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Contributions<br />

of the<br />

American Entomdogiccd Institute<br />

Volume 5, Number 2, 1970<br />

MOSQUITO STUDIES (<strong>Diptera</strong>, <strong>Culicidae</strong>)<br />

<strong>XIX</strong>. <strong>The</strong> <strong>treehole</strong> <strong>Anopheles</strong> of the New World<br />

By Thomas J. Zavortink


CONTRIBUTIONS<br />

of the<br />

AMERICAN ENTOMOLOGICAL INSTITUTE<br />

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Entomological Institute, 5950 Warren Road, Ann Arbor, Michigan, U. S. A. 48105<br />

No. 1.<br />

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No. 1.<br />

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Dasch, Clement E., 1964. <strong>The</strong> Neotropic Diplazontinae (Hymenoptera, Ichneumonidae).<br />

75 pages, 69 figures. Price: $2.25, postpaid.<br />

<strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1965.<br />

I. Belkin, John N. , -- et al. A project for a systematic study of the mosquitoes of<br />

Middle America.<br />

II. Belkin, John N., et al.<br />

-- Methods for the collection, rearing and preservation of<br />

mosquitoes.<br />

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Matthews, Robert W., I965. <strong>The</strong> biology of Heriades carinata Cresson. (Hymenoptera,<br />

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<strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1965.<br />

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IV. Belkin, John N. <strong>The</strong> mosquitoes of the Robinson-Peabody Museum of Salem<br />

expedition to the southwest Pacific, 1956.<br />

34 pages, 3 figures. Price: $1.00, postpaid.<br />

<strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1965.<br />

V. Belkin, John N. , Robert X. Schick, and Sandra J. Heinemann.<br />

<strong>Mosquito</strong>es originally described from Middle America. 95 pages. Price: $2.75 postpaid.<br />

<strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1966.<br />

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<strong>Mosquito</strong>es originally described from North America. 39 pages. Price: $1.00, postpaid.<br />

DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae<br />

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Thailand. 55 pages, 20 figures. Pricem5, postpaid.<br />

Parts of Volume 2, with prices<br />

Contributions to the mosquito fauna of southeast Asia, 1967.<br />

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296 pages, 104 figures. Price: $9.00, postpaid.<br />

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Snider, Richard J. , 1967. <strong>The</strong> chaetotaxy of North American Lepidocyrtus s. str.<br />

-7<br />

(Collembola: Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid.<br />

Contributions to the mosquito fauna of southeast Asia, 1968.<br />

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southeast Asia. 76 pages, 26 figures. Price: $2.25, postpaid.<br />

Contributions to the mosquito fauna of southeast Asia, 1968.<br />

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Aedes, subgenus Finlaya <strong>The</strong>obald. 45 pages, 12 figures. Price: $1.50, postpaid.<br />

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No. 1. <strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1968.<br />

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182 pages, 30 figures. Price: $5.50, postpaid.<br />

No. 2. <strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1968.<br />

VIII. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia.<br />

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IX. Belkin, John N. <strong>The</strong> type specimens of New World mosquitoes in European<br />

museums. 69 pages. Price: $2.00, postpaid.<br />

(Continued on back cover)


MOSQUITO STUDIES (<strong>Diptera</strong>, <strong>Culicidae</strong>)<br />

<strong>XIX</strong>. THE TREEHOLE ANOPHELES OF THE<br />

NEW WORLD’<br />

Thomas J. Zavortink2<br />

CONTENTS<br />

INTRODUCTION . . . . . . . . . .<br />

SYSTEMATICS . . . . . . . . . . .<br />

TAXONOMIC TREATMENT<br />

Description of Group . . . . . . . .<br />

Keys to Species . . . . . . . . . .<br />

1. <strong>Anopheles</strong> (An.) barberi Coquillett<br />

2. <strong>Anopheles</strong> (An.) judithae Zavortink.<br />

.<br />

3. <strong>Anopheles</strong> (An.) fausti Vargas . . .<br />

4. <strong>Anopheles</strong> (An.) arboricolus, n.sp. .<br />

5. <strong>Anopheles</strong> (An. ) xelajuensis de Leon<br />

6. <strong>Anopheles</strong> (An.) powderi, n.sp. . .<br />

REFERENCES CITED . . . . . . . .<br />

FIGURES<br />

TABLE<br />

. . . . . . . . . . . . .<br />

INDEX TO SCIENTIFIC NAMES 1 1 1 1<br />

INTRODUCTION<br />

.............. 1<br />

.............. 2<br />

.............. 3<br />

.............. 5<br />

.............. 7<br />

.............. 10<br />

.............. 12<br />

.............. 14<br />

.............. 15<br />

.............. 17<br />

.............. 18<br />

.............. 21<br />

.............. 34<br />

.............. 35<br />

<strong>The</strong> present paper is a review of the obligate <strong>treehole</strong> breeding <strong>Anopheles</strong> of the<br />

New World. It is essentially an expansion of my previous study of the <strong>treehole</strong><br />

<strong>Anopheles</strong> of the United States (Zavortink, 1969).<br />

<strong>The</strong> taxonomic procedure is that outlined for the project “<strong>Mosquito</strong>es of Middle<br />

America” by Belkin, Schick et al (1965: 10-l 1). Format, abbreviations and other<br />

details follow earlier papers in this series. I have not seen the types of barberi, fausti,<br />

or xelajuensis.<br />

I am indebted to the following individuals for loans of specimens included in this<br />

‘Contribution from project “<strong>Mosquito</strong>es of Middle America” supported by U.S. Public Health<br />

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command<br />

Research Contract DA-49-193-MD-2478.<br />

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


2 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

study: Alfonso Diaz Najera of the Instituto de Salubridad y Enfermedades Tropi-<br />

tales, Mexico [ ISET] ; Lewis T. Nielsen of the University of Utah [UTAH] ; L.L.<br />

Pechuman of Cornell University [CU] ; Lloyd E. Rozeboom of Johns Hopkins Uni-<br />

versity [HOPK] ; and Alan Stone of the United States National Museum [USNM] .<br />

Peter F. Mattingly kindly loaned specimens of extralimital species from the British<br />

Museum (Natural History) collection.<br />

I thank John N. Belkin for editing the manuscript and the following individuals,<br />

all present or past members of the “<strong>Mosquito</strong>es of Middle America” project staff,<br />

for technical assistance: Caryle Abrams, Sheila Bernstein, Sally Dieckmann, Frances<br />

DuPont, Sandra Heinemann, Christopher Ishida, L. Margaret Kowalczyk, Nancy<br />

Martsch, Michael Nelson and William Powder.<br />

SYSTEMATICS<br />

<strong>The</strong> American <strong>treehole</strong> breeding <strong>Anopheles</strong>, barberi Coquillett, 1903, fausti Var-<br />

gas, 1943, judithae Zavortink, 1969, xelajuensis de Leon, 1938, and 2 species des-<br />

cribed here, arboricolus and powderi, belong to a single phylad. At least 2 of the<br />

numerous Old World species of container breeding AnopheEes, barianensis James,<br />

1911 and plumbeus Stephens, 1828, belong to this same line. This group of 8<br />

species is differentiated morphologically and biologically, by its usual restriction to<br />

temperate or montane tropical regions, from the other phylads of container breed-<br />

ing anophelines.<br />

As members of this group have some rather unusual features in the preimaginal<br />

stages, 3 generic group taxa have been proposed for included species: CoeZodiazesis<br />

Dyar & Knab, 1906 founded on larval characteristics of barberi; CycZophorus Eysell,<br />

19 12 on the immature stages of pZumbeus; and RusseZZia Vargas, 1943 on larval pe-<br />

culiarities of xelajuensis. Despite its distinctiveness, this group of species is not cur-<br />

rently recognized at the subgeneric level and is placed in <strong>Anopheles</strong> (<strong>Anopheles</strong>)<br />

(Stone, Knight and Starcke, 1959). This is because anophelines of several distinct<br />

phylads adapted to breeding in plant containers show convergence in some of the<br />

more conspicuous elements of the larval chaetotaxy, such as reduction of the front-<br />

al hairs of the head and elongation of lateral abdominal hair 6-IV-VI, and, as a con-<br />

sequence, students of the genus have been reluctant to use features of the immature<br />

stages to define or place them.<br />

On the basis of morphology of the adult, male genitalia and larva, geographical<br />

distribution and habitat, the 6 species included here fall into at least 3 different<br />

primary groups: 1) barberi and judithae, 2) fausti and arboricolus and 3) xelajuensis.<br />

<strong>The</strong> affinities of the poorly known powderi are uncertain. <strong>The</strong> geminate species bar-<br />

beri and judithae differ most conspicuously from the others in the following fea-<br />

tures: as adults by 1) interocular scales absent or short, 2) erect head scales not<br />

brilliant white mesally and very dark laterally, 3) mesonotum shortened and arched,<br />

4) mesonotum without broad hoary median longitudinal stripe, 5) acrostichal scal-<br />

ing absent or reduced, 6) coxae more or less same color as adjacent portions of<br />

pleuron, 7) femora and tibiae entirely dark scaled and 8) wing fringe dark; in the<br />

male genitalia by the scaleless sidepieces; as larvae by hair 3-C mesad of 4-C. <strong>The</strong>se<br />

species are widespread in various types of forest in North America. <strong>The</strong> group con-<br />

sisting of fausti and arboricolus differs as follows: in the adult stage by 1) light<br />

patches on palpus, 2) hindfemur but not hindtibia conspicuously marked with white<br />

and 3) wing with a single light fringe spot at ends of branches of vein R; in the<br />

male genitalia by 1) sparsely scaled sidepiece, 2) clumped spicules on sidepiece and


Zavortink: Treehole <strong>Anopheles</strong> of New World 3<br />

3) outermost 2 setae on ventral lobe of claspette subequally developed; in the larva<br />

by plumose hairs 1 l-C, 9-III-VI, 4-IV,V and lo-VII. Both species have restricted<br />

distributions, fausti in the tropical rainforest at the base of the Sierra Madre Oriental<br />

in central Mexico and arboricozus in the humid montane forest of western Panama.<br />

An. xelajuensis is distinguished as follows: first, by its large size in all stages,<br />

then, in the adult by 1) hindfemur and hindtibia both conspicuously marked with<br />

white, 2) light patch over apex of costa and vein R, , 3) wing with 4 light fringe<br />

spots and 4) wing veins with dark spots caused by accumulation of scales; in the<br />

male genitalia by 1) long sidepiece and 2) numerous scales on sidepiece; in the larva<br />

by 1) hair 3-C thick and blunt, 2) absence of palmate hairs and 3) presence of many<br />

stellate hairs. It is apparently limited to areas of wet montane forest in southern<br />

Mexico and Guatemala. An. powderi, unknown in the male and larva, is large in<br />

size, has a single light fringe spot extending from Rr to Mr +2 and has only the<br />

hindfemur conspicuously marked with white. It has been collected in the montane<br />

cloud forest of southern Costa Rica. <strong>The</strong> differences between these groups are summarized<br />

in Table 1. I am not recognizing these groups even informally because the<br />

knowledge of the Central American species is still very fragmentary and I have not<br />

studied the Old World species in detail.<br />

Very little is known about the bionomics of species in this group. <strong>The</strong> immature<br />

stages are restricted to <strong>treehole</strong>s or those artificial containers which approach them<br />

in water composition. Larval development is unusually long. Larvae of barberi are<br />

predacious and those of other species readily eat insect fragments. Adults of both<br />

sexes are occasionally found resting in buildings, culverts and hollow trees and females<br />

of barberi and xelajuensis bite man. Stratman-Thomas and Baker (1936: 182-<br />

183) infected barberi with Plasmodium vivax and demonstrated transmission of the<br />

malaria to another person. However, this species has not been incriminated as a natural<br />

vector. American <strong>treehole</strong> AnopheZes extend from the northern United States<br />

(South Dakota to New York) to northern Panama (Chiriqui). All six known species<br />

are apparently allopatric.<br />

Since An. eiseni Coquillett, 1902, a Neotropical species unrelated to those treated<br />

here, is occasionally found breeding in <strong>treehole</strong>s, it has been included in the keys.<br />

TAXONOMIC TREATMENT<br />

DESCRIPTION OF GROUP<br />

FEMALES. Head: Adorned with erect scales only; labium entirely dark scaled;<br />

palpus long, about 0.851 .lO length of proboscis; torus and first flagellar segment<br />

with a few small dark scales. Thorax: Mesonotal bristles quite long; mesonotal scales<br />

absent or restricted to a median tuft on anterior promontory or an acrostichal line;<br />

apn, sp, pra, ppl, stp and upper mep bristles or hairs present, those on stp in a more<br />

or less continuous vertical row; pleuron without scales. Legs: Coxae without scales;<br />

femora and tibiae never mottled, light scales completely absent or restricted to small<br />

patches or rings at apex of segments; tarsi entirely dark scaled; tarsal segment 5 of<br />

all legs much shorter than segment 4. Wing: Never with conspicuous pattern of<br />

light patches or speckles, light scales completely absent or restricted to fringe and<br />

apex of anterior veins. Haltere: Knob dark scaled. Abdomen: Scales completely<br />

absent. Buccopharyngeal Armature: Not studied.<br />

MALES. Similar to females except for sexual differences. Head: Flagellum strongly<br />

plumose.


