MORMAX MARSTON Revision of New World

* Species of Anthrax

(Diptera: Bombyliidae), k

Other than the Anthrax

albofasciatus Group

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 43

Norman Marston Revision of New World

Species of Anthrax

(Diptera: Bombyliidae),

Other Than the Anthrax

albofasciatus Group

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

I97O
ABSTRACT

Marston, Norman. Revision of New World Species of Anthrax (Diptera: Bombyliidae) other than the Anthrax albofasciatus Group. Smithsonian Contributions to

Zoology, 43:1-148. 1970. A key is given to the species groups of Anthrax in North

and South America. Four of the five groups are described and keys are given for

included taxa. Each taxon is fully described, its synonymy is compiled, and its distribution is illustrated or summarized. Important taxonomic characters are illustrated.

Sixteen taxa comprise the oedipus group, including peruvianus, cordillerensis, and

insulanus, new species; and irroratus striatipennis, oedipus aquilus, and pluto nigriventris, new subspecies. Twenty-nine taxa comprise the cephus group, including

innubilipennis, inaquosum, nitidus, xanthomeros, hylaios, cathetodaithmos, snowi,

costaricensis, koebelei, austrinus, clinopictus, and laticellus, new species; and argyropygus painteri, new subspecies. Nine species comprise the trimaculatus group, including

baliopteros, latibasis, and caatingensis, new species. Five taxa comprise the tigrinus

group, including xylocopae, new species, and simson habrosus, new subspecies.

Evolution of the cephus and tigrinus groups is discussed.

Contribution Nos. 897, 898, and 931, Department of Entomology, Kansas Agricultural

Experiment Station, Kansas State University

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1970

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

Washington, D.C. 20402 - Price $1.75 (paper cover)
Norman Marston Revision of New World

Species of Anthrax

(Diptera: Bombyliidae) 3

Other Than the Anthrax

albofasciatus Group

Introduction

The genus Anthrax is one of many poorly known taxa

of Bombyliidae (Diptera) in the New World. Most

of the species are based on inadequate descriptions,

which resulted in misconceptions of the species and

in many synonyms. It often has been impossible to

determine identities of species except by studying the

types (Painter and Painter, 1962). Additionally,

many undescribed species have confused later authors

attempting to determine identities of named species.

The genus is separated into five species groups for

convenience in treatment. The albofasciatus group was

revised earlier (Marston, 1963), leaving the cephus,

oedipus, trimaculatus, and tigrinus groups to be

treated here. A future study will deal with the known

pupae. A description and the synonymy of the genus

were given in the previous paper, while the key to the

species groups is given herein.

The author wishes to thank the many people who

have given advice and encouragement during the

course of this study. Particular thanks are due to the

late Dr. Reginald H. Painter of Kansas State University, whose enthusiasm and patient guidance were a

continuous source of inspiration throughout the study.

In addition, much of the basic taxonomic work was

done with specimens in Dr. Painter's collection. Valuable assistance also was given by my wife, Barbara,

and by Dr. Carl W. Rettenmeyer of Kansas State

University.

Norman Marston, Biological Control of Insects Research

Laboratory, USDA ARS ENT, P.O. Box A, Research Park,

Columbia, Missouri 65201.

Thanks are also due to Prof. R. Defretin of the

Musee d'Histoire Naturellc, Ville de Lille, for searching for the type of Spogostylum mystaceum Macquart,

to Mr. Luis Pena of Santiago, Chile, for searching

for the types of Philippi's species, to Dr. Karl-Johan

Hedqvist of the Royal College of Forestry, Stockholm,

Sweden, for furnishing information on types of

Nemotelus tigrinus DeGeer, and especially to Dr. R. H.

Painter for supplying notes on type specimens in

European museums.

I am grateful to the following institutions and individuals for lending material on which the major portion of this study was based (abbreviations used in the

text follow the name of each institution or individual):

Academy of Natural Sciences of Philadelphia (ANSP),

American Museum of Natural History (AMNH), British Museum (Natural History) (BM), California

Academy of Sciences (CAS), Canadian National Collection (CNC), Chicago Museum of Natural History

(CMNH), Cornell University (CNLL), Florida State

Plant Board (FSPB), Illinois Natural History Survey

(INHS), Instituto Oswaldo Cruz (IOC), Museu de

la Plata (MLP), Museu de Historia Natural de Santiago (MHNS), Museum of Comparative Zoology

(MCZ), Oregon State University (OSU), Riksmuseum of Stockholm (RIKS), Secretaria da Agricultura

do Sao Paulo (SASP), Staatliches Museum fur

Naturkunde of Stuttgart (STUT), United States National Museum (USNM), Universidad de Tucuman

(TUC), Universidade do Parana (PAR), University

of Arizona (ARIZ), University of California at Davis,

Berkeley, and Riverside (UCAL), Snow Entomolog-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ical Museum at the University of Kansas (KANS),

University of Michigan (MICH), University of Nebraska (NEBR), Washington State University

(WSU), Messrs. Lawrence R. Ertle (ERTLE), Jack

C. Hall (HALL), Karol Lenko (LENKO), R. H.

Painter (RHP), Luis Pefia (PENA), John Poorbaugh

(JP), and Wolfgang K. Weyrauch (WEYRAUCH).

The author's collection is designated (NLM).

In addition to support from the Kansas Agricultural

Experiment Station, portions of this study were supported by National Science Foundation Grant G-21291

(Dr. Painter, principal investigator), a Fulbright Fellowship for study in Brazil, and National Science

Foundation Cooperative Graduate Fellowships.

The maps were made on Goode Base Map series

No. 2, published by the University of Chicago, which

owns the copyright.

Anthrax Species Groups

The species groups segregated in this paper are poorly

denned and may be polyphyletic in some cases, or at

least with some different species complexes arising

from different Old World ancestors. No good diagnostic characters have been found to fit all species in a

given group and the limits of some of the groups are

open to question. The groups are used here only for

convenience in separating the bulk of species in the

genus into workable units and should not be construed

as being equivalent to subgenera. A worldwide revision

of the genus would be required to ascertain the relationships between Old World and New World species,

and to learn the true identity and limits of the higher

taxa for which some names have been proposed. The

following key separates the groups of the genus present

in North and South America.

Key to the Species Groups of the Genus Anthrax in North and South America

1. Pigmentation of wings solid, not broken into spots (Plate 2r); if wings nearly hyaline then

setae on face extending almost to antenna (Plate lm) cephus group

Wings with isolated or coalesced spots or with bands along veins; if mostly hyaline, then

setae on face restricted to lower half and lateral margins (Plate In) 2

2(1). Cell 2Ma divided by a crossvein between m crossvein and vein Cui, wing veins margined

with brown, discs of cells mostly hyaline (Plate 2p) tigrinus group

Cell 2Ms not divided by a crossvein; wings without bands along veins 3

3(2). Medial angle of m crossvein with a short spur and usually with a distinct spot, vein R«

with a sharp medial angle and short spur (except in A. cybele Coq.); numerous spots

usually present besides those at bases of cells (Plate la-;; Plate 2a—o). . . oedipus group

Medial angle of m crossvein curved, without a spur or spot; medial angle of vein R« sinuate

or evenly curved, without a spur. Spots on wings restricted to bases of cells (sometimes

expanded), except sometimes medially in cell Ri and medially on vein 2A (indistinct

spots at tips of veins Cui and Cua and pigment extending along vein R< and m crossvein

in one species) 4

4(3). Neotropical species trimaculatus group

Nearcitc species albofasciatus group

Nomenclatorial History

Most of the descriptions of species of Anthrax by early

authors were based on one or a few specimens and were

so brief that it has often been impossible to determine

the true identity of the species. The lack of communication among the early authors also resulted in the creation of many synonyms, which has further inhibited

the work of recent students. Only since a study of the

types of North and South American Bombyliidae was

made by Painter and Painter (1962 and unpublished

notes) has it been possible to undertake a complete

revision of the genus Anthrax. The species Anthrax

midas Fabricius, for example, had not been recognized

from the time it was described and had been placed

in the genus Villa (Anthrax, auctt.) in catalogs until

it was studied by Painter and Painter.

Outside the albofasciatus group, the first New World

species of Anthrax was described as Nemotelus tigrinus

by DeGeer (1776). During the first two-thirds of the

nineteenth century most authors placed their species

in the genus Anthrax, which was then interpreted more

widely than at present. The first five species to be described were oedipus, simson, cephus, midas, and

gideon by Fabricius in 1805. All were from South

America. These were followed by Say's (1823) descriptions of analis from Georgia, irroratus from the
NUMBER 4 3

Rocky Mountains, and scriptus, a synonym of tigrinus,

from Pennsylvania. Wiedemann (1828) described

several species in the genus Anthrax, among which

were acroleucus and hyalacrus from Brazil, argyropygus for which he gave the locality "Vaterland?,"

and pluto from Kentucky. He was confused as to the

range of variation of gideon, and acroleucus is a synonym of it. Macquart (1834) described georgicus, a

synonym of analis, from Georgia, added angustipennis

and luctuosus from French Guiana, funebris from the

Dominican Republic in 1840, and "irrorata," a

synonym of irroratus Say, from Carolina and Georgia.

Also in 1840, he described mystaceus from Brazil or

Chile in the new genus Spogostylum. In 1845 he described binotatus, which has been shown to be the

female of angustipennis, in 1848, trimaculatus from

Brazil, and in 1849, Exoprosopa punctata, a synonym

of oedipus Fabricius from "Amer. Merid." In 1855,

Macquart named leucopygus from South America,

apparently without realizing that this was a homonym of leucopygus Macquart, 1840, from Timor.

D'Andretta and Carrera (1952) proposed the name

macquarti to replace the junior homonym.

Among the many species described by Walker in the

cephus group were delicatulus in 1849 from Jamaica,

followed by repertus from South America, cedens from

the United States and punctum and aequa from Brazil

in 1852, and inexactus from the Amazon in 1857. It is

shown here that cedens is simply a form of analis and

that inexactus is only a variant of repertus, while

punctum and aequa are synonyms of oedipus Fabricius.

Blanchard (1852) described vicinus from Chile, but the

name was preoccupied by vicinus Macquart (1840).

Rondani proposed the name inordinatus for the same

species in 1863. Philippi added squalidus, bellulus, and

deodecimpunctatus from Chile in 1865. The latter is

regarded as a synonym of mystaceus (Macquart).

During the last third of the nineteenth century most

authors placed the present species of Anthrax in Argyramoeba and several species were described in this

genus. Schiner (1868) described propinqua, caloptera

and imitans from Venezuela, and poecilophora from

Chile. Anthrax propinquus is now applied to a form of

gideon from northwest South America, while caloptera

is a synonym of oedipus, imitans is a synonym of trimaculatus and poecilophora is a synonym of inordinatus.

In 1869, Loew described contigua from Virginia, a

synonym of argyropygus, along with delila from California and stellans from Oregon. Osten Sacken (1886)

described sereipunctatus from "N. Sonora." Bigot

(1892) added albosparsa from "Colorado," aterrima

from Maryland, melanopogon from Washington, and

plurinota, crinita, and inappendiculata from Chile. It

is shown here that crinitus is a synonym of squalidus

while inappendiculatus is a synonym of bellulus.

