2) siphon with 4 pairs of subventral hairs (1,la-S), (3)

hair 5-C at least 4branched (4-7), and (4) hair 7-P usually single, rarely double.

We have examined material of originator from Brazil, Guyana, French Guiana and

Trinidad, and there appears to be no significant variation in the structural details of

male genitalia or in the chaetotaxy of immature stages. We have also examined |

male genitalia from Martinique and | larva from Grenada, and they also agree with

the description of those of the type. Apparently this is the only species of Anoedio-

porpa found in the Lesser Antilles. The northern limit appears to be Martinique, but

more extensive collections should be made to ascertain its presence or absence in the

other islands of this chain.

On the island of Trinidad, originator is sympatric with conservator, but is not nor-

mally found in the same treehole or bamboo, thereby apparently exhibiting ecologi-

cal isolation; out of a total of 52 lots, the 2 species have been collected only once

from the same container (TR 100, bamboo pot). It appears to be the dominant spe-

cies on this island in relation to conservator, having been recorded from 42 lots, as

opposed to 11 lots for conservator. Culex originator has not been recorded to date

from the island of Tobago, although conservator has.

We are synonymizing surukumensis Anduze 1941 (a former synonym of conserva-

tor) with originator. Although we have not seen the type of surukumensis, this nom-

inal species appears to be conspecific with originator on the basis of male genitalic

characters.

BIONOMICS. The immature stages of originator have been collected primarily in

treeholes and bamboo. Mosquito studies on the island of Marajo in Brazil show that

originator is found also in clay pots, fallen leaves on the ground and in fruit rinds

(Kumm and Novis 1938:503,511). Apparently the females at least occasionally bite

man, as | female has been taken in a biting collection in Trinidad (TR 644) at 1000-

1500 hours.

DISTRIBUTION (fig. 12). Culex originator is presently known from Venezuela

(lower Orinoco drainage), Guyana, French Guiana and Brazil (Amazon drainage),

and from the islands of Martinique and Grenada (Lesser Antilles) and Trinidad.

Material examined: 488 specimens; 64 males, 65 females, 124 pupae, 235 larvae;

115 individual rearings (83 larval, 15 pupal, 17 incomplete).

BRAZIL. Amazonas: Manaus, Macapa, 21 Dec 1921, R.M. Gordon, 1M (13.2/463) [BM, lecto-

type]. Para: Belem, IPEAN, Reserva de Aura, 1-10 m, 8 Aug 1969, B.T. and B.G. Aitken, 2 lpM

(BRA 26-11,-14), 1 pF (26-10), 1 IP (26-12), 2L (26); 1-30 m, 29 July 1970, T.H.G. Aitken, 1 lpF

(BRA 54-22), 1 IF (54-23), 2L (54) [UCLA]. Belem, Utinga Forest, 35 m, 5 Aug 1969, B.T. and

B.G. Aitken, 11 (BRA 16-11), 1L (16) [UCLA] ; det. A. Toda [FH, 15855-91,E499-507]. Curra-

linho, Recreio do Piria, Rio Aracairou, 1F (K427) [USNM]. Curralinho, Rio Canaticu, 3M, 1F (K

285) [USNM]. Curralinho, Rio Itaucu, 1935, 1M (61) [USNM]. Curralinho, Rio Aramakari, Rio

Muruaca, Rio Pariaca [INER; Xavier and Mattos 1975:246]. Curralinho, Ilha de Marajo (Kumm

and Novis 1938:503,511). :

FRENCH GUIANA. Guyane: Approuague, det. Floch [FH, 6331]. Cayenne, foret de Cabas-

sou, 5-100 m, 22 May 1967, J. Clastrier, 1 IP (FGC 3146-14), 9L (3146); 9 June 1968, J. Clastrier,

1 pM (FGC 3470-16), 1 pF (3470-14), 2M (3470-20,-22), 4F (3470-13,-15,-23,-24); 14 July 1968,

J. Clastrier, 1 pF (FGC 3519-30); 8 Sept 1968, J. Clastrier, 1 lpM (FGC 3642-33), 1 lIpF (3642-32),

2 pF (3642-31,-34), 1 IpF (FGC 3643-31), 1 lpM (FGC 3644-37), 3 IpF (3644-35,-36,-38), 2 pM

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 53

(3644-32,-33), 1 pF (3644-30); 1 Dec 1968, J. Clastrier, 3 lpM (FGC 3903-46,-51,-52), 4 lpF

(3903-42,-44,-47,-49), 2 pM (3903-46,-50), 1 pF (3903-43), 2F (3903-41,-48), 1M, 4L (3903); 9

June 1968, J. Clastrier, 1 lpM (FGC 3470-30) [UCLA]. Cayenne, foret de la Chaumiere, 5-150 m,

1 Apr 1968, J. Clastrier, 1M (FGC 3335-11), 1F (3335-10), 1 lpM (FGC 3338-14), 2M (3338-10,

-13), 1F (3338-11), 4L (3338), 1 p (FGC 3340-13) [UCLA]. Cayenne, Institut Pasteur, 5 m, 1

May 1967, J. Clastrier, 1Mgen (FGC 3116) [UCLA]. Cayenne, Raban, 5 m, 2-3 Feb 1965, T.H.G.

Aitken, A. Guerra and R. Martinez, 1F (FG 43) [UCLA]. Remire, between Lac de Remire and

Lac Lalouette, 150 m, 18 Mar 1967, R.X. Schick, 2 lpM (FG 162-10,-11) [UCLA]. Various locali-

ties (Floch and Abonnenc 1942:9; 1947:7; Fauran 1961b:43-44). Inini: Various localities, inclu-

ding Saut Tigre (Fauran 1961b:43-44).

GRENADA. St. Andrew: Grenville, Balthasar Estate, 150 m, 15 Oct 1963, R. Martinez, 1L

(GR 43) [UCLA].

GUYANA. Mazaruni River, 29 June 1936, 1M (KO 1-9) [UCLA].

MARTINIQUE. Fort-de-France, 20 July 1905, A. Busck (Howard, Dyar and Knab 1915:310).

Locality unspecified but presumably the basis for the previous record, July 1905, A. Busck, 1M

with genitalia slide (690723-3) [USNM].