4 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

MALE GENITALIA. Sidepiece: A single strongly developed internal spine and 2<br />

strongly developed parabasal spines present; lateral parabasal spine longer than mes-<br />

al; scales absent, few or numerous. Claspette: Dorsal lobe sclerotized, bearing 3<br />

(rarely 4) overlapping spatulate apically curved setae; ventral lobe conical, spicu-<br />

lose mesally, bearing 3 long and 1 short setae. Clasper: Usually longer than sidepiece.<br />

Phallosome: Aedeagus without leaflets, apex rounded.<br />

PUPAE. Setae generally weakly to moderately developed, relatively short, fine,<br />

few branched and lightly pigmented. Cephalothorax: Lightly to moderately pig-<br />

mented, with middorsal area, region caudad of trumpet and upper portion of wing-<br />

case slightly darker; hairs 6,7(: relatively close together, separated by a distance<br />

less than 0.5 that between 45-C; hair 7-C distinctly smaller than 6-C. Trumpet:<br />

Largely light to dark amber or brown in color, base lighter; rather “culicine” in<br />

shape, not strongly flared or deeply divided, the pinna relatively small. Abdomen:<br />

Generally lightly to moderately pigmented, with anterior and/or lateral portions of<br />

most segments slightly darker; hair 9-I small, subequal to or shorter than 3-1, usually<br />

single (single, double); 2-11 relatively short, single; 1 -III-VII and 5-111,IV frequently<br />

single, never with more than 5 fine branches, less than 0.4 length of corresponding<br />

segment; S-V-VII similar to 5-111,IV or thickened and then sometimes elongate; 9-<br />

IV-VIII spinelike, long, slender, pointed, becoming progressively longer on poster-<br />

ior segments, usually simple but sometimes with a few coarse branches near tip on<br />

segments VII and VIII. Terminal Segments: Hair I-IX not irregular as in most other<br />

Anophelini, but instead a simple single or double seta. Paddle: Lightly to moder-<br />

ately pigmented; more or less elliptical to oboval in shape, never more than slight-<br />

ly emarginate; outer part wider than inner; external buttress long; outer margin, to<br />

beyond end of external buttress, serrate, with teeth becoming more prominent and<br />

numerous apically; remainder of margin smooth or weakly serrate, never with long<br />

spicules; hair 1 -P thickened, spinelike.<br />

FOURTH INSTAR LARVAE. Head: Setae generally poorly developed; head long-<br />

er and collar narrower than in most Anophelini; maxillary sutures very short; mod-<br />

erately to deeply pigmented, tan to brown, with subantennal area and collar dark-<br />

er and ocular area lighter, never with pattern of contrasting colors; hairs 3-10-C rel-<br />

atively short, fine, usually single (1-4b or f), never large and plumose; 11-C weak-<br />

ly developed, short, usually not extending beyond level of hair 1-A. Antenna: Short,<br />

frequently smooth; uniformly moderately to deeply pigmented, tan to brown, or<br />

becoming lighter apically; hair 1 -A short, fine, single, arising on dorsolateral surface<br />

near middle of shaft. Thorax: Integument without conspicuous branched spicules;<br />

hair O-P apparently not developed; 1 l-P,M,T subequal, very small; 9-M,T not short<br />

and stout, at least as long as hair 10 of corresponding segment; 3-T never palmate.<br />

Abdomen: Hair 1-I not palmate, very small, shorter than hair 2 (except in xela-<br />

juensis); l-II-VII or 1 -III-VII (xelajuensis excepted) palmate, the leaflets well de-<br />

veloped, broad, usually serrate apically; 6-I-VI long, plumose, nearly equally devel-<br />

oped on all segments or 6-III-VI or 6-IV-VI less strongly developed; 4-IV,V (except in<br />

xelajuensis) elongate, longest hair on dorsal surface of respective segment; 5-IV-VII<br />

(except in xelajuensis) elongate, with relatively few weak and short lateral branches.<br />

Spiracular Lobe: Pecten with most teeth, except apically, long, never with regular<br />

sequence of long and short teeth; hair 1 -S usually single (l-3b) or with fine apical<br />

branches, never with numerous long branches from base.


Zavortink: Treehole <strong>Anopheles</strong> of New World<br />

KEYS TO SPECIES<br />

ADULTS<br />

1. Hindtibia, but not hindfemur, with a conspicuous broad apical white ring;<br />

vein R sometimes with light patches at level of ends of veins 1A and Rs<br />

(see Systematics) . . . . . . . . . . . . . . . . . . . . eiseni<br />

Hindtibia without a conspicuous broad apical white ring, apex dark or with<br />

a conspicuous white patch; hindfemur sometimes with a conspicuous api-<br />

cal white ring; vein R without light patches at level of ends of veins 1A<br />

andRs. . . . . . . . . . . . . . . . . . . . . . . . . .2<br />

2( 1). Mesonotum without a broad hoary median longitudinal stripe and with scales<br />

absent or restricted to a tuft in center of anterior promontory; hindfemur<br />

entirely dark scaled; wing fringe entirely dark scaled . . . . . . . . 3<br />

Mesonotum with a broad hoary median longitudinal stripe and a long acros-<br />

tichal scale line; hindfemur with apical white ring; wing fringe with api-<br />

cal light spot . . . . . . . . . . . . . . . . . . . . . . .4<br />

3(2). Bristles on ppl 6-l 1; anterior forecoxal bristles 19-32; anterior acrostichal<br />

bristles dark and usually not accompanied by scales; light erect head scales<br />

dingy yellowish (fig. 2). . . . . . . . . . . . . . . . .l. barberi<br />

Bristles on ppZ2-5 ; anterior forecoxal bristles 6-l 8; anterior acrostichal brist-<br />

les amber and accompanied by 5-20 whitish scales; light erect head scales<br />

pale white. . . . . . . . . . . . . . . . . . . . . 2. judithae<br />

4(2). Apex of costa and vein R, light scaled; small light fringe spots present at<br />

ends of veins M 1 +2 , M, +4 and Cur ; dark wing spots present (fig. 12). .<br />

. . . . . . . . . . . . . . . . . . . . . . . . 5. xelajuensis<br />

Apex of costa and vein Ri dark scaled; without fringe spots at ends of veins<br />

M 1+29 M3+4 and Cu 1 ; wing without dark spots . . . . . . . . . . 5<br />

5(4). Palpus with light scales at joints and apex; wing moderately scaled; scales in<br />

middle of vein 1A clasping vein or slightly spreading (fig. 12) . . . . .<br />

. . . . . . . . . . . . . . . . . . . . 3. fausti; 4. arboricolus<br />

Palpus entirely dark scaled; wing profusely scaled; scales in middle of vein<br />

IA wide spreading (fig. 11) . . . . . . . . . . . . . . 6. powderi<br />

MALE GENITALIA<br />

(6. powderi unknown)<br />

1. Aedeagus with 1 pair of conspicuous serrate leaflets; dorsal claspette lobe<br />

with 2 terete setae (see Systematics) . . . . . . . . . . . . eiseni<br />

Aedeagus without leaflets; dorsal claspette lobe with 3 flattened setae . . 2<br />

2( 1). Sidepiece long, scales numerous (fig. 9) ......... 5. xelajuensis<br />

Sidepiece shorter, scales absent or few ............... 3<br />

3(2). Lateral 2 setae of ventral lobe of claspette subequal in development; spicules<br />

of sidepiece grouped into large clumps; sidepiece usually with at least 1 or


6 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

2 scales (fig. 12) . . . . . . . . . . . . . 3. fausti; 4. arboricolus<br />

Lateral seta of ventral lobe of claspette not as strongly developed as adjacent<br />

seta; spicules of sidepiece arranged singly, in rows, or in small clumps; sidepiece<br />

usually without scales . . . . . . . . . . . . . . . . . . 4<br />

4(3). Parabasal spines much more strongly developed than internal spine; lateral<br />

parabasal spine flattened preapically; sidepiece usually more or less ellip-<br />

tical in shape (fig. 3) . . . . . . . . . . . . . . . . .l. barberi<br />

Parabasal spines only slightly stronger than internal spine; lateral parabasal<br />

spine not flattened preapically ; sidepiece usually more or less cylindrical<br />

or short conical in shape (fig. 5) . . . . . . . . . . . . 2. judithae<br />

PUPAE<br />

(3. fausti and 4. arboricolus unknown)<br />

1. Trumpet deeply divided; paddle margin with fringe of long, fine spicules (see<br />

Systematics) . . . . . . . . . . . . . . . . . . . . . . eiseni<br />

Trumpet not deeply divided; paddle margin without fringe . . . . . . . 2<br />

2( 1). Hair 7-VII single, longer than 9-VIII; hair O-VII enlarged, multiple; 5-V more<br />

strongly developed than 1-V (fig. 9). . . . . . . . . . 5. xelajuensis<br />

Hair 7-VII 2,3b (l-4), shorter than 9-VIII; hair O-VII small, single (single,<br />

double); S-V usually less strongly developed than 1-V . . . . . . . . 3<br />

3(2). Large species, width of segment VIII greater than 1 .OO mm (fig. 11) . . .<br />

6. powderi<br />

Smali species, \;idth of segment VIII less than b.85 ‘mm : : : : . . . .4<br />

4(3). Hair 9-III-VIII darkly pigmented; 9-111 more similar in size to 9-IV than 9-11;<br />

hair 5-VI,VII thickened, single (fig. 3) . . . . . . . . . . .l. barberi<br />

Hair 9-III-VIII concolorous with integument; 9-111 intermediate in size be-<br />

tween 9-11 and 9-IV; hair S-VI and usually 5-VII fine, 2,3b (fig. 5) . . .<br />

. . . . . . . . . . . . . . . . . . . . . . . . . 2. judithae<br />

FOURTH INSTAR LARVAE<br />

(6. powderi unknown)<br />

1. Hairs 4-6-C long, plumose; 3-T palmate (see Systematics) . . . . . eiseni<br />

Hairs 4-6-C shorter, never plumose; 3-T not palmate . . . . . . . . . 2<br />

2( 1). Hair l-III-VII stellate; 9-III-VI very strongly developed, stellate, long, 6,7b (5-<br />

9); hair 3-C thickened, apex usually blunt (fig. 10) . . . . 5. xelajuensis<br />

Hair l-III-VII palmate; 9-III-VI moderately developed, not stellate, elongate<br />

and plumose or shorter and 24b (l-5); hair 3-C fine, apex attenuate . . 3<br />

3(2). Hair 3C mesad of 4-C; hair 9-III-VI not elongate or plumose; 7-VII much<br />

more strongly developed than 6-VII . . . . . . . . . . . . . . 4<br />

Hair 3-C laterad of 4-C; hair 9-III-VI elongate, plumose; 7-VII not as strongly<br />

developed as 6-VII . . . . . . . . . . . . . . . . . . . . .5


4(3).<br />

5(3).<br />

Zavortink: Treehole <strong>Anopheles</strong> of New World 7<br />

Inner clypeals (2C) widely spaced, separated by a distance greater than 2.0<br />

that between inner and outer clypeals (3-C); hair 13-II-V,VII usually 3b (3-<br />