Bigot also described a headless specimen as Hemipenthes latelimbatus from "Carolina." The latter is a

synonym of the form cedens of analis. The last species

described in the 1800s was Argyramoeba cybele

Coquillett (1894) from southern California.

From 1900 to 1920, species of Anthrax were placed

in either Argyramoeba or Spongostylum. Williston

(1901), however, placed his new species, pluricellus,

from Vera Cruz, Mexico, in Anthrax, which then was

interpreted as Villa is now. Johnson (1913) studied

the species of Anthrax (then Spongostylum) related to

analis and described slossonae from Kentucky, occidentalis from the western United States, and grossbecki from Florida. It has been shown that slossonae

is a synonym of aterrimus, occidentalis is a synonym of

analis, and grossbecki is only a form of analis. In 1919,

Cole added Spongostylum argentatum from Oregon.

The last species to be described were Anthrax guianicus

Curran (1934) from British Guiana, minimaculatus

Oldroyd (1937) from Argentina, and cintalpa and

mexicanus Cole (1952) from Chiapas, Mexico.

Anthrax guianicus and mexicanus are synonyms of

midas Fabricius.

Taxonomic Characters and Techniques

The characters used for separation of species vary between groups and are therefore discussed under each

group. In general, the wings offer the most useful characters and these are illustrated for almost all species.

The Comstock-Needham system is followed in naming

the veins and cells for the most part.

The male and female genitalia present useful characters in most groups and are therefore illustrated. The

terminology is artificial in part, so the specific structures

are named in Figures 8,50, and 128.

The techniques for preparing the genitalia may be

useful to other students. First, the specimens to be

studied are placed in a relaxing jar for approximately

six hours. This is long enough to preclude breakage

during handling, yet is not so long that the insects

become wet, thus destroying the patterns of scales and

pollen. The specimen is grasped with forceps near the

apex of the abdomen and the genitalia are removed
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with an insect pin. Care should be taken to remove

all of the sixth sternum of the female or the bulbs of

the spermathecae may be broken off. The genitalia are

then placed in a solution of potassium hydroxide

(about 10%) in a shell vial and left to soak at room

temperature for 24 hours. At the end of this time the

softer tissues are removed and the genitalia are ready

for study. By this method, a relatively large number of

specimens may be handled at one time.

The genitalia are glued to the tip of an insect pin

with clear fingernail polish—the male genitalia attached ventrally at the junction of the gonocoxites and

the female genitalia exteriorly just below the cerci.

The pin is then placed in a dish of glycerine on two

paper clips. A third paper clip is placed over the pin

to hold the pin in place after the specimen has been

positioned for drawing. The glycerine prevents the nail

polish from hardening so that the specimen's position

may be adjusted on the pin, and it may be easily removed. After the drawings are completed the genitalia

may be preserved in a small vial pinned below the

specimen.

Maps illustrate the distribution of species that are

known from more than a few specimens. Each dot represents a locality from which one or more preserved

specimens have been seen. The hypothetical ranges

(shown by shading) indicate areas ecologically similar

to areas from which specimens have been recorded

and were drawn primarily from vegetation maps of the

areas under consideration.

Biology and Ecology

Species of the genus Anthrax are all parasitoids on

larvae of holometabolous insects living in tubular nests

or cells as far as is known. Life histories of the species

are probably similar to that of Anthrax limatulus fur

(Osten Sacken) described by Marston (1964), with

eggs deposited in openings of cells in a variety of nest

locations, depending upon the species. Specimens have

been collected in almost every conceivable ecological

area from the northernmost extensions of the Arctic

forests to south Chile.

The oedipus group is composed of two distinct species

complexes distinguishable by differences in wing pattern. The oedipus subgroup occurs throughout North

and South America and contains well-defined species,

which seem to have been distinct for a relatively long

time as evidenced by the diversity of morphological

structures, especially the male and female genitalia.

The pluto subgroup, on the other hand, is found only

in North America and the West Indies. The species

seem to be very closely related, except for A. cybele,

and show only minor differences in morphological

structures. It seems probable that the two subgroups

were derived from separate Eurasian ancestral stocks

with the oedipus subgroup arriving in North America

much earlier. Anthrax cybele probably also was derived from a separate ancestor, but it is included within

the pluto subgroup for convenience.

The species of the oedipus group, primarily inhabitants of open forested areas with some adapted to

desert areas with xerophytic shrubs, parasitize a wide

range of higher Hymenoptera nesting in old logs and

stumps, in banks, and in soil. The species collected by

the author seldom leave their local habitat and never

have been taken on flowers. Females often are seen

ovipositing on old stumps and logs, in cracks and

holes in vertical banks, and occasionally in holes in

open, flat areas. They sometimes are attracted to the

collector and will oviposit on dark spots on clothing.

Little is known about the biology of the species other

than host records. The little information available indicates that their life history is similar to that of A.

limatulus fur (Osten Sacken) (Marston, 1964).

With the exception of analis and argentatus, the

species of the cephus group occur predominantly in

warm temperate and tropical climates. By far the greatest number of species occur in the area between southern Mexico and southeastern Brazil. Anthrax argentatus probably was derived from an ancestral form

adapted to a cold temperate climate, since related species occur in the Palaearctic region, and it now occupies

a range from southwestern Canada south in the mountains to central California and Colorado. Anthrax

analis, on the other hand, probably has become adapted

secondarily to cold climates as well as warm ones, since

its closest relatives are Neotropical and it now may be

found into north-central Canada. Apparently no species of the cephus group has been able to adapt itself

to the temperate climates of southern South America.

The southernmost record is for gideon from Montevideo, Uruguay.

Ecologically, the species of the cephus group predominantly are found in forest habitats (including

forest glades), although several species have become

adapted to drier areas. Anthrax nitidus is found in

the desert area of southern California and southwestern

Arizona, although its closest relative, argentatus, is

found in forested areas. Several species have become
NUMBER 4 3

adapted to the dry climate of northeastern Brazil, but

these usually also occur in forest areas, indicating that

the adaptation is relatively recent or that they occupy

habitats that are similar in the two areas. The latter

may be the case with repertus, which has been collected

in grassy areas in the tropical forest of southeast Brazil.

Anthrax inaquosum is the only species restricted to

the desert of northeast Brazil as far as is known.

Anthrax analis is unique in the group (with the possible exception of funebris) in that it has become

adapted to sandy areas where it has been reared from

the larvae of tiger beetles (Cicindellidae).

Little is known of the biology of the species. With

the exception of analis, the only records of hosts are

from wasps (Hymenoptera: Sphecidae). The species

probably parasitize a wide range of holometabolous

insects within the limits of their preference for

oviposition sites.

The trimaculatus group is restricted to South and

Central America as far as is known. The species apparently are found throughout the continent, although,

due to lack of collecting, large gaps occur in such areas

as southern Argentina. The species have been collected

in a variety of habitats, along paths, in clearings, and

flying over vertical banks, where wild bees were nesting.

They seem to fill niches similar to those occupied by the

species of the albofasciatus group in North America.

Biological notes are available only for trimaculatus, a

specimen of which was reared from the nest of a species

of Diadasia (Hymenoptera: Apoidea).

The species of the tigrinus group occupy a variety of

habitats, although each is apparently restricted to a

broad ecological zone. All reared specimens have been

taken from the nests of species of Xylocopa (Hymenoptera: Apoidea), although they may also parasitize

species of related genera with similar nesting habits.

The facts known of the biologies of the species are

discussed by Hurd (1959), although he considered

the entire group to be a single species, Anthrax simson

Fabricius.

Anthrax tigrinus is known from the eastern deciduous forests so that its range is limited on the north by

the coniferous forests and on the west by the Great

Plains. Similarly, delila is found in broadleaf deciduous

and evergreen forests in southern and central California, although it occurs in areas dominated by coniferous forests in northern California. A relict population also occurs in southern Baja California. Both

tigrinus and delila have been reared from species of

Xylocopa nesting in structural timbers.

Anthrax xylocopae has become adapted to drier areas

than are inhabited by tigrinus and delila, since it occurs

in pinion-juniper forests and grasslands from eastern

Arizona and north-central Mexico to central Texas. It

has been reared from Xylocopa nests in dead-flower

stalks of Yucca and probably parasitizes bees nesting

in deadwood as well. In central Arizona xylocopae is

allopatric to simson habrosus, the latter occurring in

desert areas from southern California and Arizona

south along the Pacific coast of Mexico and in dry

tropical forests in southern Mexico as far east as Chiapas. No reared specimens of s. habrosus have been

studied, but it probably parasitizes Xylocopa brasiliorum varipuncta Patton, which nests in a variety of

woods throughout southern California, western Arizona, and northwestern Mexico. Anthrax simson simson occurs in wet tropical forests in eastern Mexico,

Central America, and northern South America. Nothing is known of its biology; the record cited by Hurd of

a bombyliid taken from the nest of Xylocopa augusti

Lepeletier at Araucaria, Parana, Brazil, might refer

to simson.

Anthrax oedipus Group

The oedipus group is a well-defined aggregate of species

in North and South America. They generally are

characterized by numerous spots on the wings at the

bases of the cells, on apices of veins, and often scattered

along veins and expanding into the cells. Setae on the

face are concentrated along the epistomal margin with

a few along the eye margins (Plate In). Light and dark

scales on the thorax and abdomen are formed in a

complex pattern with some enlarged and truncate

along the posterior and lateral margins of the abdominal terga. The gonocoxites of the male genitalia taper

apically and are not lobed; the distal segment of the

gonostylus is flat, quadrangular basally, and ends in a

deflexed hook. The base of the third antennal segment

is wider than the second segment, hemispherical, and

more or less as long as the styliform part.

Group Description

Body generally black, blue gray or brown pruinose,

gray or silvery pollinose along eye margins; femora,

tibiae, male genitalia, and basal antennal segments

sometimes dark red or orange. Front with black setae

and, on lower half, lanceolate scales. Face retreating,

epistomal margin abrupt; upper half of face bare ex-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cept for some setae and a few scales along eye margins; lower half of face with coarse black setae and

linear or oblanceolate white or yellowish scales extending along eye margins into oral cavity (Plate In).

Occiput with short, semirecumbent black setae and

linear and lanceolate scales; fringe of pile on posterior

margin dark brown or black, sometimes with light

tips. First antennal segment narrow basally, enlarged

mesad apically, slightly shorter to slightly longer than

apical width. Second segment lenticular with apical

margin rounded (Figure 15; Plate Ik) or saucer

shaped with apical margin produced as a sharp flange

(Figures 14, 16; Plate 1/); base of third segment

bulbous, slightly wider than second segment in male,

distinctly wider in female; styliform part arising

medially or toward lateral margin, slightly shorter to

distinctly longer than basal part; style short, about onefourth as long as styliform part, tuft of hairs about as

long as style.