TRINIDAD. Nariva: Archers Estate, 50m,5 Nov 1964, A. Guerra, 2 lp (TR 818-124,-125), 3L

(818) [UCLA]. Biche, Caratal Rd., 10 m, 3 Dec 1964, F. Powdhar, 7L (TR 864) [UCLA]. Char-

uma Forest, 50-150 m, 27 Aug 1964, A. Guerra, 1 lpM (TR 634-106), 2L (634), 1F (TR 644); 8

Oct 1964, A. Guerra, SL (TR 753), SL (TR 754) [UCLA]. Nariva Swamp, Bush Bush Forest, near

sea level, 17 Feb 1964, TRVL, 2 IpM (TR 73-136,-139), 3 IpF (73-137,-138,-140), 2 pF (73-131,

-132); 26 Feb 1964, TRVL, 1 IpF (TR 98-142), 3 IpF (TR 99-148-150), 1 lpM (TR 100-103), 1L

(TR 102); 4 Mar 1964, TRVL, 6 IpM (TR 155-118,-120,-185,-186,-190,-199), 4 IpF (155-101,-102,

-119,-200), 1 IF (155-184), 4L (155); 13 May 1964, TRVL, 3 lpM (TR 386-115,-116,-197), 4 lpF

(386-114,-117,-181,-198), 1 pF (386-110), 1 lpF (TR 387-109) [UCLA]. St. Andrew: Arima, det.

J. Lane [FH, 10262-3,E-1287-97] ; det. Heredia [FH, 10930-43]. Coryal, 50 m, 18 June 1964, A.

Guerra, 2 IpM (TR 491-128,-129), 3 IpF (491-121,-122,-154), 9L (491) [UCLA]. Cumaca, 150-

200 m, 22 Oct 1964, TRVL, 3 lpM (TR 785-102,-104,-106), 2 IpF (785-101,-105), 1M, 1F, 2P, 2L

(785); 14 Jan 1965, A. Guerra, 2M, 1F, 2P, 3L (TR 942); 18 Feb 1965, TRVL, 4 lpM (TR 1012-

12-15), 1 IpF (1012-11), 12L (1012) [UCLA]. Mount Harris, 100 m, 16 July 1964, F. Powdhar,

SL (TR 566) [UCLA]. Platanal Rd. near Oropuche River, 100 m, 20 May 1965, A. Guerra, 1 lpM

(TR 1179-12), 4 IpF (1179-13-16), 1 pF (1179-11), 2F, 2P, 13L (1179) [UCLA]. Turure Forest,

30 m, 7 May 1966, A. Guerra, 1 lpM (TR 1511-11), 2 IpF (1511-10,-12) [UCLA]. St. David:

Grande Riviere, 50 m, 13 Mar 1964, A. Guerra, 3 pM (TR 199-123,-125,-128) [UCLA]. St.

George: Arena Forest Reserve, 50 m, 3 Sept 1965, F. Powdhar, 1 pM (TR 1386-100) [UCLA].

Aripo Valley, 150-250 m, 16 Apr 1964, A. Guerra, 3 IpF (TR 317-125,-127,-157); 17 Sept 1964,

A. Guerra, 10L (TR 704), 2L (TR 706), 8L (TR 707), 1L (TR 708), 4L (TR 709), 10L (TR 710);

25 Feb 1965, F. Powdhar, 1 pM (TR 1022-10), 1 pF (1022-11), 1L (1022); 15 Apr 1965, A. Guer-

ra, 1 IpF (TR 1107-10) [UCLA]. Blanchisseuse, 10 m, 10 Apr 1964, A. Guerra, 4 IpM (TR 305-

136,-144,-147,-149), 4 IpF (305-143,-145,-146,-148) [UCLA]. St. Joseph, 50 m, 9 Sept 1965, A.

Guerra, 1 IpM (TR 1391-20) [UCLA]. St. Patrick Estate, 200 m, 5 Feb 1966, A. Guerra, 2L (TR

1457), 1L (TR 1458) [UCLA]. Verdant Vale, 200-300 m, 10 Sept 1964, A. Guerra, 1L (TR 675);

12 Nov 1964, A. Guerra, SL (TR 825); 11 Mar 1965, A. Guerra, 1 IpF (TR 1028-19), 2 lpM (TR

1038-11,-13), 3 IpF (1038-12,-14,-15), 9L (1038); 25 Mar 1965, M. Moody, 1 lpM (TRM 13-51),

3L (13), 1 lpF (TRM 14-50) [UCLA].

VENEZUELA. Bolivar: Headwaters of Rio Surukum, Nov 1940 (Anduze 1941a:833-834).

9. Culex (Anoedioporpa) quasioriginator Duret

Fig. 12

1972. Culex (Anoedioporpa) quasioriginator Duret 1972:3-5. TYPE: Holotype male (1334),

Pirelli, near Belem, Para, Brazil, 16 Jan 1963, J. Duret [Duret].

Culex (Anoedioporpa) quasioriginator of Xavier and Mattos (1975:246); Knight and Stone (1977:

196).

54 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

FEMALE. Unknown.

MALE. Not seen; based on description of Duret (1972:3-5). Very similar in de-

tails to originator.

MALE GENITALIA. Not seen; description based on Duret (1972:3-5). Diagnos-

tic characters as in key. Segment IX: Tergite lobes small, moundlike, each with 5

subapical setae; interlobar space wide. Sidepiece: Distal part with long bristles on

tergal surface; sternal surface with shorter setae. Lobe: Proximal division with a

long stem, bearing 2 approximate rods with hooked tip, upper rod inserted at apex,

lower slightly basad. Distal division distinct, with a moderately long subbasal seta on

stem; apex bilobed, proximal lobe with a long filament with expanded tip, a longer

lanceolate seta and a shorter pointed seta, and distal lobe with 1 longer foliform seta

with expanded apex and a lanceolate seta. Clasper: About 0.70 length of sidepiece,

sharply bent mesad at middle; distal 0.3 of outer surface with spiculelike transverse

ridges: 2 submedian setae on inner surface. Lateral Plate: Basal hook sclerotized,

strongly curved, bearing distinct denticles; apical process digitiform; sternal spine

short, pointed. Proctiger: Cercal sclerite short, broad, apex pointed. Paraproct with

9 blunt apical teeth; subapical area densely setose.

PUPA, LARVA. Unknown.

SYSTEMATICS. Culex quasioriginator, described from the state of Para in Brazil,

is known only by the males. In the male genitalia, it is distinguished from all other

members of the subgenus except originator by the presence of a densely setose area

on the paraproct, and from originator by the absence of a setose area on the stem of

proximal division of subapical lobe. We have not seen the types of quasioriginator,

but from the description this species appears to be distinct, although closely related

to originator with which it is sympatric in Para.

BIONOMICS. The immature stages of quasioriginator are unknown, but they

probably occur in treeholes or bamboo. Nothing is known about the behavior of the

adult females. Apparently all the males have been caught in sweeping collections.

DISTRIBUTION (fig. 12). Culex quasioriginator is presently known iy from

the state of Para in Brazil.

Material examined: none.