5); integument on underside of prothorax and abdominal segments I-VIII<br />

spiculose (fig. 4) . . . . . . . . . . . . . . . . . . .l. barberi<br />

Inner clypeals (2-C) closely approximated, separated by a distance less than<br />

that between inner and outer clypeals (3-C); hair 13-II-V,VII usually single<br />

(I-3b); integument of thorax and abdomen not spiculose (fig. 6) . . .<br />

. . . . . . . . . . . . . . . . . . . . . . . . . 2. judithae<br />

Hair 9-I,11 elongate, plumose; 13-11-V with many fine inconspicuous branches<br />

beyond middle; antenna1 shaft with conspicuous spicules (fig. 7) . . .<br />

. . . . . . . . . . . . . . . . . . . . . . . . . . 3. fausti<br />

Hair 9-I,11 not elongate or plumose; 13-11-V with 3,4 (2-6) conspicuous<br />

branches from base; antenna1 shaft without conspicuous spicules (fig. 8)<br />

. . . . . . . . . . . . . . . . . . . . . . . . 4. arboricolus<br />

1. <strong>Anopheles</strong> (<strong>Anopheles</strong>) barberi Coquillett<br />

Figs. l-4<br />

1903. <strong>Anopheles</strong> barberi Coquillett, 1903:310. TYPE: HoZotype 0, Plummer’s Island, Maryland,<br />

United States, 17 Aug 1903, H.S. Barber [USNM, 69591.<br />

<strong>Anopheles</strong> (<strong>Anopheles</strong>) barberi of Edwards (1921:272); Dyar (1928:454); Matheson (1944: 114-<br />

115); Darsie (1949:524-525); Penn (1949:65-66); Carpenter and La Casse (1955:32-34, in<br />

part); Stone, Knight and Starcke (1959:15, in part); Carpenter (1968:72, in part); Zavortink<br />

(1969:31-33).<br />

<strong>Anopheles</strong> barberi of Dyar (1904:243-244); Dyar and Knab (1907:49); Stratman-Thomas and<br />

Baker (1936: 182-183); Vargas (1942a: 172,173,174; 1942b:329-331); Russel, Rozeboom and<br />

Stone (1943:21,31, in part); Jenkins and Carpenter (1946:35-36, in part); Petersen, Chapman<br />

and Willis (1969: 134-135).<br />

<strong>Anopheles</strong> (Coelodiazesis) barberi of Dyar (1918: 142); Vargas (1943:64,65,66,67, in part); Vargas<br />

and Martinez(1956:l ll-113,140, in part);Vargas(1959:373).<br />

Coelodiazesis barberi of Dyar and Knab (1906: 177); Howard, Dyar and Knab (19 17: 1036-l 038).<br />

FEMALE (fig. 2). Wing: 3.49 mm. Proboscis: 1.73 mm. Forefemur: 2.10 mm.<br />

Abdomen: about 2.2 mm. A small light-colored species. Head: Integument light to<br />

dark brown; interocular bristles dark or amber; erect scales long, usually dingy yel-<br />

lowish mesally, dark or dingy yellowish laterally; interocular scales absent or very<br />

few, short and dingy yellowish; palpus entirely dark scaled, the proximal scales ap-<br />

pressed. Thorax : Mesonotum shortened and arched ; mesonotal integument very<br />

light to dark brown, largely shining, without a broad hoary median longitudinal<br />

stripe; pleural integument lighter than that of mesonotum or becoming lighter ven-<br />

trally; mesonotal bristles numerous, strongly developed, very long and conspicuous,<br />

dark in color; mesonotum without scales or sometimes center of anterior promon-<br />

tory with l-4 narrow dark scales; ppE bristles 6-l 1. Legs: Coxae more or less same<br />

color as adjacent portions of pleuron; anterior forecoxal bristles 19-32; femora, tib-<br />

iae and tarsi entirely dark scaled. Wing: Veins and fringe entirely dark scaled, scales<br />

on veins uniformly distributed and not grouped into dark spots; moderately scaled;<br />

scales in central portion of vein 1A clasping vein or slightly spreading. Abdomen:


8 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

Integument of tergites light to dark brown, usually somewhat dappled, sternites<br />

lighter.<br />

MALE (fig. 2). As for female except for sexual differences.<br />

MALE GENITALIA (fig. 3). Sidepiece: More or less elliptical in shape, short;<br />

spicules arranged singly, in rows, or in small clumps; scales usually absent; para-<br />

basal spines much more strongly developed than internal spine; lateral parabasal<br />

spine flattened preapically, apex sharply attenuate and recurved or sinuous; inter-<br />

nal spine not more strongly developed than large setae of sidepiece, apex usually<br />

sinuous. Claspette: Most lateral seta of ventral lobe distinctly shorter and/or finer<br />

than seta next mesad. Clasper: Without spicules near base.<br />

PUPA (fig. 3). Abdomen: 3.04 mm. Trumpet: 0.37 mm. Paddle: 0.88 mm. Width<br />

of segment VIII: 0.73 mm. Cephalothorax: Moderately pigmented with darker areas<br />

more extensive than in judithae; hairs 1,3-C subequal in length. Trumpet: Dark am-<br />

ber to brown in color. Abdomen: Moderately pigmented, darker areas more exten-<br />

sive than in judithae; hairs 6,7-I subequal in length; l-11,111 usually single or double;<br />

5-111-V short, fine, usually single; 5-VI,VII thickened and single, usually shorter than<br />

to subequal in length to hair 9 of corresponding segment; 9-III-VIII darkly pigment-<br />

ed, strongly contrasting with abdominal integument; 9-111 approaching 9-IV in both<br />

diameter and length; O-VII very small, usually single (single, double); 7-VII usually<br />

double (single, double), shorter than 9-VIII. Paddle: Moderately pigmented, midrib<br />

darker.<br />

FOURTH INSTAR LARVA (fig. 4). Head: 0.65 mm. Anal Saddle: 0.27 mm.<br />

Head: Inner clypeals (2-C) widely spaced, usually separated by a distance greater<br />

than 2.0 that between inner and outer clypeals, single and simple; outer clypeals<br />

(3-C) far mesad of 4-C, fine, single; 1 I-C not plumose, single or 2,3b. Antenna:<br />

Spicules absent or inconspicuous; hair 4-A simple and single. Thorax: Integument,<br />

especially on underside of prothorax, spiculose; hair 2-P short, plumose, with num-<br />

erous lateral branches; 8-P long, shaft not notably thickened, lateral branches short<br />

and moderately numerous; 8-M short, all lateral branches long; 8-T a normal plu-<br />

mose hair; 9-P,M,T spiculate. Abdomen: Integument on underside of segments l-<br />

VIII spiculose; hair 1-I very short, single, l-11 simple to palmate, l-III-VII palmate;<br />

6-I-VI similar in development and length; 9-I-VI moderately developed, with branch-<br />

es arising near base, never stellate or plumose; 13-II-V,VII moderately developed,<br />

usually 3b (3-5); hair 4-IV,V simple, single; 5-VII moderately long with 2,3b (l-3)<br />

arising near base; 7-VII moderately long, usually single or double (I-3b); hair lo-<br />

VII not plumose but single or 2,3b or 2,3f. Segment VIII: Hairs 3,5 usually with<br />

3,4b (l-5) arising near base. Anal Segment: Spicules of saddle moderately conspic-<br />

uous, located along ventral and caudal margins; hair 1 simple, usually single.<br />

SYSTEMATICS. Although barberi and judithae are a pair of closely related allo-<br />

patric species, they are easily separated in all stages on the basis of the characters<br />

given in the keys.<br />

Examination of more specimens since the publication of my earlier paper (Zavor-<br />

tink, 1969) has indicated that some changes need to be made in the description of<br />

barberi presented there. In the male genitalia the ratio of the distance from the para-<br />

basal spines to the internal spine to the distance from the latter to the apex of the<br />

sidepiece should be broadened from 0.8-l .O to 0.6-l .6 and the entire proctiger sec-<br />

tion should be deleted. In the pupa hair 5-VI,VII is sometimes longer than previ-<br />

ously indicated and is then subequal in length to hair 9 of the corresponding seg-<br />

ment. In the larva hair l-11 is frequently palmate and 7-VII is often double or triple.<br />

<strong>The</strong> following extremes in variation of chaetotaxy of the pupa have been seen:


Zavortink: Treehole <strong>Anopheles</strong> of New World 9<br />

hair 5-VI is fine, as in judithae, on a single specimen from Ithaca, New York; hair<br />

5-V is thickened, as in xelajuensis, on one side of the only specimen available for<br />

study from the Baltimore, Maryland, area; hair 9-VI,VII is shortened in some specimens<br />

from Ithaca and hair 5 of the corresponding segment is then longer than it.<br />

An. barberi is apparently the most modern species of the group. It is widespread,<br />

occurring throughout the eastern half of the United States, and the vestiture of the<br />

adult is the most derived of any New World <strong>treehole</strong> <strong>Anopheles</strong>.<br />

BIONOMICS. An. barberi is found at elevations of less than 1200 meters. <strong>The</strong><br />

immature stages are found in rot holes in trees and stumps and in artificial containers,<br />

particularly those made of wood or containing leaves and twigs. Since long<br />

periods of unfavorable weather, at least in the northern portion of its range, are<br />

passed in the larval stage, the species is usually recovered from sites which contain<br />

water more or less continuously. More than a dozen larvae are seldom collected together;<br />

this is apparently due to their predacious nature. Adults are occasionally<br />

found resting in buildings and culverts and under bridges. Both sexes are attracted<br />

to lights and females are attracted to humans.<br />

DISTRIBUTION (fig. 1). An. barberi is the most widely distributed of the New<br />

World <strong>treehole</strong> <strong>Anopheles</strong>, being found at low to moderate elevations throughout<br />

the eastern United States, from South Dakota and New York in the north to Texas<br />

and Florida in the south. Material examined: 239 specimens; 60 males, 19 male genitalia,<br />

68 females, 1 adult, 34 pupae, 57 larvae; 24 individual rearings (3 pupal, 16<br />

larval, 5 incomplete).<br />

UNITED STATES. Alabama: Guntersville Lake, 28 May-27 Aug 1942, 3 9 [UCLA] . Ozark,<br />

Camp Rucker, 11 Mar 1943, J .G. Franclemont , 1 d [CU] . Sheffield, 18 Aug 1942, J.N. Belkin, 1 9<br />

[UCLA]. Wilson Dam, 10 June-19 July 1942, J.N. Belkin, 1 lp0 (167), 1 pd(171), 1 p? (168), 2<br />

d, 2 d gen [UCLA] ; 1942, 1 d [HOPK] . Arkansas: Little Rock, Little Fourche Bayou, 28-29 Mar<br />

1943, J.N. Belkin, 2 lp (440,448), 3 P [UCLA]. Scott, 2 Ott 1908, J.K. Thibault, 1 d, 1 d gen, 1<br />

P [USNM] . Delaware: Bombay Hook, 17 June 1964, R.W. Lake, 1 L [USNM] ; 19 Aug 1965, R.W.<br />

Lake, 1 9 [USNM] . Coach’s Bay, 26 June 1961, R.W. Lake (224), 1 d, 1 d gen [USNM] . District<br />

of Columbia: Kenilworth, 2 July 1943, C.W. Travis, 1 P [USNM] . Soldier’s Home, 3 July 1943,<br />

N.E. Good and C.W. Travis, 2 d’, 1 d gen [USNM]. No locality, 12 Aug 1944, N.E. Good, 1 L<br />

[USNM] . Georgia: Atlanta, Fort McPherson, 19 Apr 1943, 2 L [USNM] . Milledgeville, 14 June<br />

1950, R.H. Foote, 1 L [USNM] . Savannah, Chatham Field, 17 Aug 1944, 2 L [USNM] . Iowa:<br />