Disc of mesonotum with sparse, fine black setae and

linear, mostly black scales; white scales in transverse

line in front of bases of wings, in submedial longitudinal lines on anterior half, along anterior edge, and in

a triangle extending forward from posterior margin;

brown scales sometimes intermixed with white and

occasionally predominating on disc. Lateral margins

of mesonotum with black setae and bristles and linear

or elongate, threadlike, white or brown and white

scales. Scutellum with linear or lanceolate black scales

on disc and lanceolate, oblanceolate or obovate white

scales on anterior and posterior margins; brown scales

often intermixed with white, white or brown scales

sometimes in medial patch on disc, black scales occasionally restricted to submedial spots. Sternopleuron,

lower half of mesopleuron, and anterior half of pteropleuron with black setae and linear or lanceolate,

recumbent to erect scales; some coarse setae on pteropleuron. Upper half of mesopleuron with coarse black

or gold setae, and curly scales; pile sometimes present.

Pile on prosternum, propleuron, and anterior margin

of mesonotum white, black or mixed, some brown hairs

often present, black setae present on anterior margin

of mesonotum. Postalar tuft of pile black, white or

yellowish brown, occasionally mixed, usually a few

white scales at base. Hypopleuron bare, metapleuron

usually bare, sometimes with a few scales behind and

below spiracle. Coxae with black setae and black or

white lanceolate or oblanceolate scales, occasionally

with some brown scales.

Wing with basal infuscation extending into basal

cells and with numerous spots in apical part of wing

at bases of cells and usually at angles and apices of

veins, sometimes also at other points on veins; spots

often more or less coalesced into medial and subapical

bands and sometimes elongated into transverse striae.

Cells C and Sc with alternating hyaline and pigmented

areas or entirely hyaline, subhyaline, or infuscated.

Spurs at bases of cells R2+3 and R4, and at medial

angle of m crossvein, and often at medial angle of vein

R4 and basal angle of m crossvein; spur at base of cell

R4 sometimes connected to vein R2+s to form a sectoral

crossvein. Anal cell slightly open or closed in margin;

r-m crossvein at basal third or two-fifths of discal

cell; junction of cells 1M2 and Cux one to three times

length of base of cell Cui. Cell 2A one to 1.5 times

as wide as cell 1A; alula well developed; calyptere

unpigmented, fringe of hairs white (Plate la-;'; Plate

2a-o). Stem of halter brown, base of knob dark brown,

apex of knob white.

Femora with oblanceolate-cuneate or obovate-cuneate scales, usually black anteriorly and white posteriorly on fore and middle pairs, entirely black or white

posteroventrally and black anterodorsally on hind

pairs; black scales sometimes more or less replaced

by brown. Scales on tibiae usually white posteriorly

and black anteriorly on fore and middle pairs, entirely

black on posterior pairs. Fore femora with anteroventral row of bristles strongest toward base; middle

femora with posteroventral row of bristles and occasionally with one or more bristles postmedially on

anterior surface; hind femora with anteroventral row

of bristles in female and additional posteroventral

row in males.

Lateral margin of first abdominal tergum with white

pile and a few black setae, occasionally some black

pile, sides of second tergum with black setae and linear,

lanceolate, oblanceolate or obovate-cuneate scales of

variable color, occasionally some black pile; sides of

posterior terga with oblanceolate and obovate-cuneate

scales of variable color, occasionally some black pile;

sides of posterior terga with oblanceolate and obovatecuneate scales and black setae. Posterior margins of

first tergum with a few black setae and with light

scales laterally and dark scales medially; discs of

remaining terga with sparse setae and complex pattern

of linear or threadlike to ovate-cuneate scales; usually

with broad white scales in submedial or medial and

sublateral patches on posterior margins, less distinct

posteriorly; smaller scales cover remainder of disc,
NUMBER 4 3

brown or white in basal and subapical, poorly defined

bands on second segment and basally on posterior

segments, black elsewhere. Sterna with sparse black

setae and linear or threadlike to ovate scales; light

scales usually predominate posterolaterally and dark

scales anteromedially.

Apex of gonocoxites of male genitalia simple, not

elongated into lobes. Basal segment of gonostylus

platelike, rounded dorsally, not produced apically; distal segment laterally flattened, quadrangular basally,

apex a deflexed hook. Apex of epiphallus small, curled

upward at tip dorsally, slightly flared laterally, with

a basally produced spine just after junction of dorsal

bands; epiphallus sometimes atrophied. Apex of aedeagus acute with gonopore ventral, or flared with gonopore apical, sometimes trilobed. Base of aedeagus

bulbous, narrowing abruptly to apical part. Epandrium

simple or with dorsal margin apically produced and

with cerci produced ventrad.

Tenth tergum of female with four to twelve spines

on each side. Sclerite on each side of gonopore hatchet

shaped; lateral lobe acute, clavate or broad and blunt;

dorsal or dorsomedial lobe short, acute or obtusely

rounded; ventral lobe narrow and parallel sided or

broadened medially, curving inward at apex. Ducts

of spermathecae uniting medially into common duct or

sometimes emptying separately into genital chamber,

with one to seven convolutions before expanding to

bulbs; bulbs elongate, expanded medially, postmedially

or sometimes apically, sometimes bent medially or

basally.

Taxonomic Characters

The wings of the species of the oedipus subgroup show

many differences between taxa, but the differences in

pattern are subtle and difficult to describe (Plate la-;).

That difficulty apparently has led some authors to

lump all the species together in despair, although other

less readily observed characters will separate the

species. Discrete differences in the pattern of the wings

often are obscured by variation in intensity of the pattern in different localities. That is especially true of

irroratus irroratus where the wings vary from lightly to

heavily pigmented in different parts of North America

(Plate \a-c). In the pluto subgroup the wing pattern

remains almost the same (except for cybele), while

differences between the species are due to restriction

or extension and coalescence of spots (Plate 2a-o).

The sexes vary in this subgroup with females having

darker wings. The spur at the base of cell R« may be

connected to vein R2+3 to form a complete sectoral

crossvein in the oedipus group. The character is stable

in some species; in others it varies.

In most species of the oedipus group the second

antennal segment is saucer shaped with the apical margin produced as a sharp flange (Figures 14, 16; Plate

1/), a character shared with many Palaearctic species.

Only two species in the group, irroratus and cybele,

have the second segment lenticular (Figure 15; Plate

Ik) as it is in all other groups of the genus in North

and South America except the tigrinus group. This is

another character used to separate the "genera" segregated in the Old World by Sack (1909) and Bezzi

(1924), but which is not applicable to New World

species.

The chaetotaxy of the legs seldom has been found

useful to separate species in the Bombyliidae and that

is generally true in the genus Anthrax. Anthrax oedipus and peruvianus, however, have a distinctive bristle

on the middle femur postmedially on the anterior side,

which has not been noted in other species in the group

(Figure 30). That is one of the qualitative characters

that lead to the conclusion that the oedipus subgroup

has evolved longer in the New World than has the

pluto subgroup.

The male genitalia are of little practical use to

separate species in the oedipus group, contrary to the

situation in the albofasciatus group. While some good

internal differences exist, external aspects of the genitalia are quite similar for almost all of the species. The

most striking differentiation is in cordillerensis and

inordinatus of the oedipus subgroup in which the apex

of the epiphallus is atrophied and the apex of the

aedeagus trilobed. Females of the species appear to be

adapted to the trilobed aedeagus in that they have

three spermathecal ducts emptying separately into the

genital chamber rather than uniting into a common

duct. Those two species also have the epandrium

elongated over the cerci so they project ventrad.

Female genitalia have several good internal characters,

the shape of the spermathecae, the number of convolutions in the spermathecal ducts, and the shape of the

sclerites on each side of the apex of the spermathecal

ducts, but a part of the abdomen has to be destroyed to

examine them.

Variation in vesture of the body is extensive. In

general, there appear to be four types of ectodermal

processes in the genus—macrochaetae (or bristles),
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

setae, pile (hairs), and scales—although the cellular

origins of the different types have not been investigated. While both setae and macrochaetae are tapered,

macrochaetae usually are larger, few in number, and

found in well-defined locations, whereas setae are finer,

more numerous, and more or less scattered where they

occur. Pile is formed of erect, parallel-sided hairs with

minutely bristly apices. No intermediate structures

have been found between pile and scales in the genus

Anthrax. Scales are the most variable component of

the vesture, in color, shape, and size, and often present

useful characters. They may be white, black or various

shades of brown, and vary from short and linear or

lanceolate to long, ovate-truncate. Some of the various

shapes discriminated are illustrated in Figures 17-26.

An accurate classification is impossible since all sizes

and shapes intergrade. "Tomentum" has been used in

other genera of Bombyliidae for dense, woolly, hairlike

scales, but such have not been observed in the genus

Anthrax.

Key to the North and South American Species of the oedipus Group of the Genus

Anthrax Scopoli

1. Cells C and Sc of wing with alternating pigmented and hyaline areas (except rarely

in A. cordillerensis from Chile). Veins in outer part of wing with numerous spots

other than at bases of cells and at apices of veins (Plate la-;). North and South

America. Anthrax oedipus subgroup 2

Cells C and Sc of wing without alternating pigmented and hyaline areas, either evenly

infuscated or partly or wholly hyaline or subhyaline. Veins in outer part of wing

without spots other than at bases of cells, apically on some veins, subapically on

vein Rj+s and submedially on R» (Plate 2a-o). North America and West Indies.

Anthrax pluto subgroup 10

Anthrax oedipus subgroup

2(1). Second antennal segment lens shaped, rounded apically (Figure 15; Plate Ik). North

America 3

Second antennal segment saucer shaped, produced apically as a sharp flange (Figures

14, 16; Plate 1/). North and South America 4

3(2). Wing with some independent rounded spots, usually many (Plate la-c). Usually

numerous white scales anteriorly on second and following abdominal sterna. Small

subspecies, wing length 4.9-9.2 mm. North America other than Arctic tundra, coastal

plain of southeastern United States and Central America (Map 1)

irroratus irroratus Say

Wing with dense pattern of transverse striae; with few, if any, independent rounded

spots (Plate Id). Abdominal sterna without white scales anteriorly. Large subspecies,

7.4-9.8 mm. Florida north to South Carolina and west to Alabama (Map 1)

irroratus striatipennis, new subspecies

4(2). Cell Sc of wing evenly infuscated or subhyaline, or with indistinct spots. Chilean and

Peruvian species 5

Cell Sc of wing with distinctly alternating pigmented and hyaline or subhyaline areas.

Southwestern United States, Mexico, West Indies, and South America other than

Chile 7

5(4). Middle femur without bristles above anteroventral row (Figure 36). Abdominal terga

without golden brown scales, wing with complete sectoral crossvein. Chile 6

Middle femur with distinctive bristle or bristles on anterior side above anteroventral

row (Figure 37). Abdominal terga with large patches of golden brown scales. Wing

without complete sectoral crossvein. Southwestern Bolivia (?) to Ecuador (Map 4).

peruvianus, new species

6(5). Wing with complete sectoral crossvein (Plate li). Femora orange. Abdomen with large

patches of golden brown scales. Central Chile and southwestern Argentina (Map 5).

inordinatus (Rondani)

Wing without complete sectoral crossvein (Plate 1;). Femora black. Few, if any, golden

brown scales on abdomen. Andes Mountains in central and southern Chile, and

southwestern Argentina (May 4) cordillerensis, new species
NUMBER 4 3

7(4). Middle femur with distinctive bristle on anterior side above anteroventral row

(Figure 37). Postalar tuft of pile white or black. Mexico and West Indies through

South America other than Chile 8

Middle femur without bristles anteriorly above anteroventral row (Figure 36). Postalar

tuft of pile usually black or with few white hairs. Southwestern United States to

Venezuela and Colombia (Map 2) cintalpa Cole

8(7). Dorsum of abdomen with black and white scales, no large patches of brown scales.