BRAZIL. Para: Belem, 18 Aug 1953, J. Duret, 2M (4702,4712); Pirelli, 14 Jan 1963, J. Duret,

2M (555,581); 16 Jan 1963, J. Duret, 10M (557,558,579,580,1327,1334,1335,3783,3786,3788)

[type series] ; Rio Guajaru (Oriboca), 23 Aug 1963, J. Duret, 2M (4710,4715); Utinga, 26 Oct

1962, J. Duret, 1M (1336). Sao Domingos do Capim, Paragominas 24 June 1964, J. Duret, 1M

(1336) (Duret 1972:4-5).

10. Culex (Anoedioporpa) luteopleurus (Theobald)

Figs. 12,26

1903. Melanoconion luteopleurus Theobald 1903:239-240. *TYPE: Holotype female, Para,

Brazil, date not specified, H.E. Durham [BM].

Culex (Anoedioporpa) luteopleurus of Belkin, Schick and Heinemann (1971:27); Knight and Stone

(1977:196).

Culex (Micraedes) luteopleurus of Belkin (1968b:11,17); Xavier and Mattos (1975:249).

Culex (Aedinus) luteopleurus of Stone, Knight and Starcke (1959:282).

Culex (Melanoconion) luteopleurus of Dyar (1928:348-349); Edwards (1932:216); Komp (1935:

10); Lane (1939:67; 1953:495-496); Rozeboom and Komp (1950:98).

Culex (Tinolestes) luteopleurus of Duret and Damasceno (1955:395,397-401 ,404,407,408); Fauran

(1961b:48); Cerqueira (1961:133).

Culex luteopleurus of Forattini, Rabello and Cotrim (1970:43; 1973:468).

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 55

Melanoconion luteopleurus of Lutz (1904a:5); Peryassu (1908:240-241); Theobald (1910:455,

456); Surcouf and Gonzalez-Rincones (1911:208).

FEMALE. Specimen greasy and therefore characters based on original description

(Theobald 1903:239-240). Similar to conservator, differing in the following. Head:

Decumbent scales on vertex narrow, dull gray. Erect scales on occiput bright yellow.

Palpus about 0.15 length of proboscis. Thorax: Scales on mesonotum black, linear.

Antealar area above paratergite with black scales. Pleural integument bright yellow.

Scales on upper margin of ppn broad, brown. Lower mep with a strong bristle.

MALE. The association of male Juteopleurus with the holotype female by Duret

and Damasceno (1955:398) is questionable, because no individual rearings were

made, without which no association can be ascertained.

MALE GENITALIA (fig. 26). Not seen; figure and description based on Duret

and Damasceno (1955:399-400). Diagnostic characters asin key. Segment IX: Ter-

gite lobes indistinct, each with 8-10 weak setae; interlobar space wide. Sidepiece:

Roughly conical; length about 2.0 of greatest width; lateral surface with longer bris-

tles; sternal surface with shorter setae. Lobe: Proximal division long, with a row of

4 setae in basal 0.5 of stem and bearing 2 distinctly separated rods with hooked tip,

upper rod inserted at apex, lower slightly basad. Distal division distinct, with 4 en-

larged striated leaves midway on stem and 1 leaf subapically; apex with a strong, sin-

uous filament with an expanded recurved tip and 1 smaller lanceolate seta. Clasper:

About 0.50 length of sidepiece, sharply curved at middle; distal 0.5 expanded, with

a row of spiculelike transverse ridges on outer surface; 2 subapical setae on inner sur-

face. Lateral Plate: Basal hook sclerotized, strongly curved; apical process long; ster-

nal spine short, slightly hooked. Proctiger: Cercal setae 3. Paraproct with 9 blunt

apical teeth.

PUPA, LARVA. Unknown.

SYSTEMATICS. The original description of luteopleurus is based on a female col-

lected in an unspecified locality (possibly in or near Belem) in the state of Para, Bra-

zil. No other specimens were reported until 1954, when Duret and Damasceno col-

lected some males in Yaguarari and Oriboca in the state of Para, which they provis-

ionally identified as luteopleurus. Therefore, the association of sexes is only pre-

sumptive.

Culex luteopleurus can be distinguished from all other members of the subgenus

in the adults by: (1) bright yellow erect scales on vertex and (2) antealar area above

paratergite with black scales; and in the male genitalia by the combination of: (1)

IX tergite lobes indistinct, (2) stem of proximal division of subapical lobe with 4 se-

tae in basal 0.5, and (3) stem of distal division of subapical lobe with 5 submedian

leaves. On the basis of male genitalic characters, Juteopleurus appears to be a very

distinct member of the subgenus Anoedioporpa.

BIONOMICS. The immature stages of Juteopleurus are unknown, but like other

members of the subgenus will probably be found in treeholes and/or bamboo. The

holotype female and the males were presumably taken in sweeping collections.

DISTRIBUTION (fig. 12). Culex luteopleurus is at present known only from the

state of Para, Brazil at low elevations.

Material examined: holotype female examined by J.N. Belkin [BM].

BRAZIL. Para: Date and exact locality unspecified, H.E. Durham [BM, holotype]. Belem,

Oriboca, Rio Guajaru (Br59,E.16); Rio Moju, Yaguarari (Br41,E.1) [Duret] ; Rio Moju, Yaguarari

(Br41,E.2) [FH, 10,551]; Utinga [FH, E485-498,E820,E821].

56 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

11. Culex (Anoedioporpa) corrigani Dyar & Knab

Figs. 12,24,25

1907. Culex corrigani Dyar & Knab 1907:203. *TYPE: Holotype female, Tabernilla, Canal

Zone, Panama, date not specified, A. Busck [USNM, 10870] .

1914. Culex chalcocorystes Martini 1914:70-74. *TYPE: Lectotype male (538), Portobelo,

Colon, Panama, Nov 1913, E. Martini [BM; designation of Mattingly 1955:31]. Synony-

my with corrigani by Dyar (1922:95).

Culex (Anoedioporpa) corrigani of Bonne and Bonne-Wepster (1925: 188,260,264); Belkin (1968b:

12,14); Heinemann and Belkin (1977a:283; 1977b:452; 1978a:183); Knight and Stone (1977:

196).

Culex (Isostomyia) corrigani of Edwards (1932:328); Komp (1936:327); Lane (1939:74); Roze-

boom and Komp (1948:403); Galindo, Carpenter and Trapido (1951: 102,104,105,108,110,

111,112,113,126); Horsfall (1955:548).

Culex (Aedinus) corrigani of Dyar (1923b:189); Stone, Knight and Starcke (1959:282); Belkin,

Schick and Heinemann (1965:56).

Culex (Melanoconion) corrigani of Dyar (1925:158,161,169; 1928:347).