Ames, 10 Sept 1919, Bishopp (9085), 2 d, 1 6 gen [USNM]. Oskaloosa, 14 Sept 1933,l 9 [US-<br />

NM]. Kentucky: Louisville, G.E. Quinby, 1 d [USNM] . Maxon Mill, 17 June 1935, G.E. Quinby,<br />

1 d, 1 d gen [USNM] . Louisiana: Alexandria, Esler Field, 13 Ott 1942, W.W. Wirth, 1 d [USNM] .<br />

Baton Rouge, 1941-1947, W.W. Wirth, 2 ~3, 1 d gen, 1 0, 1 L [USNM]. Lake Charles, 15 June<br />

1943, W.W. Wirth, 1 d [USNM] . Olla, May 1943,l 0 [USNM] .Malyland: Annapolis, 6 July 1933,<br />

F.C. Bishopp, 1 P [USNM] . Baltimore, July 1931, 1 L [HOPK] . Baltimore, Gwynns Falls Park,<br />

July 1931, 1 Ip [HOPK] . Baltimore, Patapsco State Park, 17 Ott 1965, W.A. McDonald (UCLA<br />

286), 1 L [UCLA]. Bethesda, 22 Aug 1944, G.B. Vogt, 1 d, 1 Q [USNM] . Cabin John, Aug-Oct<br />

1908, F. Knab, 3 d, 5 9, 2 L [USNM] ; July 1965, W.A. McDonald (UCLA 286A), 1 L [UCLA].<br />

Great Falls, 18 May 1919, W.L. McAtee, 1 0 [USNM]. Plummer’s Island, 23 Aug-10 Sept 1903,<br />

H.S. Barber, 2 9 [USNM] ; 5 Sept 1904, H.S. Barber and E.A. Schwarz, 1 9 [USNM] ; lo-13 July<br />

1905, H.S. Barber, 2 d, 2 C$ gen, 4 9 [USNM] ; 30 Aug 1908, F. Knab, 1 d [USNM] ; 24 May<br />

1912, E.A. Schwarz and H.S. Barber, 1 0 [USNM] ; 19 June 1912, H.S. Barber, 1 d [USNM] ; 13<br />

Sept (10263), eggs, 3 L [USNM] ; 1 0 [USNM] . Mississippi: Agricultural College, 23 July-l 5 Ott<br />

1905, G.W. Herrick, 2 9 [USNM]. Clinton, 30 May 1945,l d [USNM] . Greenwood, 1928,2 6, 1<br />

d gen, 3 0 [USNM] . Hattiesburg, Camp Shelby, 22 June 1943, 2 L [USNM] . Missouti: Neosho,<br />

Camp Crowder, 20 July 1942, A.B. Gurney (96), 1 0, 1 P [USNM] ; Sept 1942, A.B. Gurney<br />

(196), 1 9, 1 P, 1 L [USNM] . St. Louis, Aug, A. Busck, 1 9 [USNM] .iVew Jersey: Chester, 8 Sept,<br />

J.M. Aldrich, 1 0 [USNM] . New York: Ithaca, 15 June-15 Ott 1932, R. Matheson, 5 d, 10 9,l lp<br />

(1027-1065), 5 P, 8 L [CU], 2 d, 1 d gen, 3 9 [UCLA]; 1933, 1 L [CU]; 20 Aug 1935,


10 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

1 d, 1 A [CU] . North Carolina: Fayetteville, Fort Bragg, 1 Aug 1945,l 0 [USNM] . Montreat, 25<br />

June 1940, 1 0 [HOPK]. Ohio: Canton, 22 Dee 1967, T.J. Zavortink (UCLA 437), 10 lpd (437-<br />

30-36,39,41,43), 4 1~0 (437-37,38,40,42), 1 pd (437-44), 2 8 gen, 1 L [UCLA]. Oklahoma: Idabel,<br />

7 June 1938, L.E. Rozeboom, 1 lp (A-42) [HOPK] . South Carolina: Charleston, Charleston<br />

Field, 17 Aug 1944, 2 L [USNM] . Columbia, 1 Aug 1906, 1 d, 1 d gen, 1 0 [USNM] . Tennessee:<br />

Kentucky Lake, 26 Aug 1942, 1 d, 1 d gen [UCLA]. Paris, Camp Tyson, 21 June 1944, 2 d<br />

[USNM] . Reelfoot Lake, 11 Sept 1939, T.W. Simpson and G.E. Quinby, 1 L [HOPK] . Virginia:<br />

Bluemont, 29 July 1904, 1 ~3 [USNM] . Dead Run, 2 July, R.C. Shannon, 3 d, 1 ~3 gen [USNM] .<br />

Newington, 22 Aug 1910, S.A. Rohwer, 1 0 [USNM] . Williamsburg, Camp Peary, July 1943,R.<br />

Bohart, 1 L [USNM] . Woodstock, 9 Aug 1904, F.C. Pratt, 2 d, 1 0 [USNM] . Texas: Abilene, 30<br />

Aug 1953, R.X. Schick, 2 9 [UCLA]. No Data: (Mound 10180), 1 lp? (10180A3), 1 d, 2 9<br />

[USNM] ; (Mound 10253-5), 1 d gen [USNM] ; E.B. Johnson (37-268), 1 d [USNM] ; (57-3), 1 L<br />

[USNM] ; (125-l), 1 0 [USNM] ; (140-2), 1 9 [USNM] ; (150-3), 1 d gen, 1 L [USNM] ;(150-4),<br />

1 d [USNM] ; 1 0 [USNM] ; 1 L [CU] .<br />

2. <strong>Anopheles</strong> (<strong>Anopheles</strong>) judithae Zavortink<br />

Figs. 1,5,6<br />

1969. <strong>Anopheles</strong> (<strong>Anopheles</strong>) judithae Zavortink, 1969:28-3 1. TYPE: Holotype d with associated<br />

larval and pupal skins (UCLA 302-40), Cochise Stronghold Recreation Area, Dragoon<br />

Mountains, Cochise County, Arizona, United States, larva from oak <strong>treehole</strong>, 22 Mar<br />

1966, T.J. Zavortink [USNM] .<br />

<strong>Anopheles</strong> (<strong>Anopheles</strong>) barberi of Vargas (1940:319-322); Carpenter and La Casse (1955:32-34,<br />

in part); Stone, Knight and Starcke (1959: 15, in part); Carpenter (1968:72, in part).<br />

<strong>Anopheles</strong> barberi of Russel, Rozeboom and Stone (1943:21,31, in part); Jenkins and Carpenter<br />

(1946:35-36, in part); Richards, Nielsen and Rees (1956: 14); Rigby, Blakeslee and Forehand<br />

(1963:50); Burger (1965:396); Nielsen, Arnell and Linam (1967:76); Nielsen, Linam, Arnell<br />

and Zavortink ( 1968 : 363).<br />

<strong>Anopheles</strong> (Coelodiazesis) barberi of Vargas (1943:64,65,66,67, in part); Vargas and Martinez<br />

(1956:111-113,140,inpart).<br />

FEMALE. Wing: 3.3 1 mm. Proboscis: 1.92 mm. Forefemur: 2.18 mm. Abdomen:<br />

about 2.1 mm. Very similar to barberi but differing in the following. Head: Inter-<br />

ocular bristles amber; erect scales long, entirely pale white or some lateral ones dark;<br />

interocular scales numerous, white, usually none appreciably elongate. Thorax: An-<br />

terior acrostichal bristles amber; mesonotal scaling restricted to a small tuft of 5-20<br />

whitish scales in center of anterior promontory; ppl bristles 2-5. Legs: Anterior fore-<br />

coxal bristles 6- 18.<br />

MALE. Similar to female except for sexual differences.<br />

MALE GENITALIA (fig. 5). Sidepiece: More or less cylindrical or short conical<br />

in shape, short; spicules arranged singly, in rows, or in small clumps; scales absent;<br />

parabasal spines only slightly more strongly developed than internal spine; lateral<br />

parabasal spine not flattened preapically, apex gradually attenuate and strongly re-<br />

curved; internal spine more strongly developed than large setae of sidepiece, apex<br />

usually recurved. Claspette: Most lateral seta of ventral lobe distinctly shorter and/or<br />

finer than seta next mesad. Clasper: Without spicules near base.<br />

PUPA (fig. 5). Abdomen: 2.80 mm. Trumpet: 0.37 mm. Paddle: 0.83 mm. Width<br />

of segment VIII: 0.75 mm. Cephalothorax: Usually lightly pigmented with darker<br />

areas less extensive than in barberi; hairs 1,3-C subequal in length. Trumpet: Light


Zavortink: Treehole <strong>Anopheles</strong> of New World 11<br />

amber in color. Abdomen: Usually lightly pigmented, darker areas less extensive<br />

than in barberi; hairs 6,7-I more or less subequal in length; l-11,111 usually single<br />

(1-3b); hair 5-III-VI short, fine, usually 2,3b; hair 5-VII usually short, fine and 2,3b,<br />

sometimes thickened and single, but then longer than 9-VII; hair 9-III-VIII lightly<br />

pigmented, concolorous with abdominal integument; 9-111 more or less intermediate<br />

in diameter and length between 9-H and 9-IV; hair O-VII small, single; 7-VII usually<br />

2,3b (l-4), shorter than 9-VIII. Paddle: Usually lightly pigmented, midrib darker.<br />

FOURTH INSTAR LARVA (fig. 6). Head: 0.67 mm. Anal Saddle: 0.30 mm.<br />

Head: Inner clypeals (2-C) closely approximated, separated by a distance less than<br />

that between inner and outer clypeals, single and simple; outer clypeals (3-C) far<br />

mesad of 4-C fine, double; 11-C not plumose, single or 2,3b. Antenna: Spicules<br />

inconspicuous; hair 4-A simple, forked apically. Thorax: Integument without spicules;<br />

hair 2-P moderately long, with only a few (2-6) long lateral branches; 8-P moderately<br />

long, shaft not conspicuously thickened, lateral branches long and moderately<br />

numerous; 8-M moderately long, apical lateral branches very long; 8-T a normal<br />

plumose hair; 9-P,M,T simple. Abdomen: Integument without spicules; 1-I very<br />

short, usually single (1-3b), hair 1 -II-VII palmate; 6-III-VI less strongly plumose than<br />

6-1,II; hair 9-I-VI moderately developed, most branches arising near base, never stellate<br />

or plumose; 13-II-V,VII moderately developed, usually single (l-3b); hair 4-IV,V<br />

simple, single; 5-VII long with 3,4b arising from near base; 7-W moderately long<br />

with 2,3b arising from near base; lo-VII not plumose, but with 2,3b arising from<br />

near base. Segment VIZI: Hair 3 with 2-4b arising from base; 5 usually single. AnaZ<br />

Segment: Spicules of saddle inconspicuous, located along caudal margin and distal<br />

half of ventral margin; hair 1 simple, single.<br />

SYSTEMATICS. An. judithae is the <strong>treehole</strong> <strong>Anopheles</strong> of the southwestern United<br />

States. Although it is quite distinct from barberi in all stages, it was confused<br />

with that species for nearly 30 years. <strong>The</strong> two can be easily distinguished on the<br />

basis of the diagnostic features given in the keys.<br />

Although pupal hair 5-VII is fine and 2,3b in most specimens of judithae, it is<br />

thickened, single and elongate in a few individuals from nearly every collection.<br />

This is true not only in the population in the Gila River system, but also in the one<br />

in the Chisos Mountains of Texas.<br />

BIONOMICS. An. judithae occurs at elevations between 600 and 2200 meters.<br />

At the upper limit of its altitudinal range it occurs in the more or less continuous<br />

xeric evergreen forest, but at lower elevations it is restricted to the narrow bands<br />

of trees lining watercourses. <strong>The</strong> immature stages have been taken only from <strong>treehole</strong>s.<br />