Abdominal sterna three, four, and five with numerous black scales. Lateral margins

of second tergum with only black scales and pile. Mexico and West Indies through

South America other than Chile and Peru (Map 3) 9

Large patches of golden brown scales on abdominal terga. Few, if any, black scales

on third, fourth, and fifth sterna. Lateral margins of second tergum with tuft of

brown and white scales posteriorly. Southwestern Bolivia (?) to northern Peru

(Map 4) peruvianus, new species

9(8). Wing with complete sectoral crossvein (Plate In). Postalar tuft of pile black. Southern

Brazil and adjacent areas (Map 3) oedipus aquilus, new supbspecies

Wing without complete sectoral crossvein (Plate lg). Postalar tuft of pile white.

Southern Mexico to southern Brazil and northern Argentina (Map 3)

oedipus oedipus Fabricius

Anthrax pluto subgroup

10(1). Second antennal segment saucer shaped, apical margin produced as a sharp flange

(Figures 14, 16; Plate 1/). Independent spots usually present apically on vein Ra+»

or medially on R* 11

Second antennal segment lens shaped, apical margin rounded (Figure 15; Plate Ik).

Independent spots absent apically on vein R2+J and medially on R* (Plate 2o)

cybele (Coquillett)

11(10). Cell 2M of wing hyaline except at extreme base and apex. Vein 2A with at most a

faint cloud submedially in some females (Plate 1c—d) 12

Cell 2M of wing partially or completely infuscated medially. Vein 2A with at least a

small cloud submedially in males and with a distinct spot in females (Plate la-b,

e-n) 13

12(11). Abdominal sterna one through three with long, linear, white scales predominating.

Sternopleuron and lower halves of mesopleuron and pteropleuron with white scales

only. Terga two through five usually with linear white scales across entire posterior

margins or narrowly broken medially by black scales. Texas to southern California,

south into Sonora and Baja California (Map 8) seriepunctatus (Osten Sacken)

Abdominal sterna one through three with few, if any, linear white scales. Sternopleuron,

and lower halves of mesopleuron and pteropleuron with mixed black and white

scales. Posterior margins of terga two through five with submedial and sublateral

patches of lanceolate or cuneate white scales interrupted by areas of black scales.

Texas to southern' California, south into Mexico and north into Great Basin

(Map 9) atriplex, new species

13(11). Scales on hind femur entirely black or with few white ones posteriorly at base. Underside of abdomen with few, if any, golden brown scales; numerous black scales often

present 14

Hind femur with golden brown scales ventrally and black scales dorsally. Long, curly

scales on abdominal sterna two through seven white on posterior margins, golden

brown subapically and sometimes basally; few, if any, black scales present. Canada

south in Coast Ranges and Sierra Nevada to southern California, in Rockies to

Colorado and in Appalachians to Georgia (Map 8) stellans (Leow)

14(13). Upper half of mesopleuron with mixed black, white, and often brown hairs, and black

setae. Pile on lateral margins of first abdominal tergum often yellowish white or with

some black hairs posteriorly. Continental North America 15

Upper half of mesopleuron with dense, snowy white pile and few yellow or black

bristles, no black or brown hairs present. Lateral margins of first abdominal tergum

with snowy white pile. West Indies (Map 7) insulanus, new species
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

15(14). Cell 2M entirely infuscated or with at most a small medial subhyaline area in some

males. Submedial spot on vein 2A broadly connected to base (Plate 2e-f). Eastern

United States, Texas, and eastern and southern Mexico (Map 6) 16

Cell 2M with at least a small subhyaline area in females, largely hyaline in males.

Submedial spot on vein 2A separated from base (Plate 2k-n). Western United

States, Texas, and northern Mexico 17

16(15). Abdominal sterna one through three with threadlike, mostly white scales, with a few

black ones on two and three. Lateral margins of first abdominal tergum with white

pile ventrally, with few black hairs posteriorly. Mississippi valley from Appalachians

to Great Plains, south to Oklahoma and Tennessee (Map 6)

pluto pluto Wiedemann

Scales on abdominal stema two through seven entirely black, or few white ones present

on posterolateral margins. Ventral part of lateral margins of first abdominal tergum

with black pile or with few white hairs intermixed. Texas to Florida, south along

Gulf coast to south-central Mexico (Map 6) pluto nigriventris, new subspecies

17(15). Scales on abdominal sterna two through seven mostly black, few white or yellow ones

sometimes present along posterior margins. Postalar tuft of pile usually predominantly

white. Dry oak or mesquite forests from southern California to Texas and into

northern Mexico (Map 9) atripUx, new species

Abdominal sterna with long, threadlike white scales, sometimes with few black scales

medially on three through seven. Postalar tuft of pile usually predominantly black.

Coniferous forests from southern California to British Columbia and south in Rocky

Mountains to northern New Mexico (Map 10) melanopogon (Bigot)

Anthrax irroratus irroratus Say

Anthrax irroratus Say, 1823, p. 46.—Osten Sacken, 1858,

p. 40 [irrorata].—Say, 1859, p. 61.—Curran, 1927, p. 85

[irrorata].—Maughan, 1935, p. 33 [part] [irrorata].—

Priddy, 1939, p. 45 [irrorata].—Brooks, 1952, p. 370

[irrorata].—Painter and Painter, 1965, p. 432.—Krombein,

1967, p. 403.

Anthrax irroratus Macquart, 1840, p. 60 [irrorata].—Macquart, 1848, p. 34 [irrorata].—Osten Sacken, 1858, p. 40

[irrorata].

Anthrax oedipus.—Wiedemann, 1828, p. 262.—Osten Sacken,

1858, p. 41 [part].—Johnson, 1925, p. 108.—Curran, 1927,

p. 85.—Edwards, 1930, p. 173 [part].—Brimley, 1938,

p. 341.—Strickland, 1938, p. 195.—Hall, 1954, p. 146

[not Fabricius, 1805; misidentification].

Argyramoeba oedipus.—Osten Sacken, 1877, p. 243 [part].—

Osten Sacken, 1878, p. 90 [part].—Osten Sacken, 1886,

p. 102 [part].—Townsend, 1893, p. 60.—Coquillett, 1894,

p. 95 [part].—Davidson, 1894, p. 170.—Baker, 1895,

p. 173.—Kertte, 1909, p. 66 [part].—MaUoch, 1917,

p. 392.—Rau, 1926, p. 231 [Argyromoeba] [not Fabricius,

1805; misidentification].

Spongostylum oedipus.'—Aldrich, 1905, p. 223 [Spogostylum].—Cole and Lovett, 1921, p. 244 [Spogostylum].—Cole,

Malloch, and McAtee, 1924, p. 185 [Spogostylum] [not

Fabricius, 1805; misidentification].

MALE.—Head with black setae and lanceolate,

black and white scales. Second antennal segment lenticular, apex not produced as sharp flange (Figure 15;

Plate Ik); base of third segment bulbous, about 1.33

times wider than second segment, styliform part about

1.33 to 1.5 times longer than base, about 2 to 3 times

longer than style.

Mesonotum and scutellum with linear, black and

white scales, curly along margins, brown scales sometimes present. Anterior half of pteropleuron, mesopleuron, and sternopleuron with black setae, and black,

white, and sometimes brown pile and scales. Pile on

prosternum, propleuron, and anterior margin of mesonotum white or mixed black and /or brown and white.

Postalar tuft of pile black, often with some white or

brown hairs, rarely completely white. Metapleuron

sometimes with patch of lanceolate white or yellowishwhite scales in specimens from western United States.

Coxae with black setae and white or mixed black and

white scales.

Cells C and Sc with distinct spots alternating with

hyaline areas; remainder of wing with dark brown

spots varying greatly in number and extent (Plate

la-c), specimens from eastern United States with spots

coalesced into broad subapical, medial, and basal

bands and with some independent spots elongated

transversely; specimens from western United States

usually with spots smaller, rounded and not coalesced

into bands; specimens from Canada grading from

heavily pigmented eastward to lightly pigmented westward; specimens from Mexico with spots sometimes

coalesced into bands, but with independent spots

rounded. Spur at basal angle of cell R4 rarely connected to vein R2+3 to form a sectoral crossvein. Basal
NUMBER 43 11

part of vein Cux 0.33 to 0.5 as long as complete vein.

Scales on fore and middle femora usually black anteriorly, white or yellowish white posteriorly, sometimes with few black scales posteriorly, often with

white scales anteriorly; scales on hind femur varying

from entirely black to predominantly white. Middle

femur without a bristle anteriorly above anteroventral

row.

Sides of first abdominal tergum with white pile and

some black setae; posterior margin with black scales

medially and white scales laterally. Lateral margins

of remaining terga with short, obovate-truncate to long

oblanceolate black scales and few black setae and hairs,

with white scales posteriorly on three, sometimes on four

and five, and predominating on six and seven. Discs of

terga with linear black scales anteriorly, and with submedial and sublateral patches of ovate-truncate white

scales on posterior margins; lanceolate white, yellowishwhite and/or brown scales often present anteriorly;

white scales on posterior margins often coextensive on

posterior terga and spreading over discs of sixth and

seventh terga. Venter with sparse black setae and sparse

lanceolate scales, usually entirely white or yellowish

white, sometimes partly or entirely black.

FEMALE.—Similar to male. Base of third antennal

segment more enlarged, about 1.5 times wider than second segment. White scales on posterior abdominal

terga less extensive.

MALE GENITALIA (Figure 9).—Apical part of gonocoxites about three times longer than basal part; distal

segment of gonostylus about twice as long as wide. Apex

of epiphallus with sharp flange dorsally; no medial

process above apex of aedeagus. Apex of aedeagus not

flared; gonopore ventral. Epandrium rounded apically

in dorsal view; cercus projecting apically beyond

epandrium.

FEMALE GENITALIA (Figure 38).—Tenth tergum

with nine spines apically on each side; ventral extension

from base narrowed apically. Dorsal part of sclerite on

each side of gonopore strongly produced above lateral

lobe and broadly rounded; lateral lobe truncate, apex

acute dorsally, obtuse ventrally; ventral extension narrowed apically and strongly curved mesally. Ducts of

spermathecae united into medial tube before emptying

into genital chamber, with four or five convolutions before expanding to bulbs; neck of bulbs slightly recurved, remainder geniculate submedially and postmedially, and enlarged apically.

BODY LENGTH.—3.9 to 9.2 mm.

WING LENGTH.—4.1 to 10.1 mm.

DISTRIBUTION.—Anthrax irroratus irroratus occurs

in forested areas throughout North America except in

southern Mexico, Central America, and Florida and

the coastal plain in southeastern United States. It is

allopatric to irroratus striatipennis in the last area

(Map 1).