Culex (Micrades) corrigani of Gordon and Evans (1922:327); Dyar (1922:95,96; 1923:177).

Culex (Tinolestes) corrigani of Lane (1953 :392-393); Galindo and Blanton (1955:70).

Culex corrigani of Theobald (1910:614); Howard, Dyar and Knab (1915:222,223,229,386-387);

Galindo, Carpenter and Trapido (1955:159,161); Stone and Knight (1957:58); Forattini, Rabel-

lo and Cotrim (1970:39).

Culex (Micraedes) chalcocorystes of Dyar (1918:90,102).

FEMALE. Wing: 2.8 mm. Proboscis: 1.9 mm. Forefemur: 1.6 mm. Abdomen:

about 1.8 mm. Essentially similar to conservator, differing in the following. Head:

Decumbent scales on vertex white. Erect scales on occiput pale brown. Palpus

about 0.14-0.15 length of proboscis. Thorax: Acrostichal bristles represented by

3,4 short setae cephalad of prescutellar space. Antealar area above paratergite with

scattered white scales. Pleural integument pale yellow, finely spiculose. Lower mep

area bare.

MALE. Wing: 2.7 mm. Proboscis: 2.1mm. Forefemur: 1.75 mm. Similar to fe-

male in general coloration. Palpus about 0.14 length of proboscis.

MALE GENITALIA (fig. 24). As figured; diagnostic characters as in key. Seg-

ment IX: Tergite lobes distinct, moundlike, each with 6-9 short to moderately long

setae; interlobar space wide. Sidepiece: Roughly conical; length about 2.0-2.3 of

greatest width; tergal and sternal surfaces with shorter setae; lateral surface with long

bristles; tergomesal surface caudad of proximal division with a few short fine setae;

1 seta laterad of distal division of lobe usually lanceolate. Lobe: Proximal division

with a short stem bearing 2 rods with hooked tip, larger rod inserted at apex, smaller

slightly basad. Distal division poorly developed, wider than long, bearing | foliace-

ous seta, 4 specialized setae with recurved apexes and 2,3 shorter lanceolate setae.

Clasper: About 0.60 length of sidepiece, stout, smoothly curved; distal 0.3 of outer

surface with closely set spiculelike transverse ridges; 1 submedian and 1 subapical se-

ta on inner surface. Lateral Plate: Basal hook sclerotized, strongly curved, bearing a

few subapical spines; apical process narrow basally, broadly truncate at tip; sternal

spine short, pointed. Proctiger: Cercal sclerite long, broad, smoothly rounded at

apex, directed caudomesad; cercal setae 2-4. Paraproct with a row of 6-8 blunt api-

cal teeth,

PUPA (fig. 24). Abdomen: 2.4-2.7 mm. Trumpet: 0.45 mm; index 5.0-6.0. Pad-

dle: 0.55 mm. As figured; diagnostic characters as in key. Cephalothorax: Integu-

ment lightly pigmented. Hairs 1-3-C usually double; 4,5-C triple; 6,7-C double or

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 57

triple; 8,9-C single; 10-C at least 3-branched. Trumpet: Moderately pigmented and

strongly contrasting with cephalothorax. Abdomen: Integument lightly pigmented.

Hair 1-II short, weakly branched, easily distinguishable at 100X; hair 7-II usually sin-

gle; 4-VI short, weakly branched, cephalomesad of 5-VJ; hair 1-VI,VII at least dou-

ble; 9-VII at least 4-branched (4-6); hair 9-VIII usually with 7 branches (6-8). Pad-

dle: Lightly pigmented, longer than wide, about 2.0 of segment VIII. Male genital

lobe extending to 0.35 and female to 0.20 of paddle.

FOURTH INSTAR LARVA (fig. 25). Head: 0.85-0.90 mm. Siphon: 2.1-2.3 mm;

index about 20.0-30.0. Anal Saddle: 0.35-0.40 mm. As figured; diagnostic charac-

ters asin key. Head: Width about 1.1 of length. Subantennal pouch distinct, dome

shaped. Hair 4+C single; 5-C usually with 4 branches (3-5); hair 9-C usually with 4

branches (4-6); hair 14-C single; 15-C usually with 3 branches (2-4), short, not ex-

tending to base of mental plate. Mental plate with a strong median shouldered tooth

and 7 distinct teeth on either side. Antenna: Length about 0.5 of head, distinctly

spiculose to hair 1-A; all hairs single except 1-A (22,20-29). Thorax: Hair 0-P short,

weak, with dendritic branches; 1,2-P long, single; 3-P long, about 0.3 of 1-P, usually

single; 4-P usually with 3 branches (3,4); hairs 5,7-P long, single; 14-P short, single;

4-M usually double (1-3). Abdomen: Hair 3-I usually with 3 branches (2,3); hair 6-I

usually with 3 branches (2-4); hair 7-I single; 6-I] longer than segment, usually triple

(2,3); hair 6-III-V moderately long, always shorter than segment, triple; 1-V short,

never extending beyond segment V, weakly branched(4,3-6). Segment VIII: Comb

scales in a patch of 3,4 irregular rows, about 48-76 in number; individual scale with

spatulate fringed apex. Siphon: Very long, index variable. Integument moderately

pigmented, with a narrow basal dark ring; distinctly spiculose, spicules wartlike. Sub-

ventral hairs (1,la-S) usually 6 pairs (rarely 5), progressively smaller distad; subdorsal

hairs (2a-S) 2 pairs, located in distal 0.15 of siphon. Pecten short, extending barely

to proximal 0.15; individual tooth long, pointed apically, with minute ventral denti-

cles. Anal Segment: Integument strongly pigmented, imbricate and slightly spicu-

lose, spicules visible at 200X. Hair 1-X short, weakly branched; 2,3-X long, single.

Gills long; dorsal 2.0-2.5 length of saddle; ventral slightly shorter than dorsal.

SYSTEMATICS. Culex corrigani is distinct from all other members of the subge-

nus in both the adult and immature stages. In the adults, it is distinguished by the

combination of: (1) 3,4 acrostichal bristles near prescutellar area and (2) lower mep

bare; in the male genitalia by the combination of: (1) poorly developed distal divi-

sion of subapical lobe, (2) moderately broad, stout clasper, and (3) broad digitiform

cercal sclerite; in the pupa by the combination of: (1) hair 1-VII at least double and

(2) 4-VI cephalomesad of 5-VI; and in the larva by: (1) hair 1-V short, barely extend-

ing to apex of segment, multiple, (2) long siphon with an index of about 20.0-30.0,

and (3) usually 6 pairs of subventral siphon hairs (1, 1a-S).

We have seen material of corrigani from Nicaragua, Costa Rica and Panama, and

have not noticed any significant variations either in the adults or in the immature

stages.