Like barberi, judithae overwinters in the larval stage and, as a result, is more<br />

frequently found in permanent than temporary <strong>treehole</strong>s. In contrast to barberi, it<br />

is not unusual to find great numbers of judithae larvae in the same <strong>treehole</strong>. Except<br />

for 3 males collected in a building in Sonora, Mexico (Vargas 1940: 3 19-322), adults<br />

have not been encountered outside the laboratory.<br />

DISTRIBUTION (fig. 1). An. judithae is widely distributed at moderate to high<br />

elevations in the southwestern United States and, undoubtedly, northern Mexico.<br />

In Arizona and western New Mexico it is common in the Gila River system. In<br />

Texas it has been collected in the southern Trans-Pecos area. Material examined:<br />

1002 specimens; 200 males, 15 male genitalia, 190 females, 304 pupae, 293 larvae;<br />

168 individual rearings (56 pupal, 102 larval, 10 incomplete).<br />

MEXICO. Sonora: Imuris, 1940, A. Martinez Palacios, 1 d, 1 cf gen [ISET].<br />

UNITED STATES. Arizona: Chiricahua National Monument, Headquarters, 24 Dee 1966, L.T.<br />

Nielsen (N-36-66), 1 d [UTAH] ; same data (N-37-66), 3 d, 4 0 [UTAH] . Cochise Stronghold Rec-


12 Contrib. Amer. Ent. Inst ., vol. 5, no. 2, 1969<br />

reation Area (Dragoon Mts.), 22 Mar 1966, T.J. Zavortink (UCLA 302), holotype lpd (30240),<br />

allotype 1~9 (30242) [USNM] , 6 lpd (302-25,41,43,45,46,48), 5 lp? (302-44,47,49,5 1,52), 3 pd<br />

(302-100,101,103), 1 p? (302-102), 1 0, 1 P, 6 L [UCLA] ; 23 Mar 1966, T.J. Zavortink (UCLA<br />

306), 1 1~0 (306-20) [UCLA] ; 4 Sept 1966, T.J. Zavortink (UCLA 328), 1 lpd (328-22) [UCLA] ;<br />

6 Sept 1966, T.J. Zavortink (UCLA 342), 7 1~~3 (342-13,15,30,33,38,43,44), 11 1~0 (342-10,31,<br />

32,34-37,3942), 10 pd (342~100~106,109,110,113), 5 p? (342~107,108,111,112,114), 17 d, 2 d<br />

gen, 25 0, 48 P, 27 L [UCLA]. Coronado National Memorial, Headquarters, 20 Mar 1968, L.T.<br />

Nielsen, J H. Arnell and J.H. Linam (HA-10-68), 2 c$, 2 9 [UTAH]. Coronado National Memorial<br />

(1 mi E), 20 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-g-68), 2 d, 1 9 [UTAH].<br />

Douglas (34 mi ENE), 13 Sept 1968, L.T. Nielsen (N-43-68), 2 0, 2 L [UTAH]. Fort Huachuca,<br />

17 Sept 1962, Bates, 1 d gen [USNM] . Kitt Peak National Observatory (Quinlan Mts.), 12 Sept<br />

1968, T.J. Zavortink (UCLA 448), 1 Ipd (448-27), 6 lp? (448-20-25), 1 p? (448-15), 1 lp (448-28)<br />

[UCLA] ; 8 Sept 1969, T.J. Zavortink (UCLA 630), 1 pd (630-103), 1 P, 14 L [UCLA]. Lochiel<br />

(2 mi E), 21 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-13-68), 1 d [UTAH].<br />

Lochiel (11 mi E), 21 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-12A-68), 4 L<br />

[UTAH]. Nogales (13 mi NW), 21 Mar 1966, T.J. Zavortink (UCLA 297), 2 1~0 (297-20,21)<br />

[UCLA]. Nogales (9 mi NE), 21 Mar 1966, T.J. Zavortink (UCLA 298), 3 lpd (29840,42,49), 4<br />

lp? (29843,45,50,51), 3 pd (298-lOl-103), 4 lp (29841,4648), 1 d gen, 3 ?,4 P, 22 L [UCLA] ;<br />

same data (UCLA 299), 5 lpd (299-21,26,28-30), 6 lp? (299-20,23-25,27,31), 1 lp (299-22), 1 ~3<br />

gen, 2 L [UCLA] . Palominas ( 1 mi E), 20 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam<br />

(HA-8-68), 1 cl, 1 9 [UTAH]. Patagonia (14 mi WSW), 24 Aug 1954, W.A. McDonald (UCLA<br />

137), 2 L [UCLA] ; 6 Sept 1963, J. Burger (UCLA 413A), 2 L [UCLA] ; 18 Aug 1964, J. Burger<br />

(UCLA 253), 17 d, 11 9 [UCLA] ; 13 Sept 1964, J. Burger (UCLA 270), 3 L [UCLA] ; 20 Sept<br />

1964, J. Burger (UCLA 260), 17 d, 1 d gen, 9 ? [UCLA] ; 25 July 1965, J. Burger (UCLA 281), 4<br />

L [UCLA] ; 27 July 1965, J. Burger (UCLA 282), 5 L [UCLA] ; 21 Mar 1966, T.J. Zavortink<br />

(UCLA 300), 2 Ipd (300-30,38), 6 lp9 (300-32-36,39), 2 pd (300-100,102), 2 p? (300~101,103), 3<br />

d, 3 9,6 P, 12 L [UCLA] ; 5 Sept 1966, T.J. Zavortink (UCLA 333), 1 cl, 1 P, 2 L [UCLA] ; same<br />

data (UCLA 334), 1 1~~3 (334-15), 4 lp? (334-12-14,16), 2 pd (334-102,103), 3 p0 (334-100,101,<br />

104), 1 0, 3 P, 2 L [UCLA] ; 21 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-14-68),<br />

18 d, 17 0, 11 L [UTAH] ; 14 Sept 1968, T.J. Zavortink (UCLA 458), 1 pQ (458-100). Portal (3<br />

mi W), 20 Mar 1968, L.T. Nielsen, J H. Arnell and J.H. Linam (HA-7A-68), 1 d [UTAH]. Portal<br />

(15 mi WNW), 6 Sept 1966, T.J. Zavortink (UCLA 343), 9 lpd(343-14,15,17,31-34,36,37), 8 lp?<br />

(343-12,16,19,30,38-41) 10 pcl (343~100~104,107,109,110,113,114), 4 p0 (343~105,108,111,<br />

112), 2 lp (343-13,35), 30 d, 2 d gen, 25 9, 69 P, 39 L [UCLA]. Prescott, 19 Mar 1966, L.T.<br />

Nielsen (N-2-66), 2 cl, 1 0, 1 P, 5 L [UTAH]. Prescott (7 mi NE), 1 Sept 1966, T.J. Zavortink<br />

(UCLA 315), 2 lpd (315-16,18), 1 dgen [UCLA]. Santa Rita Mts., 20 Ott 1940, R.A. Flock, 1 d,<br />

1 0 [CU] , 1 c3, 1 d gen [UCLA]. Wickenburg (5 mi S), 18 Mar 1966, L.T. Nielsen (N-l-66), 1 d,<br />

1 dgen, 1 ?,9L [UTAH].NewMexico: Animas(lSmiS), 19Mar 1968,L.T.Nielsen,JH.Arnell<br />

and J.H. Linam (HA-6C-68), 1 0 [UTAH]. Glenwood (5 mi E), Catwalk Cmpg., 21 Mar 1967, L.T.<br />

Nielsen and J.H. Linam (N-5-67), 1 Cs, 1 ~3 gen, 1 P, 1 L [UTAH]. Texas: Big Bend National Park,<br />

Chisos Mts., 31 Aug 1969, T.J. Zavortink and J.A. Bergland (UCLA 603), 1 lp$ (603-23), 4 1~0<br />

(603~21,22,26,33), 6 pd (603-100,101,102,103,108,109), 2 p? (603~104,l lo), 2 lp (603-20,27), 2<br />

d gen, 1 I’, 4 L [UCLA] ; 1 Sept 1969, T.J. Zavortink (UCLA 612), 2 lpd (612-10,l l), 2 lp? (612-<br />

20,24), 1 L [UCLA] ; same data (UCLA 613), 1 lpd (613-14) [UCLA].<br />

3. <strong>Anopheles</strong> (<strong>Anopheles</strong>) fausti Vargas<br />

Figs. 1,7,12<br />

<strong>Anopheles</strong> (Coelodiazesis) fausti Vargas, 1943:66-68. TYPE: Holotype larva, Tamazun-<br />

chale, San Luis Potosi, Mexico, from <strong>treehole</strong>, Apr or May 1942, M. Macias [ISET].<br />

<strong>Anopheles</strong> (<strong>Anopheles</strong>) fausti of Stone, Knight and Starcke (1959: 18, in part); Belkin, Schick and


Zavortink: Treehole <strong>Anopheles</strong> of New World 13<br />

Heinemann (1965:34); Zavortink (1969:30, in part).<br />

<strong>Anopheles</strong> (Coelodiazesis) fausti of Vargas and Martinez (1956: 113-l 16,141); Senevet (1958:43);<br />

Vargas (1959:377).<br />

<strong>Anopheles</strong> xelajuensis of Vargas (1942a:169-175); Russel, Rozeboom and Stone (1943:35, in<br />

part).<br />

FEMALE (fig. 12). Wing: 3.36 mm. Proboscis: 2.06 mm. Forefemur: 1.98 mm.<br />

Abdomen: about 2.0 mm. A small dark species. Head: Integument dark brown; in-<br />

terocular bristles white; erect scales apparently short, dark brown except for a small<br />

patch of bright white scales in anterior dorsal region; interocular scales very long,<br />

fine, wavy, white, forming a conspicuous tuft; palpus with white scales at joints and<br />

apex, proximal scales slightly outstanding. Thorax: Mesonotum elongate and rather<br />

flat; mesonotal integument dark brown except for broad hoary median longitudinal<br />

stripe; pleural integument largely dark brown; mesonotal bristles not as long, strong<br />

or conspicuous as in barberi and judithae, dark except for light anterior acrostichals;<br />

center of anterior promontory with a conspicuous tuft of white scales; acrostichal<br />

scale line extending about halfway to scutellum; ppl bristles 3-5. Legs: Coxae and<br />

trochanters white, strongly contrasting with dark pleuron; anterior forecoxal bris-<br />

tles 13-20; femora, tibiae and tarsi dark scaled except as follows: apex of fore- and<br />

midfemora with a few white scales, more conspicuous on midfemur; apex of hind-<br />

femur with a broad ring of semierect white scales; apex of tibiae usually with a few<br />

white scales. Wing: Dark scaled except for a single large cream-colored apical fringe<br />

spot at ends of branches of vein R; scales of veins uniformly distributed and not<br />

grouped into dark spots; moderately scaled; scales in middle of vein IA nearly clasp-<br />

ing vein. Abdomen: Tergites and sternites dark brown.<br />

MALE. As for female except for usual sexual differences,<br />

MALE GENITALIA (fig. 12). Sidepiece : More or less cylindrical or short conical<br />

in shape, short; spicules grouped into large clumps; 1 to 5 scales usually present;<br />

parabasal spines slightly to considerably stronger than internal spine; lateral para-<br />

basal spine flattened preapically, apex attenuate and straight to recurved; internal<br />

spine as or more strongly developed than large setae of sidepiece, apex nearly<br />

straight, curved or recurved. Claspette: Lateral 2 setae of ventral lobe subequal in<br />

length and stoutness. Clasper: Sometimes with a few spicules near base of ventral<br />

surface.<br />

PUPA. Unknown.<br />

FOURTH INSTAR LARVA (fig. 7). Head: 0.64 mm. Anal Saddle: 0.27 mm.<br />

Head: Inner clypeals (2-C) closely approximated, separated by a distance much less<br />

than that between inner and outer clypeals, single, weakly plumose at least apically;<br />

outer clypeals (3X) distinctly laterad of 4-C, fine, usually forked apically; 11-C<br />

plumose, with 9-l 3 lateral branches. Antenna: Spicules conspicuous; hair 4-A sing-<br />

le, plumose. Thorax: Integument apparently not spiculose; hair 2-P moderately long,<br />

plumose, with numerous lateral branches; 8-P long, shaft not notably thickened,<br />

lateral branches short and moderately numerous; 8-M long, all lateral branches short;<br />

8-T a normal plumose hair; 9-P,M,T barbed. Abdomen: Integument apparently not<br />

spiculose; hair 1-I very short, single or double, l-II-VII palmate; 6-IV-VI shorter<br />

and less strongly plumose than 6-I-111; hair 9-I-VI distinctly plumose; 13-II-V,VII<br />

rather weakly developed with numerous (4-12) fine branches apically; 4-IV,V weak-<br />

ly plumose; 5-VII long, plumose; 7-VII short, plumose or forked apically; lo-VII<br />

plumose. Segment VIII: Hair 3 plumose; hair 5 4,Sf. Anal Segment: Spicules of sad-<br />

dle moderately conspicuous, located along caudal margin and ventral margin distad<br />

of hair 1; hair 1 single, weakly plumose apically.