TYPES.—Anthrax irroratus irroratus was described

by Say (1823) from material collected in the Rocky

Mountains. The specimen or specimens have since been

destroyed. The types of Anthrax irrorata Macquart

(1840) were not mentioned by Painter and Painter

(1962). Macquart's species may be synonymous with

irroratus striatipennis since his specimens were from

Carolina and Georgia, but the name is a primary

homonym of irroratus Say.

BIOLOGY.—This species seems to be a nonspecific

parasite of a variety of hymenopterous insects nesting

in logs, banks, and open, flat areas. Females often are

observed ovipositing in openings of tunnels in stumps

and logs and are sometimes attracted to the collector,

flipping eggs at dark spots on clothing.

Townsend (1893) described the pupal skin of A.

irroratus irroratus (as Argyramoeba oedipus

(Fabricius) from a specimen reared from a species of

Odynerus (Hymenoptera: Vespidae) taken in Colorado. Davidson (1894) reared this species from the

nest of a small bee "made 6 inches deep in adobe

soil." Baker (1895) recorded A. irroratus irroratus

as a parasite of several species of Odynerus; Brooks

(1952) described and figured the pupa reared from

Megachile nivalis Friese (Hymenoptera: Megachilidae); and Hall (1954) reared it from a "Hymenopteran nesting in an old anobiid burrow in a

log." Recently, Krombein (1967) recorded this species

as a parasite of Megachile gentilis Cresson, Dianthidium heterulkei fraternum Timberlake, Ashmediella

bucconis denticulata (Cresson) (Hymenoptera:

Megachilidae), and Hylaeus asininus (Cockerell and

Casad) (Hymenoptera: Colletidae), as well as an

unidentified vespid wasp.

In collections studied by the author there are

specimens reared from Megachile nivalis Friese, M.

rotundata (Fabricius), M. sp., Osmia rostrata Sandh.,

and Osmia clarescens Ckll. (Hymenoptera: Megachilidae) ; Anthophora sp. (Hymenoptera: Apidae) ;

Odynerus sp. and Pseudomasaris coquilletti Rohwer

(Hymenoptera: Vespidae); and Sceliphron sp.

(Hymenoptera: Sphecidae). Others were reared from

"Tapoxylon albitarsi" "solitary wasp cocoon" and

"from mud wasp in cocoon of moth."
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

iVroratus stratipennis, n. ssp. =

irroratus irroratus Say m

MAP 1.—Distribution of Anthrax irroratus irroratus and A. irroratus striatipennis.
NUMBER 43 13

DISCUSSION.—Anthrax irroratus irroratus and irroratus striatipennis differ from other species in the

oedipus subgroup in having the second antennal segment lenticular rather than saucer shaped with the

apical margin produced as a sharp flange. Specimens

from southwestern United States and Mexico also are

usually smaller than those of cintalpa, have the wings

more lightly infuscated, and have oblanceolate and

obovate rather than linear and lanceolate scales on the

sides of the second abdominal tergum. Characters

separating irroratus irroratus from irroratus striatipennis are discussed under the latter.

Anthrax irroratus irroratus is an extremely variable

taxon. Specimens from western United States have

less heavily pigmentcd wings and lighter pile and

scales than those from eastern United States. There is

an even cline across Canada, however, indicating that

the two populations are consubspecific, although the

two extremes are quite different. Specimens from

Mexico also vary in degree of pigmentation of the

wings and extent of dark scales. Darkest specimens are

from wetter areas while the lightest specimens are from

drier areas, which may indicate that differences in

color are due to climatic factors.

Anthrax irroratus striatipennis, new subspecies

Argyramoeba oedipus.—Johnson, 1895, p. 325.—Kert&z,

1909, p. 66 [part] [not Fabricius, 1805; misidentification].

Spongostylum oedipus.—Johnson, 1913, p. 55 [Spogostylum]

[not Fabricius, 1805; misidentification].

MALE.—Head with black setae, and black and white

scales. Second antennal segment lenticular, not produced as sharp flange apically (Figure 16; Plate Ik).

Mesonotum and scutellum with sparse black setae;

scales black and white, curly along margins. Pile on

prosternum and propleuron black, on anterior portion

of mesonotum, dorsal part of mesopleuron, and anterodorsal part of pteropleuron mixed black and white;

postalar tuft of pile black with few white hairs.

Mesopleuron and sternopleuron with sparse black and

white scales. Scales on coxae black and white.

Cells C and Sc of wing (Plate Id) with alternating

hyaline and pigmented areas; remainder of wing pigmented with dark brown, transverse striae, most dense

and coalesced from basal cells out to bases of cells

R-2+3, Rs, and 2M2, and in broad preapical band. Basal

section of vein Cux two-fifths as long as entire vein.

Scales on fore and middle femora mixed black and

white; on hind femur black; middle femur without

a bristle above anteroventral row.

Scales on hind margin of first abdominal tergum

black medially, white laterally; scales on discs of remaining terga black anteriorly, white submedially and

laterally on hind margins, white scales more extensive

posteriorly. Lateral margins of first tergum with white

pile intermixed with black setae and few brown hairs;

remaining terga with lanceolate-truncate and ovatetruncate black scales, and black setae. Venter with

sparse black setae and linear black scales, few linear

white scales present on first sternum.

FEMALE.—Propleuron with some white hairs; scales

on mesopleuron white. Pile on lateral margins of first

abdominal tergum light yellow.

VARIATION.—The prosternal pile may be partially

white. The scales on the coxae and pleurae may be

entirely black. The number of white scales on the fore

and middle femora varies; a few white scales may be

present on the posterior femur. The basal part of vein

Cux varies from two-fifths to one-third the length of

the entire vein. The pile on the lateral margins of

the first tergum varies from yellow to white.

MALE GENITALIA.—Similar to irroratus irroratus

Say.

FEMALE GENITALIA (Figure 39).—Similar to irroratus irroratus. Tenth tergum with 15 spines apically

on each side. Lateral lobe of dorsal part of sclerite on

each side of gonopore acute. Spermathecae less

strongly geniculate subapically.

BODY LENGTH.—7.4 to 9.8 mm.

WING LENGTH.—8.1 to 11.2 mm.

DISTRIBUTION.—Anthrax irroratus striatipennis

occurs in Florida and the coastal plain of southeastern

United States as far west as Alabama and as far north

as South Carolina. It is allopatric to irroratus irroratus

(Map 1).

HOLOTYPE.—cf, Billy's Island, Okefenokee Swamp,

Georgia, VI-1912 (Cornell U. Lot. 482, Sub. 148).

ALLOTYPE.—Same data as holotype (in copula).

PARATYPES.—Alabama: Lee Co., cf, $5 Auburn,

VII-9-1917 (J. Bequaert).

Florida: Alachua Co., d, Gainesville, VI-4-1958

(J. W. Perry) (in light trap) ; ° , VI-19-1961 (H. A.

Denmark) (in black light trap). Brevard Co., $ ,

Titusville, IV-25-1923 (F. 4676). Duval Co., <f,

Jacksonville (Mrs. A. T. Slosson) ; $ , Jacksonville,

VIII-24-1958 (Dave Ribble). Highlands Co., <?, ? ,

Highlands Hammock St. Pk., 111-31-1956 (H. V.

Weems, Jr.); $ , Archbold B. Sta., Lake Placid,
14

111-28-1957 (H. E. Evans). Orange Co., <?, Gotha,

111-96 (W. M. Wheeler). Osceola Co., ? , Osceola

National Forest, VII-1-1961 (A. and H. Dietrich)

(ocean pond camp). Palm Beach Co., o*, Lake Worth

(Mrs. A. T. Slosson) (Ace. 26226). Santa Rosa Co.,

? , VIII-15-1956 (F. W. Mead). Suwannee Co.,

2 ? , Branford, VII-31-1930 (R. H. Beamer). ?,

"W. Park," IV-14-1939.

Georgia: Decatur Co., d, Spring Creek, VII-16-

29-1912 (Cornell U. Lot 482, Sub. 148). Tift Co.,

? , Tifton, VI-13-1896 (C. W. Johnson). Screven

Co., $, VII-12-1946 (A. K. Wyatt). 2? , Billy's

Island, Okefenokee Swamp, VI-1912 (Cornell U. Lot

482, Sub. 148). 2c?, "Georgia" (Osten Sacken).

South Carolina: Aiken Co., d, Aiken, VI-12-1957

(J. R. Vockeroth).

OTHER MATERIAL EXAMINED.—Florida: Pinellas Co.,

d, Gulfport, III-3-1925 (F. M. Gaige). Orange Co.,

? , Gotha, III-1896 (W. M. Wheeler).

Georgia: Bibb Co., d, Macon, VI-3-1923 (F. W.

Walker). ?, "Prattsburg," VII-25-1930 (L. D.

Tuthill); d, "Georgia" (Osten Sacken).

TYPES.—The holotype and allotype are deposited in

the Cornell University collection. Paratypes may be

found in the collections of the author, R. H. Painter,

Cornell University, the Museum of Comparative

Zoology, the Canadian National Collection, the American Museum of Natural History, the United States

National Museum, the University of Kansas, the University of Michigan, and the Florida State Plant Board.

DISCUSSION.—Anthrax irroratus striatipennis usually may be separated from irroratus irroratus by the

dense pattern of transverse striae on the wing. Specimens of irroratus usually have the wing pattern less

dense with most of the independent spots rounded. In

some specimens of irroratus from eastern United

States the spots may be elongated, in which case the

absence of white scales on the anterior parts of

abdominal sterna two through four and the larger

size of striatipennis will serve to separate the subspecies.

The differences between irroratus and other species are

noted under the nominate subspecies.

Anthrax cintalpa Cole

Argyramoeba oedipus.—Osten Sacken, 1877, p. 243 [part].—

Osten Sacken, 1878, p. 90.—Osten Sacken, 1886, p. 102

[part].—Coquillett, 1894, p. 95 [part].—Kerte"sz, 1909,

p. 66 [part] [not Fabricius, 1805; misidentification].

Anthrax oedipus.—Edwards, 1930, p. 173 [part] [not Fabricius, 1805; misidentification].

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Anthrax cintalpa Cole, 1957, p. 200.—Painter and Painter,

1965, p. 431.

MALE.—Head with fine black setae and black and

white lanceolate scales, sometimes with a few brown

scales. Basal antennal segments brown to red orange;

second segment short, saucer shaped, apical margin

produced as sharp flange (Figure 14; Plate 1/).

Mesonotum and scutellum with linear or lanceolate,

black and white scales; often with some brown scales,

especially in anterior patch on disc of scutellum. Scales

on mesopleuron, sternopleuron, and anterior half of

pteropleuron black, brown, white, or mixed. Pile on

prosternum, propleuron, and anterior margin of

mesonotum black, white, or mixed, often some brown

hairs present. Postalar tuft of pile black, often with

some white hairs and rarely entirely white. Scales on

coxae white, or mixed black and white.

Cells C and Sc of wing (Plate \e), with alternating

pigmented and hyaline areas. Remainder of wings with

numerous irregular dark brown spots, sometimes coalesced into preapical, submedial, and basal transverse

bands, sometimes largely discrete. Discrete spots mostly

rounded, few elongated transversely. Basal part of vein

Cui one-fifth to one-third as long as entire vein. Spur

at basal angle of vein R4 very rarely connected with

vein R2+3 forming a sectoral crossvein.