We have also examined the type of chalcocorystes Martini 1914, and in our opin-

ion its synonymy with corrigani by Dyar (1922:95) is justified.

BIONOMICS. The immature stages of corrigani are usually found in treeholes.

According to Galindo, Carpenter and Trapido (1951:127), the immature stages show

preference for breeding in deep holes in trees growing in virgin forest. The larvae

were also found in bamboo traps near the ground and in the forest canopy. Nothing

is known about the behavior of the adult females.

58 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

DISTRIBUTION (fig. 12). Culex corrigani is Central American in distribution,

having been reported from Nicaragua (Atlantic), Costa Rica (Atlantic) and Panama

(Atlantic and Pacific) at low (near sea level) to moderate (600 m) elevations. The re-

cord from Colombia (Knight and Stone 1977: 196) is apparently in error.

Material examined: 194 specimens; 52 males, 45 females, 33 pupae, 64 larvae; 35

individual rearings (18 larval, 10 pupal, 7 incomplete).

COSTA RICA. Heredia: Puerto Viejo, Finca La Selva, OTS Field Station, 100 m, 8 Aug 1974,

A. Berrios A., 3 lpM (CR 433-30,-31,-33), 3 IpF (433-32,-34,-35), 1L (433) [UCLA].

PANAMA. Bocas del Toro: Almirante, 10 m, 27 Apr 1963, A. Quinonez, 1 pM (PA 259-1 19),

2L (259); 7 May 1963, A. Quinonez, 1 IpF (PA 335-107), 1 pM (335-110), 3 pF (335-103,-105,

-111), 1 1P (335-106), 2L [UCLA]. Canal Zone: Barro Colorado Is., 18 Jan 1935, 1Mgen [USN

M]; 7 May 1943, W.H.W. Komp, 1M (KO 37-18), 5M, 4F (KO 37-24), 1M (KO 41-9), 1M (KO 41-

18), 1F (KO 41-19), 1M, 1F (KO 41-32), 3M (KO 41-24), 1F (KO 41-27), 6M, 2F (KO 41-29), 1F

(KO 41-30), 1M, 1F (KO 41-33); 21 May 1943, W.H.W. Komp, 1M (KO 41-14), 1F (KO 41-15), 1

F (KO 41-16), 2F (KO 41-20), 1M (KO 41-23), 2F (KO 41-25), 1M, 1F (KO 207A-35); 21 May

1943, G.B. Fairchild, 2M (KO 37-40); 13 May 1945, 3L (5-336,-366); 15 May 1945, 1L (5-392);

18 May 1945, 1L (5-414); 23 May 1945, 4L (5-367,-382,-389,-406); 26 June 1945, 1L (5-411)

[UCLA] ; no date, det. J. Lane [FH, 6369,6370]. Fort Sherman, 5 Jan 1926, D. Baker, 1M, 3F

[USNM]. Mojinga Swamp, 5 m, 13 Oct 1964, A. Quinonez, 1 pF (PA 722-101), 3L (722) [UCL

A]. S. Gamboa Trail, 9 June 1943, Elton, 2M (KO 37-12) [UCLA]. Tabernilla, A. Busck, 1 L

[USNM], 1F [USNM, 10870; holotype of corrigani|. Colon: Portobelo, no data [FH, 8938];

18-24 Feb 1911, A. Busck, 1M, 11F; 25 Feb 1911, A. Busck, 1M, 2F; 13 Mar 1911, A. Busck, 3M

[USNM] ; Nov 1913, E. Martini, 1Mgen, 1F [USNM, 18466; type series of chalcocorystes| ; Nov

1913, E. Martini, 1M (538) [BM, lectotype]. Darien: El Real, Piriaque, near sea level, 13 Jan

1964, A. Quinonez, 4L (PA 622) [UCLA]. Morti, Morti Hydro, 80 m, 1 Dec 1966, O.G.W. Berlin

and R. Hinds, 1 pM (PA 967-100), 1L (967) [UCLA]. Pucro, Rio Tacarcuna valley, 600 m, 5 July

1963, A. Quinonez, 2 pM (PA 438-101,-103), 1 pF (438-102); 8 July 1963, A. Quinonez, 6 IpM

(PA 442-101,-103-105,-108,-110), 6 IP (442-102,-107,-109,-111-113), 1L (442) [UCLA]. Santa

Fe, 20 m, 22 Nov 1966, O.G.W. Berlin, 1 lpM (PA 945-11), 1 IpF (945-10) [UCLA]. Panama:

Juan Mina, 40 m, 18 Jan 1963, A. Quinonez, 1 lpM (PA 5-104), 1 IM (5-109), 2L (5) [UCLA]. Pa-

cora, 22 Dec 1950, S.J. Carpenter, 1L; 26 Dec 1950, S.J. Carpenter, 4L [UCLA]. Province not

specified: Santa Clara, La Venta, 9 Apr 1943, 1M, IF [USNM]. No data, det. Galindo [FH,

9011].

NICARAGUA. Zelaya: Bluefields, near sea level, 14 July 1964, A. Quinonez, 1 lpM (NI 48-10),

5L (48) [UCLA].

Restrictor Group

FEMALE. Small to moderate in size, inornate species. Head: Erect scales

forked apically, brown. Proboscis slightly longer than forefemur, entirely dark

scaled. Palpus short, about 0.20 length of proboscis, 4-segmented, segment 4 about

2.0 of segment 3. Antenna slightly shorter than proboscis. Thorax: Mesonotal

scales linear, auburn. Acrostichal bristles distinct, extending from anterior promon-

tory to prescutellar space. Pleural integument yellowish; scaling restricted to upper

ppn and stp; lower mep with | strong bristle. Abdomen: Tergites [I-VU with baso-

lateral light patches, rest dark scaled; sternites predominantly creamy, dark scaled

distally. |

MALE. Similar to female in coloration. Palpus long, subequal to proboscis

length, 5-segmented; segment 3 long; segment 4 short, about 0.30-0.33 of segment 3;

segment 5 shorter, about 0.60 of segment 4.

MALE GENITALIA. Segment IX: Tergite lobe moundlike, with 12-18 moderate-

ly long setae. Sidepiece: Roughly conical, length about 2.0 of greatest width. Lobe

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa x9

situated at 0.7, directed caudomesad, divided into distinct, approximated proximal

and distal divisions; proximal division with 2 distinct rods; distal division with 4 api-

cal leaves in addition to 2,3 specialized setae. Clasper: About 0.65 length of side-

piece, relatively simple and curved inward. Phallosome: Lateral plate with apical

process broadly truncate from dorsal aspect; sternal spine directed basolaterad.