14 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

SYSTEMATICS. An. fausti is readily separated from all species except arbori-<br />

COZUS in all known stages by the characters given in the keys. It is distinguished<br />

from arboricolus only in the larval stage; the differences are indicated in the key<br />

and the systematics section of arboricolus.<br />

<strong>The</strong> distal portion of many of the larval hairs of fausti is plumose or forked;<br />

because of the fineness of these branches it has not been possible to count them<br />

accurately.<br />

In contrast to the other austral species of New World <strong>treehole</strong> <strong>Anopheles</strong>, fausti<br />

appears to be a low elevation form.<br />

BIONOMICS. <strong>The</strong> larvae of the type series were collected in <strong>treehole</strong>s at elevations<br />

of less than 200 meters in a tropical rain forest. Nothing is known of the<br />

habits of the adults.<br />

DISTRIBUTION (fig. 1). Known definitely only from Tamazunchale and environs,<br />

at the base of the Sierra Madre Oriental in central Mexico. I have not seen<br />

the material upon which the Nicaragua record in Stone, Knight and Starcke (1959:<br />

18) is based. Material examined: 11 specimens; 2 males, 3 male genitalia, 2 females,<br />

4 larvae.<br />

MEXICO. San Luis Potosi: Tlapexhuacan, near Tamazunchale, Apr 1942, M. Macias, 1 0<br />

[HOPK] ,2 c?, 1 d gen, 1 0, 1 L [ISET], 2 d gen, 3 L [USNM] .<br />

4. <strong>Anopheles</strong> (<strong>Anopheles</strong>) arboricolus Zavortink, n.sp.<br />

Figs. 1,8,12<br />

TYPES: Holotype larva (PA 647), “Bajo Grande,” near Cerro Punta, Chiriqui, Panama, eleva-<br />

tion about 2070 meters, from <strong>treehole</strong>, 18 Mar 1964 [USNM] . Paratypes: 5 larvae (PA 647), same<br />

data as holotype [UCLA].<br />

<strong>Anopheles</strong> (<strong>Anopheles</strong>) fausti of Stone, Knight and Starcke (1959: 18, in part); Zavortink (1969:<br />

30, in part).<br />

?AnopheZes (<strong>Anopheles</strong>) xelajuensis of Lane (1953: 171-172, in part); Stone, Knight and Starcke<br />

(1959:30, in part).<br />

?AnopheZes xelajuensis of Galindo (1947 : 23).<br />

FEMALE. Unknown.<br />

MALE. Wing: 3.67 mm. Proboscis: 2.54 mm. Forefemur: 2.23 mm. Abdomen:<br />

about 2.3 mm. A small dark species. Very similar to fausti and possibly not distinguishable<br />

from it since the following minute differences may be individual rather<br />

than specific. Head: Erect scales short, especially mesally, patch of bright white<br />

scales larger than in fausti. Legs: Femora, tibiae and tarsi dark scaled except for a<br />

ring of semierect white scales at apex of hindfemur, this ring narrower and with<br />

the scales less outstanding than in fausti.<br />

MALE GENITALIA (fig. 12). Probably indistinguishable from fausti. Sidepiece:<br />

More or less short conical in shape, short; parabasal spines considerably stronger<br />

than internal spine; internal spine more strongly developed than large bristles of<br />

sidepiece, apex recurved. Clasper: Without spicules near base.<br />

PUPA. Unknown.<br />

FOURTH INSTAR LARVA (fig. 8). Head: 0.83 mm. Anal Saddle: 0.35 mm.<br />

Head: Inner clypeals (2-C) closely approximated, separated by a distance much less<br />

than that between inner and outer clypeals, single and simple; outer clypeals (3-C)<br />

distinctly laterad of 4-C, fine, single or forked apically; 11-C plumose, with 6-8 lat-


Zavortink: Treehole <strong>Anopheles</strong> of New World 15<br />

era1 branches. Antenna: Spicules apparently absent; hair 4-A simple, single or double.<br />

Thorax: Integument apparently without spicules; hair 2-P moderately long, plumose,<br />

with numerous lateral branches; 8-P moderately long, shaft very thick, lateral<br />

branches short and numerous; 8-M moderately long, all lateral branches moderately<br />

long; 8-T a normal plumose hair; 9-P,M simple; 9-T very weakly plumose, with 4-6<br />

lateral branches. Abdomen: Integument apparently without spicules; hair 1-I very<br />

short, usually single (l-3b), hair l-11 multiple or palmate, l-III-VII palmate; 6-I-VI<br />

similar in development and length; 9-I,11 moderately developed, with multiple branches<br />

arising near base, 9-III-VI distinctly plumose; 13-II-V,VII moderately developed,<br />

usually 3b (2-6); hair 4-IV,V usually weakly plumose; 5-VII moderately long with<br />

3,4b usually arising near base; 7-VII short with 4-7b usually arising near base; lo-<br />

VII plumose. Segment VIII: Hair 3 single, simple; hair 5 usually with 2,3b from<br />

base. Anal Segment: Spicules on saddle inconspicuous, located along ventral margin<br />

distad of hair 1 and along caudal edge; hair 1 single, simple.<br />

SYSTEMATICS. An. arboricolus is at present definitely known from a single collection<br />

of 3 fourth and 3 third instar larvae from the Chiriqui Volcano region of<br />

western Panama. Although these larvae agree with those of fausti in many significant<br />

details, they differ in so many others that I do not hesitate to recognize them<br />

as a distinct species in the absence of associated adults. In addition to the characters<br />

given in the key, larvae of the 2 species differ in morphology of hairs 4-A, 8-P,M,<br />

9-P,M,T, 6-IV-VI, S-VII and 3-VIII. <strong>The</strong> description of the male and male genitalia<br />

of arboricolus is based on a single individual; the presumptive association of this<br />

specimen with the larvae is probably correct since the specimen is from the same<br />

region as the larvae and, like them, similar to fausti. In fact, as indicated in the description<br />

above, adults and male genitalia of fausti and arboricolus are probably indistinguishable.<br />

<strong>The</strong> original record of xelajuensis from the Chiriqui Volcano region of Panama<br />

(Galindo, 1947:23) and the subsequent ones based on it (Lane, 1953:172; Stone,<br />

Knight and Starcke, 1959:30) possibly refer to arboricoZus. <strong>The</strong> record of fausti<br />

from Panama (Stone, Knight and Starcke, 1959: 18) is apparently based on the same<br />

male that I have seen and tentatively associated with arboricolus.<br />

BIONOMICS. <strong>The</strong> larvae of the type series were collected in a <strong>treehole</strong> at an elevation<br />

of about 2070 meters. <strong>The</strong> report of finding 6 males and 4 females of xelajuensis<br />

resting in hollow trees at an elevation of about 1900 meters on Volcan Chiriqui<br />

(Galindo, 1947:23) may refer to this species.<br />

DISTRIBUTION (fig. 1). Currently known only from high elevations in the Chiriqui<br />

Volcano area of western Panama. Material examined: 8 specimens; 1 male, 1<br />

male genitalia, 6 larvae.<br />

PANAMA. Chiriqui: “Bajo Grande,” near Cerro Punta, type series, see above. El Volcan Chiriqui,<br />

30 June 1943, T.H.G. Aitken, 1 c$, 1 c$ gen [USNM] .<br />

5. <strong>Anopheles</strong> ( <strong>Anopheles</strong>) xelajuensis de Leon<br />

Figs. 1,9,10,12<br />

1938. <strong>Anopheles</strong> xelajuensis de Leon, 1938:421-423. TYPE: Holotype c3, Cerro Quemado, near<br />

Quezaltenango, Quezaltenango, Guatemala, 2 Jan 1936, J. Romeo de Leon [ESPG] .<br />

<strong>Anopheles</strong><br />

(<strong>Anopheles</strong>) xelajuensis of Lane (1953:171-172, in part); Stone, Knight and Starcke<br />

(1959:30, in part); Belkin, Schick and Heinemann (1965:28); Zavortink (1969:30).


16 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

<strong>Anopheles</strong> xeiizjuensis of Komp (1941:92,96); Russel, Rozeboom and Stone (1943:35, in part);<br />

Forattini(l961:174,180,181).<br />

<strong>Anopheles</strong> (Coelodiazesis) xelajuensis of Vargas and Martinez (1956: 116-l 19,142); Vargas (1959:<br />

377).<br />

<strong>Anopheles</strong> (Russellia) xelajuensis of Vargas ( 1943 :65,67,68-70); Senevet (1958:44).<br />

FEMALE (fig. 12). Wing: 5.16 mm. Proboscis: 2.92 ,mm. Forefemur: 2.75 mm.<br />

Abdomen: about 2.6 mm. A large dark species. Head: Integument dark brown; in-<br />

terocular bristles whitish; erect scales long, mostly black, anterior ones brilliant<br />

white; interocular scales white, much elongate, forming a conspicuous tuft; palpus<br />

entirely dark scaled, basal scales semierect. Thorax: Mesonotum long and only slight-<br />

ly arched; mesonotal integument dark brown with a broad hoary median longitudi-<br />

nal stripe; pleural integument dark brown; anterior mesonotal bristles not as long,<br />

strong or conspicuous as in barberi and judithae, dark except for light anterior acros-<br />

tichals; narrow white scales in tuft in center of anterior promontory and extending<br />

through anterior third of acrostichal line; ppl bristles 3-5. Legs: Coxae and trochan-<br />

ters white; anterior forecoxal bristles 8-16; femora, tibiae and tarsi dark scaled ex-<br />

cept as follows: femora with apical white scales, few on foreleg, many, forming a<br />

conspicuous ring, on hindleg; fore- and midtibiae with a few white scales at apex;<br />

hindtibia with long broad apical white patch covering anterior, dorsal and posterior<br />

surfaces. Wing: Largely dark scaled; apex of costa and vein R, with yellowish-white<br />

scales; 4 fringe spots usually present, 1 large, extending from end of costa to vein<br />

R 4 +5, 3 smaller, at ends of veins M1 +z, Ma +4 and Cu, ; dark scales not uniformly<br />

distributed, but grouped into strong spots at base of veins Rs and R4+5 and at furca-<br />

tion of veins Rz +3, M and Cu; profusely scaled; scales in middle portion of vein 1A<br />

somewhat spreading. Abdomen: Tergites black, sternites black with basal white<br />

band.<br />

MALE. As for female except for sexual characters.<br />

MALE GENITALIA (fig. 9). Sidepiece: More or less cylindrical in shape, long;<br />

spicules arranged singly, in rows or in small clumps; scales numerous, 6-l 6; para-<br />

basal spines stronger than internal spine; lateral parabasal spine not flattened pre-<br />

apically, apex gradually attenuate and recurved; internal spine slightly stronger than<br />

large bristles of sidepiece, apex curved. Claspette: Lateral 2 setae of ventral lobe<br />

subequal in stoutness, but outermost usually shorter. Clasper: With at least a few<br />

spicules near base of ventral surface.<br />

PUPA (fig. 9). Abdomen: not measured. Trumpet: 0.55 mm. Paddle: 1.19 mm.<br />

Width of segment VIII: 0.92 mm. Cephalothorax: Moderately pigmented with dark-<br />

er areas more extensive than in powderi; hair 3-C much longer than 1-C. Trumpet:<br />