Femora usually black, occasionally reddish brown;

tibiae black to reddish yellow. Scales on femora lanceolate and ovate-truncate, usually black anteriorly and

white posteriorly. Middle femur without a bristle anteriorly above anteroventral row (Figure 36).

Lateral margins of first abdominal tergum with

white pile and few black setae; second tergum with

black pile, setae, and erect, lanceolate- and/or

oblanceolate-truncate scales laterally; lateral margins

of remaining terga with black setae and scales

anteriorly, and white or golden-brown scales posteriorly, especially on posterior terga. Posterior margin

of first tergum with white scales laterally and black

scales medially; discs of remaining terga with linear

black scales anteriorly, sometimes with a few goldenbrown or white scales; posterior margins with obovatetruncate white scales in submedial and sublateral

patches, patches sometimes coalesced on two and five,

sometimes reduced or absent on four, and expanded

over six and seven. Venter of abdomen with sparse

black setae and linear scales, predominantly black in

specimens from southern Mexico, predominantly white

in specimens from United States.

FEMALE.—Similar to male.
NUMBER 43 15

MALE GENITALIA (Figure 8).—Apical part of gonocoxites about three times longer than basal part; distal

segment of gonostylus about three times as long as

wide. Apex of epiphallus with sharp dorsal flange extending ventrally as sharp carina, a serrate process

present above apex of aedeagus. Apex of aedeagus not

flared, gonopore ventral. Epandrium rounded apically

in dorsal view; cercus projecting apically beyond

epandrium.

FEMALE GENITALIA (Figure 47).—Tenth tergum

with nine spines apically on each side; ventral extensions at base broadened at right angle apically. Apical

part of sclerite on each side of gonopore twisted, lateral

lobe arising mesally, narrow, acute apically; ventral

extension narrow, slightly acuminate and curved

mesally. Ducts of spermathecae united into medial

tube before emptying into genital chamber; ducts with

six or seven convolutions before expanding to bulbs;

bulbs S-shaped basally, widest postmedially, sharply

constricted to nipple-like apices.

BODY LENGTH.—6.2 to 10.6 mm.

WING LENGTH.—6.2 to 10.8 mm.

DISTRIBUTION.—Anthrax cintalpa occurs in forested

areas from southwestern United States through Mexico and Central America into Venezuela (Map 2).

The distribution probably is limited on the south by

the Amazonian rain forest in Venezuela and by the

Andes Mountains in Colombia. In the United States

it is limited by the Great Plains and eastern deciduous

forests. The northern limit is approximately at the 41st

parallel except for a population in northwestern

Nebraska.

MAP 2.—Distribution of Anthrax cintalpa.

360-673 O—7C
16

TYPES.—Anthrax cintalpa was described from four

specimens collected at Francia, 8 mi. NE of Cintalpa,

Chiapas, Mexico, by R. C. Bechtel and E. I. Schlinger.

The types are in the University of California

(Berkeley) collection. The paratypes have been studied

by the author.

BIOLOGY.—Cole (1957) stated that the type specimens of A. cintalpa were reared from the nests of

Melitoma euglossoides Lepeletier and Serville (Hymenoptera: Apidae). Other specimens have been reared

from Megachile chilopsidis, M. concinna, and

Dianthidium sp. (Hymenoptera: Megachilidae). This

species has been collected in several localities in Mexico

by the author, always along steep embankments.

DISCUSSION.—Anthrax cintalpa differs from A. irroratus in having a saucer shaped rather than lenticular second antennal segment and in lacking oblanceolate and obovate scales on the sides of the second

abdominal tergum. Anthrax oedipus and peruvianus

differ from cintalpa by having a distinctive bristle

or bristles on the anterior side of the middle femur

above the anteroventral row; also, cintalpa usually

has the postalar tuft of pile black or mixed black and

white, whereas in Mexican and Central American

specimens of oedipus it is white.

Anthrax oedipus oedipus Fabricius

Anthrax oedipus Fabricius, 1805, p. 123.—Osten Sacken,

1858, p. 41 [part].—Curran, 1927, p. 85.—Edwards, 1930,

p. 173 [part].—Stuardo Ortiz, 1946, p. 93.—Wolcott, 1951,

p. 450.—Painter and Painter, 1962, p. 78.

Argyramoeba oedipus.—Lynch Arribalzaga, 1879?.—Wulp,

1882, p. 85.—Osten Sacken, 1886, p. 102 [part].— Williston, 1901, p. 275.—Kertesz, 1909, p. 66 [part].

Spongostylum oedipus.—Aldrich, 1905, p. 223 [parti

[Spogostylum.]

Anthrax irroratus.—Painter, 1930, p. 794.—Wolcott, 1951,

p. 450 [not Say, 1823].

Exoprosopa punctata Macquart, 1850, p. 108.

Anthrax punctum Walker, 1850, p. 253.

Anthrax aequa Walker, 1852, p. 192.

Argyramoeba aequa.—Kertesz, 1909, p. 59.

Argyramoeba caloptera Schiner, 1868, p. 121.—Kertdsz, 1909,

p. 62.

MALE.—Head with black setae and mixed black,

white and sometimes brown, lanceolate scales. Antennae black or with first two segments reddish; second

segment short, with apical edge produced as sharp

flange (Figure 16).

Mesonotum and scutellum with sparse black setae,

and black, white and sometimes brown, linear and

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lanceolate scales, curly along margins; scutellum usually

with a spot of brown scales medially on anterior side.

Mesopleuron, sternopleuron, and pteropleuron with

white or yellowish-white pile, black or mixed black

and yellow setae, and white or mixed brown and white

scales; scales on coxae white. Prosternum, propleuron,

and anterior margin of mesonotum with white or

yellowish-white pile, often with some black hairs;

postalar tuft of pile white, rarely with few black hairs.

Cells C and Sc of wing (Plate \g) with alternating

hyaline and pigmented areas; remainder of wing pigmented with brown spots, most dense and more or less

coalesced from the basal cells out of bases of cells R2+3,

R5, 2M2, and Cu1} and in band from apex of vein Sc to

apices of veins M2 and Cui; basal band usually broken

midway on vein Cu2, apical band usually broken on

m crossvein and often on vein M2; specimens from

Mexico with posterior spots much reduced. Spur at

basal angle of vein R4 not connected with vein R2+3 to

form a sectoral crossvein. Basal segment of vein Cuj

one-fourth to two-fifths as long as entire vein.

Femora with scales usually predominantly black

anteriorly and predominantly white posteriorly; middle femur with distinctive bristle or group of bristles on

anterior side about three-fifths of way to apex above

anteroventral row (Figure 37).

Pile on sides of first abdominal tergum white or

yellowish white, few black setae present posteriorly;

posterior margin with white scales laterally and black

scales medially. Sides of terga two through five with

dense black pile and linear black scales; sides of posterior terga with black setae and obovate-truncate

white scales. Discs of terga with black scales anteriorly

and white scales submedially and laterally on posterior

margins, expanded on third segment, reduced on

fourth, and expanding over most of terga six and seven;

few brown scales sometimes intermixed with white.

Sterna with sparse black setae and sparse linear scales,

predominantly white anteriorly and mixed black and

white posteriorly, sometimes with a few brown scales.

FEMALE.—Similar to male.

MALE GENITALIA (Figure 13).—Apical part of gonocoxites about 2.5 times longer than basal part; distal

segment of gonostylus about 3 times longer than wide.

Apex of epiphallus with sharp dorsal flange apically,

without medial process above aedeagus. Apex of aedeagus flared, gonopore posteroventral, entire laterally.

Epandrium rounded apically in dorsal view; cercus

projecting apically beyond epandrium.
NUMBER 4 3

FEMALE GENITALIA (Figure 49).—Tenth tergum

with 10 spines apically on each side; ventral extensions

at base slightly expanded and bluntly rounded apically.

Dorsal part of sclerite on each side of gonopore bluntly

rounded and slightly produced apically toward meson;

ventral extension narrow and parallel sided, slightly

curved mesally. Ducts of spermathecae uniting into

common tube before emptying into genital chamber;

ducts with two convolutions before expanding to bulbs;

neck of bulbs recurved, remainder straight, broadest

medially and slightly constricted before lightly sclerotized apex.

WING LENGTH.—5.8 to 9.5 mm.

BODY LENGTH.—5.8 to 10.4 mm.

DISTRIBUTION.—Anthrax oedipus oedipus occurs

throughout South America except in the Amazon

Basin and the Andes Mountains, and in southern

Brazil where it is replaced by oedipus aquilus. It probably occurs throughout the West Indies, Central America, and southern Mexico, ranging northward along

the Pacific coast to Sinaloa and along the Gulf coast

into south-central Texas (Map 3).

TYPES.—The type-specimens (two males) of A.

oedipus Fabricius are in the Copenhagen Museum.

Painter and Painter (1962) state that one male is preserved in excellent condition; the other is headless and

badly rubbed. The better preserved specimen was designated lectotype. The photograph and redescription

of the lectotype leave little doubt as to the identity

of this species. The type-locality is the West Indies

where only oedipus oedipus occurs of the species in

the oedipus subgroup.

Painter and Painter (unpublished notes) state that

there are four specimens listed as Exoprosopa punctata

Macquart (1849) in the Bigot collection in the British

Museum. The first carries the label "Exoprosopa punctata Macq., 4 supp." (Macquart's label); the second

carries the label "A. punctata. Exoprosopa id. Macq.,

Ost. Sacken., Amer. Merid."; the third specimen is

labeled "B. Ayres"; and the fourth has no label. Painter

and Painter considered the second, if any, specimen to

be the type. It is badly covered with fungus and lacks

the right wing, but their notes and wing photograph

indicate that it is a typical specimen of oedipus oedipus.

The type-locality is "Amer. Mer."

According to Painter and Painter (unpublished

notes), the type of Anthrax aequa Walker (1852) is

in the British Museum. It carries the labels "One of

Walker's types so named. E. A. W.," "Brasil, Mrs. Noel"

and the green type label (without a name on the back).

oedipus oedipus Fabricius mi

oedipus aquilus, n. ssp. •»:•>

MAP 3.—Distribution of Anthrax oedipus oedipus and A.

oedipus aquilus.

The type is now headless, lacks all but one leg, has the

vesture rubbed off one side of the abdomen, has the

left wing glued on, and has the body and wings dirty.

The description and photographs of the type made by

Painter and Painter indicate that this species is a

synonym of oedipus oedipus.

The type of Argyramoeba caloptera Schiner (1868)

was found and marked by Painter and Painter in the

Vienna Museum. According to their notes, it is a male,

quite well preserved except that its head is covered

with fungus obscuring the antennae, and only one middle leg and one foreleg remain. The type carries the

labels "Lindig 1864 Venezuela," "Calopterus alte

sammlung" and "Argyramoeba calopterus Schin." The

photographs and redescription of the type indicate

that the species is a synonym of oedipus oedipus.

The type of Anthrax punctum Walker (1849) was

not found in any European museum visited by Painter

and Painter. The description leaves little doubt, how-
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ever, that the species is a synonym of oedipus oedipus.

The type-locality is "Brasil."