Proctiger: Paraproct with 6-8 apical teeth; cercal setae 2-5.

PUPA. Cephalothorax: Hair 5-C moderately long, weakly developed, subequal

in length to 4-C; hair 10-C strongly developed, single or double, distinctly longer

than 11-C. Abdomen: Hair 1-II with 3-5 moderately long branches, not resembling

float hair (1-1); hair 1-IJI-VII varied in branching; 6-I,II long, about 2.0 of 7-I,II; hair

5-IV,V moderately long, double, barely extending to apex of succeeding segment; 2-

II-VI subequal in length to 9-II-VI; hair 9-VII,VIII strongly developed, at least 4-

branched; 9-VIII subequal in length to segment VIII; hair 1-[X absent. Posterior

margin of sternum VIII lobed laterad of 9-VIII. Paddle: Elongate, lightly pigment-

ed; hairs 1,2-P absent.

FOURTH INSTAR LARVA. Head: Hair 5-C usually 3-branched; 6-C single; 7-C

with 7-9 and 11-C with 4,5 branches. Central tooth of mental plate simple. Anten-

na: Hair 1-A inserted about 0.70 from base. Thorax: Hairs 5-7-P single, strongly

developed. Abdomen: Hair 6-III-V moderately long, about 0.75 length of 6-I,I];

hair 9-II-VI short, single. Segment VIII: Hair 2-VIII on an oval sclerotized plate.

Siphon: Subventral hairs (1,1la-S) 5 pairs; subdorsal hairs (2a-S) 5 pairs, middle ones

sometimes unpaired; proximal subdorsals always within last pecten tooth. Anal Seg-

ment: Ventral brush (4-X) with 6 pairs of hairs on a grid. Gills short, ventral slightly

shorter than dorsal.

DISCUSSION. The Restrictor Group is recognized here for the nominate form.

This species, originally described from Mexico, was included in the subgenus Micro-

culex by Stone, Knight and Starcke (1959:280) and Knight and Stone (1977:269).

On the basis of correlated features of adults and immatures, it appears to be more

closely related to species in the subgenus Anoedioporpa than to any other subgenus

of Culex in the New World. Within the subgenus, it is distinct from all the other

members in the adults and immature stages. The adults are readily distinguishable by

the presence of acrostichal bristles on the mesonotum from the anterior promontory

to prescutellar space. The pupa is easily separated by the longer, strongly developed

hair 10-C. The larva is diagnosed by the presence of 6 pairs of hairs in the ventral

brush, and a sclerotized plate at the base of hair 2 on segment VIII. This group has

retained more ancestral characters than the Conservator Group and is restricted to

the Middle American area.

The Restrictor Group is known at present only from the mainland of Central

America, from Mexico as far north as the states of Jalisco and San Luis Potosi, and

from the state of Aragua in Venezuela (fig. 27). The immature stages are found in

treeholes at low to moderately high (1700 m) elevations.

12. Culex (Anoedioporpa) restrictor Dyar & Knab

Figs, 27,28,29,30

1906. Culex restrictor Dyar & Knab 1906:222. *TYPE: Holotype larval skin (311c) with asso-

ciated whole female pupa on slide, Almoloya, Oaxaca, Mexico, 21 July 1905, F. Knab

[USNM]. Information in Belkin, Schick and Heinemann 1965:38.

1908. Culex consternator Dyar & Knab 1908:59. *TYPE: Lectotype male (429.1), Cordoba,

Veracruz, Mexico, 7 Mar 1908, F. Knab [USNM 11969; designation of Stone and Knight

1975:46]|. Synonymy with restrictor by Howard, Dyar and Knab 1915:333.

60 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Culex (Anoedioporpa) restrictor of Dyar (1923b:190); Heinemann and Belkin (1977a:283; 1977b:

427,433; 1977c:528; 1978a:183; 1978b:393).

Culex (Melanoconion) restrictor of Dyar (1925:169; 1928:348); Edwards (1932:217); Martini

(1935:60); Lane (1939:70).

Culex (Microculex) restrictor of Lane (1953:527-528); Galindo and Blanton (1955:73); Stone,

Knight and Starcke (1959:280); Belkin, Schick and Heinemann (1965:38); Diaz Najera and Var-

gas (1973:120); Knight and Stone (1977:269).

Culex (Micraedes) restrictor of Dyar (1918:102).

Culex (Aedinus) restrictor of Dyar (1923:189-190,191).

Culex restrictor of Dyar and Knab (1906:208); Howard, Dyar and Knab (1915:331); Galindo, Car-

penter and Trapido (1955:159,160); Forattini, Rabello and Cotrim (1970:47).

FEMALE (fig. 28). Wing: 3.2 mm. Proboscis: 2.0 mm. Forefemur: 1.7 mm. Ab-

domen: 2.2 mm. As described for subgenus and group, with the following additional

features. Head: Decumbent scales on vertex narrow, white; sides and venter with

broader white scales. Erect scales auburn. Palpus short, about 0.20 length of probos-

cis. Thorax: Mesonotal scales narrow, auburn except for white scales on anterior

promontory and along anterior part of lateral prescutal area. Antealar area above

paratergite with narrow auburn scales, Pleural integument yellowish. Upper margin

of ppn with auburn scales. Lower mep with a strong bristle. Abdomen: Scales on

tergites predominantly dark except for basolateral light areas. Sternites predominant-

ly creamy, dark scaled distally.

MALE (fig. 28). Wing: 2.7 mm. Proboscis: 1.8 mm. Forefemur: 1.5 mm. Simt-

lar to female in general coloration. Palpus subequal to length of proboscis.

MALE GENITALIA (fig. 29). As figured; diagnostic characters asin key. Readily

separated from all members of the subgenus by the presence of 4 apical leaves on dis-

tal division of subapical lobe. Segment IX: Tergite lobes distinct, widely separated,

each with 12-14 moderately long setae. Sidepiece: Roughly conical; length about

2.0 of greatest width; tergal surface with a few longer bristles; sternal surface with

shorter setae and a few scales basally. Lobe: Proximal division with a moderately

long stem bearing 2 rods with hooked tip, upper rod inserted at apex, lower slightly

basad. Distal division distinct, stem short, bearing apically 1 short and | longer spec-

ialized seta with hooked tip, and 4 leaves. Clasper: About 0.60 length of sidepiece,

smoothly curved mesad; 1 submedian and | subapical seta on inner surface. Lateral

Plate: Basal hook sclerotized, strongly curved; apical process long, broadly truncate

at tip; sternal spine short, pointed. Proctiger: Apex of cercal sclerite broad, triangu-

lar; cercal setae 2-8. Paraproct with a row of 6-8 blunt apical teeth.