Brown. Abdomen: Moderately pigmented, darker areas more extensive than in pow-<br />

deri; hair 6-I much longer than 7-I; hair l-11,111 multiple (3-5b); hair 5-111,IV short,<br />

fine, usually 3,4b ; hair 5-V-VII thickened, much longer than hair 9 of correspond-<br />

ing segment, and with 3-5 fine lateral or apical branches; 9-III-VIII moderately pig-<br />

mented, concolorous with to slightly darker than abdominal integument; 9-111 ap-<br />

proaching 9-IV in both diameter and length; O-VII enlarged, 5,6b; hair 7-VII single,<br />

longer than 9-VIII. PaddIe: Moderately pigmented, midrib not darker.<br />

FOURTH INSTAR LARVA (fig. 10). Head: 0.90 mm. Anal Saddle: 0.39 mm.<br />

Head: Inner clypeals (2-C) closely approximated, separated by a distance much less<br />

than that between inner and outer clypeals, single and simple; outer clypeals (3-C)<br />

distinctly laterad of 4-C, single, thickened; 11-C not plumose, with 2-5b from near<br />

base. Antenna: Spicules inconspicuous; hair 4-A single and simple. Thorax: Integu-


Zavortink: Treehole <strong>Anopheles</strong> of New World 17<br />

ment, at least on ventral surface, spiculose; hair 2-P stellate, 9-15b; hair 8-P moderately<br />

long, shaft very stout, lateral branches very short and moderately numerous;<br />

8-M moderately long, basal lateral branches very long; 8-T with thickened shaft, not<br />

plumose to apex, distal end blunt; 9-P,M,T barbed. Abdomen: Integument, at least<br />

on more posterior segments, spiculose; l-I-VII stellate; &IV-VI less strongly plumose<br />

than 6-I-111; hair 9-I-VI stellate, very strongly developed; 13-II-V,VII strongly developed,<br />

stellate; 4-IV,V simple, single or double; 5,7-VTI stellate, 8-l lb; hair IO-VII<br />

stellate, 4-6b. Segment VIII: Hair 3 with 3-6 very stout branches from base; hair 5<br />

stellate, 18-22b. Anal Segment: Spicules on saddle very conspicuous, located on all<br />

but basal portion of saddle; hair 1 with 3-5b, usually from base.<br />

SYSTEMATICS. An. xelajuensis is the most differentiated New World <strong>treehole</strong><br />

<strong>Anopheles</strong>. <strong>The</strong> most striking characteristics of the species have been indicated in<br />

the keys and the systematics chapter.<br />

According to the original description, which is based on a single male from Guatemala,<br />

the hindtibia of xezajuensis is entirely dark scaled. <strong>The</strong> adults from Mexico<br />

examined during this study have a conspicuous large white patch at the apex of the<br />

hindtibia. It is possible that these specimens represent a different species, but this<br />

cannot be determined until additional material, especially topotypic xelajuensis, is<br />

obtained.<br />

BIONOMICS. Larvae of xelajuensis have been collected in a <strong>treehole</strong> at an elevation<br />

of 2500 meters. According to Vargas (1943 :70) the larvae feed on bottom sediments<br />

and spend relatively little time at the surface, the pupal stage lasts 12 to 15<br />

days and females readily bite humans. <strong>The</strong> holotype male was collected between<br />

rocks in an oak forest in the highlands of Guatemala.<br />

DISTRIBUTION (fig. 1). Known only from high elevations in the Sierra Madre<br />

de1 Sur of southern Mexico and the Sierra Madre in Guatemala. <strong>The</strong> Panama records<br />

of Galindo (1947:23), Lane (1953: 172) and Stone, Knight and Starcke (1959:<br />

30) refer to arboricozus or powderi. Material examined: 13 specimens; 2 males, 3<br />

male genitalia, 1 female, 1 pupa, 6 larvae.<br />

MEXICO. Oaxaca: Galera Vieja, between Ixtlan and Tepanzacoalco, Sept 1942, M. Macias, 1 d<br />

gen, 1 L [CU] ,2 d, 2 d gen, 1 0,2 L [ISET] , 1 P, 2 L [USNM] , 1 L [UCLA].<br />

6. <strong>Anopheles</strong> (<strong>Anopheles</strong>) powderi Zavortink, n.sp.<br />

Figs. 1 ,l 1<br />

TYPE: Holotype 0 with associated pupal skin (CR 50-102), about 10 km SE of La Sierra, San<br />

Jose, Costa Rica, elevation about 2400 meters, from rothole in fallen tree, 24 Nov 1962, C.L.<br />

Hogue and W.A. Powder [USNM] . This species is dedicated to William A. Powder, in recognition<br />

of his contributions to the “<strong>Mosquito</strong>es of Middle America” project.<br />

?AnopheZes (<strong>Anopheles</strong>) xelajuensis of Lane (1953:171-172, in part); Stone, Knight and Starcke<br />

(1959:30, in part).<br />

?<strong>Anopheles</strong> xelujuensis of Galindo ( 1947 : 23).<br />

FEMALE (fig. 11). Wing: 5 .15 mm. Proboscis: 3.13 mm. Forefemur: 2.88 mm.<br />

Abdomen: about 2.7 mm. A large dark species. Quite similar to xelajuensis, differ-<br />

ing mostly in ornamentation of leg and wing. Head: Interocular bristles amber. Tho-<br />

rax: Whitish acrostichal scales extending about halfway to scutellum. Legs: Femora,<br />

tibiae and tarsi dark scaled except for conspicuous ring of semierect white scales at


18 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

apex of hindfemur and a few white scales at apex of hindtibia. Wing: Veins and<br />

fringe dark scaled except for a single large apical fringe spot extending from end of<br />

vein R, to end of vein M, +2 ; scales of veins uniformly distributed and not grouped<br />

into dark spots; profusely scaled; scales in middle portion of vein 1A wide spread-<br />

ing. Abdomen: Tergites and sternites black.<br />

MALE and MALE GENITALIA. Unknown.<br />

PUPA (fig. 11). Abdomen: 4.75 mm. Trumpet: 0.54 mm. Paddle: 1.27 mm.<br />

Width of segment VIII: 1 .l 1 mm. Cephalothorax: Lightly pigmented with darker<br />

areas less extensive than in xelajuensis; hairs 1,3-C subequal in length. Trumpet:<br />

Light amber in color. Abdomen: Lightly pigmented, darker areas less extensive than<br />

in xelajuensis; hairs 6,7-I subequal in length; l-11,111 multiple; S-III-VII short, fine,<br />

usually 2,3b; hair 9-III-VIII lightly pigmented, more or less concolorous with ab-<br />

dominal integument; 9-111 approaching 9-IV in diameter and length; O-VII small,<br />

single; 7-VII double, shorter than 9-VIII. Paddle: Very lightly pigmented, midrib<br />

not noticeably darker.<br />

LARVA. Unknown.<br />

SYSTEMATICS. An. powderi is known from a single female with its associated<br />

pupal skin. <strong>The</strong> adult, as indicated in the key, is adequately distinct but the pupa,<br />

except for its much larger size, is scarcely differentiated from that of judithae. An.<br />

xelajuensis is the only other New World <strong>treehole</strong> <strong>Anopheles</strong> of comparable size.<br />

This species could extend through the Cordillera de Talamanca into western Pan-<br />

ama, and, if so, it might be the xelajuensis of Galindo (1947:23), Lane (1953:172)<br />

and Stone, Knight and Starcke (1959:30).<br />

BIONOMICS. <strong>The</strong> pupa of powderi was taken from a rothole in a fallen tree at an<br />

elevation of about 2400 meters in a cloud forest.<br />

DISTRIBUTION (fig. 1). Presently known solely from the holotype which was<br />

collected high in the Cordillera de Talamanca in southern Costa Rica. Material ex-<br />

amined: 2 specimens; 1 female, 1 pupa; 1 pupal rearing.<br />

COSTA RICA. San Jose: La Sierra (10 km SE), holotype, see above.<br />

REFERENCES CITED<br />

Belkin, John N., R.X. Schick, P. Galindo and T.H.G. Aitken<br />

1965. <strong>Mosquito</strong> Studies (yiptera, <strong>Culicidae</strong>). I. A project for a systematic study<br />

of the mosquitoes of Middle America. Amer. Entomol. Inst., Contrib. l(2):<br />

1-17.<br />

Belkin, John N., R.X. Schick and S.J. Heinemann<br />

1965. <strong>Mosquito</strong> Studies (<strong>Diptera</strong>, <strong>Culicidae</strong>). V. <strong>Mosquito</strong>es originally described<br />

from Middle America. Amer. Entomol. Inst., Contrib. l(5). 95 p.<br />

Burger, John F.<br />

1965. Aedes kompi Vargas and Downs 1950, new to the United States. <strong>Mosquito</strong><br />

Iqews 25:396-398.<br />

Carpenter, Stanley J.<br />

1968. Review of recent literature on mosquitoes of North America. Calif. Vector<br />

Views 15:7 l-98.<br />

Carpenter, Stanley J. and W.J. LaCasse<br />

1955. <strong>Mosquito</strong>es of North America (North of Mexico). Berkeley, Univ. Calif.<br />

Press. 360 p.<br />

Coquillett, Daniel W.<br />

1903. A new <strong>Anopheles</strong> with unspotted wings. Can. Entomol. 35 :3 10.


Zavortink: Treehole <strong>Anopheles</strong> of New World 19<br />

Darsie, Richard F.<br />

1949. Pupae of the anopheline mosquitoes of the northeastern United States<br />

(<strong>Diptera</strong>, <strong>Culicidae</strong>). Rev. Entomol. 20:509-530.<br />

Dyar, Harrison G.<br />

1904. Brief notes on mosquito larvae. N.Y. Entomol. Sot., J. 12:243-246.<br />

19 18. Notes on American <strong>Anopheles</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>). Insecutor Inscitiae<br />

Mens. 6:141-151.<br />

1928. <strong>The</strong> mosquitoes of the Americas. Wash.,Carnegie Inst. (Publication 387).<br />

616 p.<br />

Dyar, Harrison G. and F. Knab<br />

1906. <strong>The</strong> larvae of <strong>Culicidae</strong> classified as independent organisms. N.Y. Entomol.<br />

Sot., J. 14: 169-230.<br />

1907. On the classification of the mosquitoes. Can. Entomol. 39:47-50.<br />

Edwards, Frederick W.<br />

192 1. A revision of the mosquitoes of the Palearctic region. Bull. Entomol. Res.<br />

12:263-35 1.<br />

Forattini, Oswald0 P.<br />

196 1. Chaves para a identificacao do genero <strong>Anopheles</strong> Meigen, 18 18, da Regiao<br />

Neotropical (<strong>Diptera</strong>, <strong>Culicidae</strong>). Rev. Brasil. Entomol. 10: 169-l 87.<br />

Galindo, Pedro<br />

1947. <strong>Anopheles</strong> xelajuensis de Leon, a new addition to the known anopheline<br />

fauna of Panama. Pan-Pacific Entomol. 23:44.<br />

Howard, Leland O., H.G. Dyar and F. Knab<br />

19 17. <strong>The</strong> mosquitoes of North and Central America and the West Indies. v.4.<br />

Wash., Carnegie Inst. (Publication 159). p. 525-1064.<br />

Jenkins, Dale W. and S.J. Carpenter<br />

1946. Ecology of the tree hole breeding mosquitoes of Nearctic North America.<br />

Ecol. Monogr. 16:31-47.<br />

Komp, William H.W.<br />

1941. <strong>The</strong> classification and identification of the <strong>Anopheles</strong> mosquitoes of Mexi-<br />

co, Central America and the West Indies. In Moulton, Forest, R., A symposium<br />

on human malaria. Wash., Amer. Ass. Advance. Sci. (Publication 15). 398 p.<br />

Lane, John<br />

1953. Neotropical <strong>Culicidae</strong>. Sao Paula, Univ. Sao Paulo. 2 v. 1112 p.<br />

de Leon, J. Romeo<br />

1938. El anophelismo de altura en Guatemala. Bol. Dir. Gen. Sanid. Publica (Gua-<br />

temala) 9:41 l-424.<br />

Matheson, Robert<br />

1944. Handbook of the mosquitoes of North America. ed. 2. Ithaca, Comstock<br />

Publishing Co. 3 14 p.<br />

Nielsen, Lewis T., J H. Arnell and J.H. Linam<br />

1967. A report on the distribution and biology of the tree hole mosquitoes in the<br />

western United States. Calif. <strong>Mosquito</strong> Control Ass., Proc. Pap. 35 :72-76.<br />

Nielsen, Lewis T., J.H. Linam, J H. Arnell and T.J. Zavortink<br />

1968. Distributional and biological notes on the tree hole mosquitoes of the west-<br />

ern United States. <strong>Mosquito</strong> News 28:361-365.<br />

Penn, George H.<br />

1949. Pupae of the Nearctic anopheline mosquitoes north of Mexico. Nat. Malar-<br />

ia Sot., J. 8:50-69.