BIOLOGY.—The only reared specimens in collections

seen by the author emerged from cells of Odynerus

guadelupensis Sauss. at St. Vincent, B.W.I. Specimens

of this species have been collected by the author in

Parana and Rio Grande do Norte, Brazil, and Guerrero, Tamaulipas, and San Luis Potosi, Mexico, along

paths through forested areas and around stumps and

sandy paths in drier areas. Two specimens were observed ovipositing on a stump at Sao Martinho,

Parana, Brazil.

DISCUSSION.—Anthrax oedipus may be distinguished from all other species in the oedipus subgroup

except peruvianus by the presence of a postmedial bristle on the anterior side of the middle femur above the

anteroventral row. Characters separating oedipus oedipus from oedipus aquilus and peruvianus are discussed

under those taxa.

Anthrax oedipus aquilus, new subspecies

Argyramoeba oedipus.—Kertlsz, 1909, p. 66 (part).

Anthrax oedipus.—Edwards, 1930, p. 173 (part).

MALE.—Scales on head black and white. Basal antennal segments black; second segment saucer shaped,

apical margin produced as sharp flange.

Mesonotum and scutellum with black setae and

black and white scales; scutellum with golden-brown

scales in large medial patch on anterior half. Mesopleuron, pteropleuron, and stemopleuron with black

setae and mixed black and white scales, some black

and white pile on mesopleuron. Pile on prosternum,

propleuron, and anterior margin of mesonotum mixed

black and white; postalar tuft of pile black. Coxae with

black setae and white scales.

Cells C and Sc of wing (Plate \h) with alternating

pigmented and hyaline areas. Remainder of wings with

numerous irregular dark brown spots, mostly coalesced

into preapical, submedial, and basal transverse bands.

Submedial and basal bands not distinctly separated;

preapical band extending unbroken from tip of vein

Sc to tip of vein Cux; submedial band continuous in

cell Cu! on basal half of vein Cu2. Spur at basal angle

of cell R4 connected with vein R2+3 to form a complete sectoral crossvein. Basal part of vein Cuj about

one-fourth as long as entire vein.

Femora black, tibiae reddish brown. Scales on fore

and middle femora mixed black and white posteriorly,

black anteriorly; scales on hind femur entirely black.

Middle femur with distinctive bristle three-fifths of

way to apex on anterior side in addition to anteroventral row.

First abdominal tergum with white pile and few

black setae laterally; sides of second tergum with black

setae and dense black pile and scales; third, fourth,

and fifth terga with black setae, hairs, and scales

laterally, some obovate-truncate white scales posteriorly on two and four; sixth and seventh terga with white

scales laterally. Posterior margin of first tergum with

white scales laterally, and black scales medially and

among white scales; discs of remaining terga with

black scales anteriorly and white scales along posterior

margins in submedial and sublateral patches, white

scales united medially on five and spreading over discs

on six and seven. Venter with black setae, and linear

and lanceolate scales, white on first sternum, anteriorly

on second sternum and laterally on posterior margins

of second and following sterna, black elsewhere.

FEMALE.—Similar to male. Postalar tuft of pile with

few golden-brown and white hairs.

VARIATION.—The basal antennal segments vary

from black to reddish brown. The spur at the basal

angle of cell R4 may not be connected to vein R2+3 in

one wing or rarely in both wings; the subapical band

of spots may be broken on the m crossvein; the submedial band is rarely broken on vein Cu2. The femora

may be reddish brown or black, the tibiae vary from

reddish brown to yellow. White scales on venter of

abdomen may be slightly more extensive than in the

type, but usually are not more numerous than the black

scales on segments three through seven.

MALE GENITALIA.—Similar to oedipus oedipus.

Distal lobe of gonostylus sometimes less than three

times as long as wide.

FEMALE GENITALIA.—Similar to oedipus oedipus.

BODY LENGTH.—6.3 to 11.0 mm.

WING LENGTH.—6.7 to 9.7 mm.

DISTRIBUTION.—Anthrax oedipus aquilus occurs in

the plateau of southern Brazil and adjacent areas as

far north as southern Minas Gerais and west into Paraguay (Map 3). A specimen from Buenos Aires may

indicate that this subspecies has a more extensive southern distribution than shown on the map. Specimens

collected by the author were found in rather dense

forested areas.

HOLOTYPE.—0% Jacarezinho, Parana, Brazil, II14-1961 (N. Marston-9).

ALLOTYPE.— $ , Bage, Rio Grande do Sul, Brazil,

III-1-1961 (N.Marston-1).
NUMBER 4 3 19

PARATYPES.—Argentina: Missiones, Parambi, I-II1950(J.H.Jibson).

Brazil: Santa Catarina, $, Nova Teutonia, II—2—

1938 (F. Plaumann); $, Nova Teutonia, XII-1955

(F. Plaumann); ? , Nova Teutonia, 1-14-1956 (F.

Plaumann); $, Nova Teutonia, X-1956 (F. Plaumann) ; 2 ? , X-31-XII-7-1956 (F. Plaumann); 2 ? ,

Nova Teutonia, 1-27-30-1957 (F. Plaumann); $,

Nova Teutonia, IX-22-1957 (F. Plaumann); 2? ,

Nova Teutonia, 1-21-11-2-1959 (F. Plaumann); ?,

Nova Teutonia, XII-4-1959 (F. Plaumann). Sao

Paulo, $, Barueri, IV-19-1955 (K. Lenko); d, 2 ? ,

Barueri, III-31-V-24-1957 (K. Lenko); ? , Embu,

IX-1949 (F. Lane) ; c?, 3 ?, Faz. do Bonito, Serra da

Bocaina, S. Jose do Barreiro, I-1-II-10-1960 (Vulcano); d, Itii, 111-22-1961 (N. Marston-6); d,

Osasco, IX-1951 (M. A. V. A.); <?, Sao Paulo, Cid.

Jardim, 1-1945 (Barretto); 4c?, ? , Sao Paulo, Ypiranga, IV-2-17-1936 (Lange de Morretas); d, Sao

Paulo, Ypiranga, 1-1940 (Carrera) ; d, "Sao Paulo"

(Barbiellini); <?, Ste. Andre, V-1937 (R. Spitz).

Forty-six additional specimens have been examined

from these localities:

Argentina: Buenos Aires, Buenos Aires.

Brazil: Cidade Azul, Minas Gerais; Curitiba, Parana; Cauna and Nova Teutonia, Santa Catarina;

Barueri, Campos do Jordao, Cantaricira, Guaruja,

Monte Alegres, Piedade and Sao Paulo, Sao Paulo.

Paraguay: Col. Independencia and Molinesque.

TYPES.—The holotype and allotype are in the U.S.

National Museum collection. Paratypes are deposited

in the collections of the author, R. H. Painter, the

University of California, the Canadian National Collection, the American Museum of Natural History, the

Departmento de Zoologia, Secretaria de Agricultura,

Sao Paulo, Brazil, the Instituto Oswaldo Cruz, Rio de

Janeiro, Brazil, and the Instituto Miguel Lillo, Universidad de Tucuman, Tucuman, Argentina.

DISCUSSION.—The margin between the ranges of

A. oedipus aquilus and oedipus oedipus seems poorly

defined. Intermediates have been studied from these

localities: Jacarezinho, Parana; Nova Teutonia, Santa

Catarina; Jatai, Goias; Porto Cabral, Sao Paulo; and

Caceres, Bodoquena, and Corumba, Mato Grosso.

Since it appears that oedipus is adapted to arid areas,

whereas aquilus occurs in areas with denser vegetation

and higher rainfall, the broad range of intermediates

could be accounted for by recent restriction of the

range of aquilus and expansion of the range of oedipus

due to progressively drier climate in south-central

Brazil and Paraguay.

Anthrax oedipus aquilus differs from the nominate

subspecies in having a complete sectoral crossvein and

in having the postalar tuft of pile black. There are also

several other less reliable differences between the subspecies. The wings usually are more heavily infuscated

in aquilus than in oedipus with the submedial band of

spots continuous to the apex of vein 2A in the former,

whereas it usually is broken on vein Cu2 in the latter.

Also, aquilus generally has fewer white scales, especially ventrally on the posterior femur, ventrally on

the abdomen and posteriorly on the mesonotum.

Differences between A. oedipus and other species in

the oedipus subgroup are discussed under the nominate

subspecies.

Anthrax peruvianus, new species

MALE.—Head with black setae, and black, white,

and yellow scales. Fringe of pile on posterior margin

of occiput black with light tips. Antennae black; second segment short, with apical margin produced as

sharp flange.

Disc of mesonotum with yellow, white, and black

scales; scutellum with lanceolate golden-brown scales

medially. Scales on mesopleuron, pteropleuron, and

sternopleuron white. Pile on prosternum white, on

propleuron and anterior margin of mesonotum white

with some black hairs and setae intermixed; postalar

tuft of pile white. Scales on coxae white.

Cell C of wing (Plate 1/) with alternating hyaline

and pigmented areas; cell Sc with alternating pigmented and subhyaline areas. Remainder of wings pigmented with brown spots, most dense and mostly coalesced in medial and preapical bands. Spur at basal

angle of cell R4 not connected with vein R2+3 to form

a sectoral crossvein. Basal section of vein Cux about

one-fourth as long as entire vein.

Femora with white and yellowish-white scales, a

few brown scales on anterior surfaces of fore and middle pairs; middle femur with distinctive bristle anteriorly above anteroventral row.

Sides of first abdominal tergum with dense white

pile, with few black setae posteriorly; posterior margin

with yellowish-white scales. Sides of second tergum

with black pile and lanceolate scales anteriorly, and

with some yellow hairs and lanceolate scales posteriorly; sides of third through sixth terga with black hairs

and scales anteriorly and yellowish-white hairs and
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

scales posteriorly; sides of seventh tergum with white

scales. Discs of second and following terga with linear

black and brown scales anteriorly; posterior margins

with white scales submedially and laterally, black scales

decreasing and white scales increasing in number posteriorly. Sterna with yellow setae and rather dense

linear and lanceolate white scales.

FEMALE.—Fore and middle femora with patches of

brown scales anteriorly.

VARIATION.—The pile on the thorax and sides of

the first abdominal tergum varies from white to yellowish white with some brown hairs sometimes present.

The lateral margins of terga two and four may or may

not have tufts of white scales posteriorly. The scales on

the anterior surfaces of the femora may be predominantly yellowish white or brown. The pigmented areas

in the cell Sc of the wing may be well or poorly defined.

The spots on the wing vary from light brown to dark

brown; they may or may not be coalesced into medial

and preapical bands. The spur at the basal angle of

vein R4 may connect to vein R2+3 to form a sectoral

crossvein.

MALE GENITALIA (Figure 10).—Apical part of gonocoxites about twice as long as basal part; distal segment of gonostylus about 1.5 times longer than wide.

Apex of epiphallus with a sharp dorsal flange, without

medial process above aedeagus. Apex of aedeagus

flared, with distinct lateral and ventral emarginations.

Epandrium rounded apically in dorsal view; cercus

projecting apically beyond epandrium.