PUPA (fig. 29). Abdomen: 2.5 mm. Trumpet: 0.45-0.50 mm; index about 7.0.

Paddle: 0.6 mm. As figured; diagnostic characters as in key. Cephalothorax: Light-

ly to moderately pigmented, wing case darker. Hairs 1-5-C moderately long, always

double; 6-C at least 3-branched (3-5); hair 7-C double; 8-C usually double; 9-C single;

10-C single or double, longer than 11-C; hairs 11,12-C single. Trumpet: Very strong-

ly pigmented and strongly contrasting with cephalothorax. Abdomen: Integument

lightly to moderately pigmented, progressively lighter caudad, imbricate. Hair 1-II

moderately long, extending beyond middle of succeeding segment, branched; 7-II

double; 4-VI short, usually double, cephalad of 5-VI; hair 1-VI,VII at least double (2-

5); hair 9-VII with 4-6 branches; 9-VIII with 4-7 branches. Paddle: Lightly pigment-

ed, longer than wide, about 2.0 of segment VIII and finely spiculose ventrally; apex

slightly produced. Male genital lobe extending to 0.4 and female to 0.2 of paddle.

FOURTH INSTAR LARVA (fig. 30). Head: about 1.0mm. Siphon: 1.9-2.2 mm;

index 16.0-19.0. Anal Saddle: 0.32-0.34 mm. As figured; diagnostic characters as in

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 61

key. Head: Width about 1.1 of length. Hair 4-C usually double; 5-C usually triple

(3-5); hair 9-C usually with 5 branches (4-7); hair 14-C double; 15-C with 3,4 branch-

es. Mental plate with a strong median tooth and 9-11 distinct teeth on either side.

Antenna: Length about 0.5 of head, distinctly spiculose to hair I-A; all hairs single

except 1-A (11,8-15). Thorax: Hair 0-P moderately long, weakly stellate, with 8-15

branches; 1,2-P long, single; 3-P moderately long, usually double (1-4); hair 4-P usu-

ally double (1-3); hairs 5-7-P single; 14-P small, single; 4-M double. Abdomen: Hairs

6,7-I, 6-II-V long, double; 6-VI single; 1-V moderately long, barely reaching apex of

segment, with 3-5 branches. Segment VIII: Comb scales in a patch of 4,5 irregular

rows, about 49-73 in number; individual scale with spatulate fringed apex. Siphon:

Integument moderately pigmented, darker basally, with fine wartlike spicules. Sub-

ventral hairs (1,la-S) 5 pairs, usually single, progressively smaller distad; subdorsal

hairs (2a-S) 5 pairs, median ones usually unpaired. Pecten extending barely to proxi-

mal 0.2; individual tooth long, pointed apically, without lateral denticles. Anal Seg-

ment: Integument moderately pigmented, imbricate, spiculose. Hair 1-X short,

weakly branched; 2,3-X long, single. Gills short, subequal to length of saddle, atten-

uated apically; ventral slightly shorter than dorsal.

SYSTEMATICS. Culex restrictor can be readily distinguished from other mem-

bers of Anoedioporpa in the adults by the presence of acrostichal bristles extending

from anterior margin to prescutellar area of mesonotum; in the male genitalia by the

short distal division of subapical lobe with 4 apical leaves in addition to other spec-

ialized setae; in the pupa by hair 10-C strongly developed, single or double and dis-

tinctly longer than 11-C; and in the larva by: (1) ventral brush (4-X) with 6 pairs of

hairs, (2) siphon with both subdorsal (2a-S) and subventral hairs (1,la-S) composed

of 5 pairs, and (3) hair 2-VIII on a sclerotized plate.

We have examined material of restrictor from Mexico, Guatemala, El Salvador,

Costa Rica, Panama and Venezuela, and have not come across any significant varia-

tions between populations. The characters that separate restrictor from the members

of Conservator Group are constant.

BIONOMICS. The immature stages of restrictor are usually found in treeholes

(31 collections), and occasionally in cut bamboo internodes (2 lots) or artifical con-

tainers (tires in 2 lots, gallon can in 1 lot). They have been collected at low (10 m)

to moderately high (1700 m) elevations. The immatures have been collected in as-

sociation with many typically treehole-breeding mosquitoes, including 10 different

species of Aedes (Protomacleaya), Culex (Culex) declarator Dyar & Knab 1906, Cx.

(Cux.) mollis Dyar & Knab 1906, Corethrella appendiculata Grabham 1906, Haema-

gogus (Haemagogus) equinus Theobald 1903, Hg. (Hag.) lucifer (Howard, Dyar &

Knab 1913), Hg. (Hag.) mesodentatus Komp & Kumm 1938, Orthopodomyia kum-

mi Edwards 1939 and Toxorhynchites (Lynchiella) moctezuma (Dyar & Knab 1906).

Nothing is known about the behavior of the adult females.

DISTRIBUTION (fig. 27). Culex restrictor is primarily Central American in dis-

tribution. We have seen specimens from Mexico as far north as the states of Jalisco

(Pacific side) and San Luis Potosi (Atlantic side), from Guatemala, El Salvador, Hon-

duras, Costa Rica and Panama. It has also been collected from Venezuela, from 2 ad-

jacent localities just N of Maracay.

Material examined: 973 specimens; 121 males, 124 females, 262 pupae, 466 larvae;

148 individual rearings (98 larval, 40 pupal, 10 incomplete).

COSTA RICA. Alajuela: Alajuela, 880 m, 18 July 1971, D.W. Heinemann, 7L (CR 270) [UCL

A]; no data [FH, 8931]. Desamparados, Rio Machuca, 260 m, 1 Nov 1971, D. Schroeder, 1 lpM

(CR 497-11), 4 IpF (497-12-15), 3 pM (497-100-102), 6L (497) [UCLA]. Desmonte, 600 m, 2

62 Contrib. Amer. Ent. Inst., vol. 17, no. 2, 1980

Aug 1971, A. Berrios A., 1L (CR 330) [UCLA]. Cartago: Orosi, 1070 m, 19 July 1971, J.N. Bel-

kin, 2 lpM (CR 280-10,-11), 2 lpF (280-12,-13) [UCLA]. Guanacaste: Puerto Humo, Palo Verde

OTS Field Station, 10 m, 18 Aug 1971, S.J. and D.W. Heinemann, 1 pM (CR 443-102), 2 pF (443-

100,-101), 10L (443), 1 IpF (CR 444-11), 2 IP (444-10,-12), 8L (444); 19 Aug 1971, D.W. Heine-

mann, 2 lpM (CR 449-10,-11), 5 lpF (449-12-14,-16,-18), 3 pM (449-100-102), 2 IP (449-15,-17),

20L (449); 20 Aug 1971, D.W. Heinemann, 6 lpM (CR 452-21-23,-25-27), 8 IpF (452-20,-24,-28-

33), 4 pM (452-100,-101,-103,-109), 3 pF (452-102,-107,-108), 1L (452) [UCLA]. Puntarenas:

El Roble, 16 June 1943, T.H.G. Aitken, 1F (KO 207B-12) [UCLA]. San Jose: Limonal, 1080 m,

4 Aug 1971, D. Schroeder and S.J. Heinemann, 2 IpM (CR 335-10,-11), 2 IpF (335-14,-16), 2 pF

(335-100,-101), 3L (335) [UCLA]. San Isidro del General, 750 m, 20 June 1964, C.L. Hogue, 1

pF (CR 176-102), 1P (176) [UCLA].