20 Contrib. Amer. Ent. Inst., vol. 5, no. 2, 1969<br />

Petersen, J. J., H.C. Chapman and O.R. Willis<br />

1969. Predation of <strong>Anopheles</strong> barberi Coquillett on first instar mosquito larvae.<br />

<strong>Mosquito</strong> News 29 : 134-l 35.<br />

Richards, Charles S., L.T. Nielsen and D.M. Rees<br />

1956. <strong>Mosquito</strong> records from the Great Basin and the drainage of the lower Colo-<br />

rado River. <strong>Mosquito</strong> News 16: 10-17.<br />

Ribgy, Paul T., T.E. Blakeslee and C.E. Forehand<br />

1963. <strong>The</strong> occurrence of Aedes taeniorhynchus (Wiedemann), <strong>Anopheles</strong> barberi<br />

(Coquillett), and Culex thriambus (Dyar) in Arizona. <strong>Mosquito</strong> News 23:50.<br />

Russel, Paul F., L.E. Rozeboom and A. Stone<br />

1943. Keys to the anopheline mosquitoes of the World. Philadelphia, Amer. Ent-<br />

omol. Sot. 152 p.<br />

Senevet, Georges<br />

1958. Les <strong>Anopheles</strong> due globe. Revision generale. Encycl. Entomol. (A) 36.<br />

215 p.<br />

Stone, Alan, K.L. Knight and H. Starcke<br />

1959. A synoptic catalog of the mosquitoes of the World (<strong>Diptera</strong>, <strong>Culicidae</strong>).<br />

Wash., Entomol. Sot. Amer. (Thomas Say Found. Publication 6). 358 p.<br />

Stratman-Thomas, W.K. and F.C. Baker<br />

1936. <strong>Anopheles</strong> barberi Coquillett, as a vector of Plasmodium vivax Grassi and<br />

Feletti. Amer. J. Hyg. 24: 182-183.<br />

Vargas, Luis<br />

1940. <strong>Anopheles</strong> (<strong>Anopheles</strong>) barberi en Mexico. Inst. Salubr. Enferm. Trop.,<br />

Rev. 1:319-322.<br />

1942a. <strong>Anopheles</strong> xelajuensis Romeo de Leon, 1938 en Mexico. Inst. Salubr. En-<br />

ferm. Trop., Rev. 3: 169-175.<br />

1942b. El huevo de AnopheZes barberi Coquillett, 1903. Inst. Salubr. Enferm.<br />

Trop., Rev. 3:329-33 1.<br />

1943. Los subgeneros Americanos de AnopheEes (<strong>Diptera</strong>, <strong>Culicidae</strong>). <strong>Anopheles</strong><br />

(Russellia) xelajuensis de Leon, 1938 n.subgn. y <strong>Anopheles</strong> (Coelodiazesis)<br />

fausti n.sp. Inst. Salubr. Enferm. Trop., Rev. 4:57-77.<br />

1959. Lista de <strong>Anopheles</strong> de las Americas y su identification por caracteres mas-<br />

culinos. Inst. Salubr. Enferm. Trop., Rev. 19:367-386.<br />

Vargas, Luis and Amado Martinez Palacios<br />

1956. Anofelinos Mexicanos Taxonomia y distribution. Mexico, D.F., Secretar.<br />

Salubr. Asistencia. 181 p.<br />

Zavortink, Thomas J.<br />

1969. <strong>Mosquito</strong> Studies (<strong>Diptera</strong>, <strong>Culicidae</strong>). XV. A new species of <strong>treehole</strong> breed-<br />

<strong>Anopheles</strong> from the southwestern United States. Amer. Entomol. Inst., Con-<br />

trib. 4(4):27-38.


Zavortink: Treehole <strong>Anopheles</strong> of New World 21<br />

FIGURES<br />

1. Distribution of collections examined<br />

2. <strong>Anopheles</strong> (An.) barberi; adult<br />

3. <strong>Anopheles</strong> (An.) barberi; male genitalia and pupa<br />

4. <strong>Anopheles</strong> (An.) barberi; larva<br />

5. <strong>Anopheles</strong> (An.) judithae; male genitalia and pupa<br />

6. <strong>Anopheles</strong> (An.) judithae; larva<br />

7. <strong>Anopheles</strong> (An.) fausti; larva<br />

8. <strong>Anopheles</strong> (An. ) arboricolus ; larva<br />

9. <strong>Anopheles</strong> (An.) xelajuensis; male genitalia and pupa<br />

10. <strong>Anopheles</strong> (An. ) xelajuensis; larva<br />

11. <strong>Anopheles</strong> (An.) powderi; pupa and pleuron and mesonotum of adult<br />

12. <strong>Anopheles</strong> (An.) arboricolus; male genitalia. <strong>Anopheles</strong> (An.) fausti; male gentalia,<br />

wing and hindleg. <strong>Anopheles</strong> (An.) xelajuensis; wing and hindleg.


Muryland<br />

United States<br />

VO.lW<br />

FORE MID


ANOPHELES


)<br />

Fig. 5<br />

0.2<br />

I


1.0<br />

A MP


\ ANOPHELES ,,<br />

1 Fig. 9<br />

I 0.2<br />

I<br />

xelajuensis<br />

OUXUCU<br />

Mexico


ANOPHELES<br />

powderi<br />

CR 50<br />

San Jose<br />

Costa Rica<br />

ii


Gin I3 ANOPHELES<br />

Mexico<br />

fausti<br />

srm L&s Potosi<br />

Maico<br />

1<br />

fausti<br />

smz Luis Potosi<br />

M&&O<br />

I _ / / 0.2<br />

~-l.o-


TABLE 1.<br />

Groups of New World <strong>treehole</strong> <strong>Anopheles</strong><br />

CHARACTER barberi and judithae arboricolus and fmsti xelajuensis powderi<br />

SIZE small small large large<br />

ADULT<br />

interocular scales<br />

erect head scales<br />

light palpal patches<br />

mesonotum<br />

mesonotal hoary<br />

stripe<br />

acrostichal scaling<br />

color of coxae<br />

apex of hindfemur<br />

apex of hindtibia<br />

dark spots on<br />

wing veins<br />

light patches on<br />

wing veins<br />

light fringe spots<br />

MALE GENITALIA<br />

absent to numerous<br />

and short<br />

light mesally, light or<br />

darker laterally<br />

absent<br />

shortened and arched<br />

absent<br />

absent or tuft on<br />

promontory<br />

same as adjacent portions<br />

of pleuron<br />

dark<br />

dark<br />

absent<br />

absent<br />

absent<br />

numerous and long<br />

bright white mesally,<br />

very dark laterally<br />

present<br />

longer and flatter<br />

present<br />

long line<br />

much lighter than<br />

pleuron<br />

conspicuous white<br />

ring<br />

dark<br />

absent<br />

absent<br />

1, from R 1 to R,+s<br />

numerous and long<br />

bright white mesally,<br />

very dark laterally<br />

absent<br />

longer and flatter<br />

present<br />

long line<br />

much lighter than<br />

pleuron<br />

conspicuous white<br />

ring<br />

conspicuous white<br />

patch<br />

present at base of<br />

branches<br />

present at apex of<br />

costa and Rr<br />

4,1 from Rr to R,+s,<br />

1 each at end of<br />

Mr+z, Ma+4 and Cur<br />

numerous and long<br />

bright white mesally,<br />

very dark laterally<br />

absent<br />

longer and flatter<br />

present<br />

long line<br />

much lighter than<br />

pleuron<br />

conspicuous white<br />

ring<br />

dark<br />

absent<br />

absent<br />

1,fromRr toMr+a<br />

sidepiece short short long unknown<br />

arrangement of side- singly, rows or large clumps singly, rows or unknown<br />

piece spicules small clumps small clumps<br />

sidepiece scales absent few many unknown<br />

ventral claspette lobe lateral seta not as strongly lateral 2 setae subequally lateral seta not as strongly unknown<br />

developed as 1 next developed developed as 1 next<br />

mesad mesad<br />

LARVA<br />

hair 3-C<br />

1lC<br />

l-III-VII<br />

9-III-VI<br />

4-IV,V<br />

IO-VII<br />

fine, mesad of 4C<br />

1-3b from base<br />

pahnate<br />

branched from base, moderately<br />

developed<br />

simple, single<br />

2,3b from base or middle<br />

DISTRIBUTION widespread<br />

United States, northern<br />

Mexico<br />

fine, laterad of 4C<br />

plumose<br />

palmate<br />

plumose<br />

plumose<br />

plumose<br />

restricted<br />

central Mexico, west-<br />

em Panama<br />

thick, laterad of 4C<br />

2-5b from base<br />

stellate<br />

stellate, very strongly<br />

developed<br />

simple, single or double<br />

4_6b, stellate<br />

restricted<br />

southern Mexico,<br />

Guatemala<br />

unknown<br />

unknown<br />

unknown<br />

unknown<br />

unknown<br />

unknown<br />

restricted<br />

Costa Rica<br />

HABITAT temperate deciduous forest, humid montane forest, wet montane forest montane cloud<br />

xeric evergreen forest,<br />

gallery forest<br />

tropical forest forest<br />

i


Zavortink: Treehole <strong>Anopheles</strong> of New World 35<br />

INDEX TO SCIENTIFIC NAMES<br />

arboricolus Zavortink, 2,3, 5-7k, 14, 14-15, 17; I, 8, 12<br />

barberi Coquillett, 1,2, 3, 5-7k, 7-10, 10, 11, 13, 16; 1-4<br />

barberi of authors, 10<br />

barianensis James, 2<br />

Coelodiazesis Dyar & Knab, 2<br />

Cyclophorus Eysell, 2<br />

eiseni Coquillett, 3, S-6k<br />

fausti Vargas, 1,2,3, 5-7k, 12-14, 14, 15; 1, 7, 12<br />

fausti of authors, 14, 15<br />

judithae Zavortink, 2,5-7k, 8,9, 10-12, 13, 16, 18; I, 5, 6<br />

plumbeus Stephens, 2<br />

powderi Zavortink, 2,3,5-6k, 16, 17, 17-18; I, II<br />

Russellia Vargas, 2<br />

xelajuensis de Leon, 1,2,3,4,5-6k, 9, 15-17, 17, 18; I, 9, 10, 12<br />

xelajuensis of authors, 13, 14, 15, 17, 18


(Continued from inside front cover)<br />

Parts of Volume 4, with prices<br />

No. 1. <strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1968.<br />

X. Peters, T. Michael. Dixinae originally described from North America. 7 pages.<br />

XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. <strong>Mosquito</strong>es<br />

originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay.<br />

29 pages. Price: $1.25, postpaid.<br />

No. 2. <strong>Mosquito</strong> <strong>studies</strong> (<strong>Diptera</strong>, <strong>Culicidae</strong>), 1969.<br />

XII. Berlin, 0. G. W. A revision of the Neotropical subgenus ~ Howardina of - Aedes.<br />

190 pages, 58 figures. Price: $6.00, postpaid.<br />

No. 3. Carlson, Robert W. and Fred B. Knight. Biology, taxonomy, and evolution of four<br />

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