EL SALVADOR. Sonsonate: Izalco, 430 m, 6 Nov 1971, J.N. Belkin and S.G. Breeland, 1 IpF

(SAL 53-50) [UCLA]. Los Planes, det. W. Kumm [FH, 4271,6310]. Sonsonate, Finca San Dioni-

sio,4 Aug 1964, A. Quinonez, 1 lpF (SAL 8-20), 1 pF (8-101), 3L (8) [UCLA]. Sonsonate, Can-

ton El Castano, 300 m, 1 Aug 1964, A. Quinonez, 1 IpF (SAL 1-20), 1L (1), 1 IpM (SAL 3-11), 3

IpF (3-13-15), 1P, 24L (3) [UCLA]. Usulutan: San Juan del Gozo, H.W. Kumm, 3M, 2F (KO

207A-28) [UCLA].

GUATEMALA. Chimaltenango: Locality unspecified, Jun-Oct 1950, H.T. Dalmat, 4 lpF (GUA

151-23,-24,-26,-28), 8M (151-10,-11,-13,-14,-19,-22,-29,-31), 11F (151-12,-15-18,-20,-25,-27,-30,

-32,-33), 4P (151) [UCLA]. Escuintla: Escuintla, 160-180 m, 10 July 1964, J. and T. Zavortink,

6 IpF (GUA 39-10,-11,-13,-14,-16,-18), 1 lp (39-17), 3 pM (39-101-103), 4 pF (839-105-108), 2 1P

(39-12,-15), 3M, 3F, 11P, 7L (39), 3 lpM (GUA 40-20-22) [UCLA]. San Jose de Guatemala

[=Puerto de San Jose], 16 July 1943, D.G. Hall, 2M (KO 41-2); July1943, D.G. Hall, 2F (KO 37-26,

-39), 2M (GUAK 27) [UCLA]. Guatemala: Bethania, Rio El Naranjo, 1500 m, 15 July 1964, T.

Zavortink and P. Cowsill, 2 lpF (GUA 44-10,-11), 1 IP (44-12), 2L (44) [UCLA]. Guatemala City,

1500 m, 2 Sept 1964, W. Almengor and P. Cowsill, 2L (GUA 118) [UCLA]. Villa Canales, 1300

m, 22 July 1964, W. Almengor and P. Cowsill, 1L (GUA 60) [UCLA]. Retalhuleu: San Felipe,

580 m, 2 July 1964, P. Cowsill, 6 IlpF (GUA 22-20,-22-26), 1 IF (22-21), 6F, 6P, 6L (22) [UCLA].

HONDURAS. Atlantida: Lancetilla, 50m, 19 Aug 1964, A. Quononez, 1L (HON 55-10)

[UCLA].

MEXICO. Guerrero: Chilpancingo, 34 km S of, ?900 m, 7 Aug 1966, D. Schroeder, 1L (MEX

424), 2L (MEX 426); 38 km S of Chilpancingo, ?700 m, 8 Aug 1966, D. Schroeder, 6L (MEX

420) [UCLA]. Jalisco: El Tuito, 19 km N of Boca de Tomatlan, 610 m, 30 Aug 1972, J.N. Bel-

kin, 16 lpM (MEX 722-10-15,-19,-22,-24-29,-80,-81), 8 lpF (722-16-18,-20,-21,-23,-82,-83), 1 pM

(722-101), 2 pF (722-102,-103), 48M, 16F, 80P, 172L (722) [UCLA]. Quililla, El Mirador, 1700

m, 8 June 1971, L.T. Nielsen and T. Zavortink, 4 lpF (MEX 671-30-33) [UCLA]. Oaxaca: Almo-

loya, 21 July 1905, F. Knab, 1 IP (311c) [USNM, holotype of restrictor]. Salina Cruz, ?10 m, 9

Aug 1966, D. Verity, 1P (MEX 105-10), 2L (105) [UCLA]. Tehuantepec, Puentes Tortugas, 100

m, 16 Aug 1966, D. Schroeder, 1 pM (MEX 436-100), 1 IP (436-10), 12L (436), 2 lpM (MEX 437-

10,-11), 1L (437) [UCLA]. San Luis Potosi: Tamazunchale, 200 m, 20 July 1965, R.X. Schick

and D. Schroeder, 1 IpM (MEX 213-21), 1 lpF (213-90), 1M (213-20), 27L (213); 21 July 1965,

R.X. Schick and D. Schroeder, 1L (MEX 233) [UCLA]. Veracruz: Cordoba, 900 m, 18 July

1964, E. Fisher, 1L (MEX 52); 22 July 1964, E. Fisher and D. Verity, 1L (MEX 70) [UCLA] ; 7

Mar 1908, F. Knab, 1M (429.1) [USNM, 11969; lectotype of consternator] .

PANAMA. Bocas del Toro: Highlands of Chiriqui above 1000 m (Galindo and Blanton 1955:

73). Canal Zone: Corozal Dam, 28 July 1943, 1F (KO 41-21) [UCLA]. Cocle: El Valle, 600 m,

13 Aug 1963, A. Quinonez, 1 pM (PA 507-101) [UCLA]. Darien: Pucro, northern slopes of Cerro

Mali, 1470 m, 27 May 1963, A. Quinonez, 1 lp (PA 366-107) [UCLA].

VENEZUELA. Aragua: Guamitas, 740 m, 15 July 1969, J. Bergland and T. Zavortink, 2 lpF

(VZ 198-30,-32), 1 pM (198-102), 4 pF (198-100,-101,-103,-104), 1 lp (198-31), 12P, 16L (198);

18 Aug 1969, J. Clavijo and J. Pulido, 3L (VZ 374) [UCLA]. El Castano, 600 m, 20 Aug 1969, J.

Clavijo and J. Pulido, 9L (VZ 380) [UCLA].

Berlin and Belkin: Subgenera Aedinus, Tinolestes and Anoedioporpa 63

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