FEMALE GENITALIA (Figure 46).—Tenth tergum

with 12 spines apically on each side; ventral extensions

at base sharply rounded apically. Dorsal part of sclerite

on each side of gonopore broad laterally with sharp

dorsal angle and rounded ventral angle, produced as

narrow dorsal lobe toward meson; ventral extension

narrow, parallel sided and slightly curved toward

meson. Ducts of spermathecae united before emptying

into genital chamber, with two convolutions before

expanding to bulbs; neck of bulbs recurved, remainder

broadest medially and slightly constricted before lightly

sclerotized apex.

BODY LENGTH.—8.1 to 9.6 mm.

WING LENGTH.—7.6 to 9.2 mm.

DISTRIBUTION.—Anthrax peruvianus is known from

only a few localities in Peru (Map 4).

HOLOTYPE.— <S, Pachacamac, Lima, Peru, IX-1 -

1964 (R. H. Painter).

ALLOTYPE.— 9 , same data as holotype.

PARATYPES—Peru: Arequipa, cf, IV-10-1938.

peruvianus, n. sp.

cordillerensi*. n. sp.

MAP 4.—Distribution of Anthrax peruvianus and A. cordillerensis.

Guayas, cf, 8 mi. S Manglar Alte, 1-31-1955 (E. I.

Schlinger and E. S. Ross). Huanuco, $ , IV-6-1940,

1900 m (Weyrauch) (W. K. W., 3480-C). Lambayeque, $ , Chiclayo, XII-12-1930; $ , 42 mi. E Olmos,

1-18-1955 (E. I. Schlinger and E. S. Ross). Lima,

$ , Chaclacayo, 750 m (E. Escomel); 2 cf, $ , Chosica,
NUMBER 43 21

XI-5, XII-13-14-1964 (P. Hocking); 3? , Lima,

1-1-1939 (Weyrauch) (W. K. W., 3480); ? , Lima

(Soukeep); lld \ ? , Matucana, X-31-1964, 8000

feet (R. H. Painter); 2cf, "W. K. W., 3480." Piura,

? , Piura. $ , "La Areha," VII-14-1944 (Berry).

TYPES.—The holotype and allotype are in the collection of R. H. Painter. Paratypes may be found in

the collections of the author, Prof. Wolfgang Weyrauch, R. H. Painter, the American Museum of

Natural History, the Museum of Comparative Zoology, the California Academy of Sciences and the U.S.

National Museum.

DISCUSSION.—Anthrax peruvianus is most closely

related to oedipus. The male genitalia are quite similar,

but peruvianus has the apex of the aedeagus shallowly

three-lobed while in oedipus it is entire and rounded.

Externally, peruvianus has patches of golden-brown

scales on the abdominal terga and has few, if any, black

scales on the third, fourth, and fifth sterna. It usually

has a tuft of brown and white scales posteriorly on the

lateral margins of the second tergum. Anthrax oedipus is usually darker, lacking large patches of brown

scales on the abdominal terga and having numerous

black scales on the third, fourth, and fifth sterna. There

are no brown or white scales posteriorly on the lateral

margins of the second tergum.

Anthrax peruvianus sometimes has pigmented areas

in the cell Sc poorly defined and separated by subhyaline areas, although it does not appear to be closely

related to the Chilean species, so it may be keyed

through both choices of couplet 2. It may be readily

separated from all species, except oedipus, in the

oedipus subgroup, by the bristle or bristles anteriorly

above the anteroventral row on the middle femur.

Anthrax cordillerensis, new species

Anthrax oedipus.—Edwards, 1930, p. 173 [part] [not Fabricius, 1805].

MALE.—Scales on head black, and white or yellowish white. Basal antennal segments black; second segment saucer shaped, apical margin produced as sharp

flange.

Scales on mesonotum black and white, few goldenbrown ones intermixed. Scutellum with white and

black scales, few golden-brown scales present around

white patches. Pile on dorsal half of mesopleuron and

anterodorsal part of pteropleuron white, few goldenbrown hairs present. Scales on lower half of mesopleuron, anteroventral part of pteropleuron, and

sternopleuron mostly white, few black and goldenbrown scales present. Pile on prosternum, propleuron,

and anterior margin of mesonotum mixed black and

white, few brown hairs present on the propleuron.

Postalar tuft of pile black. Scales on coxae brown and

white.

Cell C of wing (Plate 1;) with alternating pigmented and hyaline areas; cell Sc mostly subhyaline, without well-defined pigmented areas. Remainder of wing

with numerous rounded brown spots not coalesced into

transverse bands. Sectoral crossvein incomplete. Basal

part of vein Cux two-fifths as long as entire vein.

Femora black, tibiae orange. Fore and middle femora

with black and white scales; hind femur with black

scales. Middle femur without bristle anteriorly above

anteroventral row.

Pile on sides of first abdominal tergum white, few

black hairs and setae present posteriorly. Sides of second tergum with black pile and erect, linear scales;

sides of terga three to six with black setae and oblanceolate and oblanceolate-truncate scales, black anteriorly, white posteriorly, white scales most extensive

on three; seventh tergum with oblanceolate-truncate

white scales. Discs of remaining terga with linear black

scales anteriorly and oblanceolate or oblanceolatetruncate white scales submedially and laterally along

margins, white scales expanding over discs on posterior

terga. Scales on venter long and white on first sternum

and lateral margins of posterior sterna, short and black

elsewhere.

FEMALE.—Similar to male. Basal antennal segments

orange. Posterior femur with white scales ventrally.

VARIATION.—There may be a few white scales

ventrally on the posterior femora. The basal antennal

segments may be red orange or black. Cell C of the

wing may be entirely infuscated or subhyaline. The

spots on the wing may be fewer in some specimens,

and the basal and anterior part of the wing may be

more heavily infuscated than in the types.

MALE GENITALIA (Figure 12).—Apical part of

gonocoxites about 2.75 times longer than basal part;

distal segment of gonostylus about twice as long as

wide. Dorsal part of apex of epiphallus atrophied,

remainder trilobed apically; medial carina arched dorsally, lateral carina produced basad of medial carina.

Epandrium acuminate apically in dorsal view, extending past cercus.

FEMALE GENITALIA (Figure 43).—Tenth tergum

with nine spines apically on each side; ventral extensions from base broadened and rounded apically.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sclerite on each side of gonopore rounded dorsally,

produced above lateral lobe; lateral lobe not tapering,

bluntly rounded distally; ventral extensions tapering,

sharply curved mesad distally. Ducts of spermathecae

emptying together into genital chamber, with two convolutions before expanding to bulbs; neck of bulbs

curved at right angle, remainder straight, broadest

medially, and tapering to each end.

BODY LENGTH.—7.2 to 11.7 mm.

WING LENGTH.—6.9 to 10.5 mm.

DISTRIBUTION.—Apparently A. cordillerensis inhabits a rather small area along the east and west

sides of the Andes Mountains in central Chile and

western Argentina (Map 4).

HOLOTYPE.—cf, Las Cruces, Cord. Parral, Linares,

Chile, XI-11/25-1960 (Luis Pena).

ALLOTYPE.— $ , same data as holotype.

PARATYPES.—Argentina: Rio Negro, °.,ElBols6n,

1-1955.

Chile: Bio Bio, d, ?, El Abanico, XII-31-1950

(Ross and Michelbacher). Conception, d, Conception, XII-1926 (R. and E. Shannon). Coquimbo,

d, 5 mi. N Illapel, XI-30-1950 (Ross and Michelbacher) ; 2cT, 5 mi. N Laguna Dam, XII-6-1950

(Ross and Michelbacher); ? , 20 mi. SW of Ovalle,

XII-12-1950 (Ross and Michelbacher); ? , 12 mi.

E Rivadavia, XII-4-1950 (Ross and Michelbacher).

Linares, d, Fundo Malcho, XI-1956 (Luis Pena);

5o", Fundo Malcho, 1-1957 (Luis Pena); d, Fundo

Malcho, XII-1957 (Luis Pena); d, Fundo Malcho,

1-1958 (Luis Pena); d, Fundo Malcho, 11-1958

(Luis Pena); 2d, Las Cruces, Cord. Parral, XI11/25-1960 (Luis Pena). Malleco, d, Angol, XII-4-

1945; d, Angol, XII-4-1946; 3d, Angol, XI-6-1947

(D. S. Bullock); 2d\ Angol, XII-8-1956; d, Angol,

XI-15-1957 (W. Salinas). Maule, d, Constitucion

(P. Garulla; 2d, $, "Constitucion." Nuble, 2c?, 50

km E of San Carlos, XII-28-1950 (Ross and Michelbacher). Santiago, $, Las Mercedes, 11-1937

(P. Flamino Ruiz); d, $, Qta. Normal, Santiago,

XII-11-1950 (E. Zomasa); ?, Santiago, "06"; ? ,

Pichinahuel, Cord. Nahuelabuta, 1-10/20-1959 (Luis

Pena). d, "Pua," II-4-1958 (G. Dorman). d,

"Chile," 1-12-1959 (T. Schmidt and D. Lepin).

Additional specimens have been seen from the following localities in Chile: Fundo Malcho, Linares

Prov.; El Peumo, O'Higgins Prov.; Laja, Nuble Prov.;

"Estero Leira"; "C. Chile" and "Chile."

TYPES.—The holotype and allotype are in the collection of the U.S. National Museum. Paratypes may

be found in the collections of the author, the Museum

of Comparative Zoology, the Instituto Oswaldo Cruz

in Rio de Janeiro, the University of California, Michigan State University, the U.S. National Museum, the

California Academy of Sciences, Jack C. Hall, and

R. H. Painter.

DISCUSSION.—Anthrax cordillerensis differs from

inordinatus in lacking a complete sectoral crossvein,

in having the femora black rather than orange, and

in lacking patches of golden-brown scales on the

abdomen. It may be distinguished from peruvianus by

the lack of bristles on the anterior side of the middle

femur above the anteroventral row and by the lack of

white scales on the posterior side of the posterior

femur. It differs from other species in the oedipus subgroup in lacking well-defined spots in the subcostal

cell of the wing.

A few specimens of cordillerensis have the cell C

mostly or entirely subhyaline as in the pluto subgroup.

These specimens usually also have the wings somewhat

darker than usual and have cell 2M entirely infuscated.

Anthrax inordinatus Rondani

Anthrax vicinus Blanchard, 1852, p. 381.—K6rtesz, 1909, p.

57.—Philipi, 1865, p. 663 [preoccupied Marquart, 1846].

Anthrax inordinatus Rondani, 1863, p. 64.—Kgrtesz, 1909,

p. 42.

Argyramoeba poecilophora Schiner, 1868, p. 121.—Ke'rtesz,

1909, p. 66.

Anthrax oedipus.—Edwards, 1930, p. 173 [part] [not Fabricius, 1805].

MALE.—Scales on head mostly white to brown, some

black scales present. Basal antennal segments yellow;

second segment saucer shaped, with the apical margin

produced as a sharp flange.

Discs of mesonotum and scutellum with black,

brown, and white scales. Scales on mesopleuron, pteropleuron, and sternopleuron golden brown; pile on

mesopleuron brown and white. Prosternum with mixed

brown and white pile; pile on propleuron and anterior

margin of mesonotum brown and white. Postalar tuft

of pile black with some brown or white scales basally.

Scales on coxae white or mixed brown and white.

Cell C of wing (Plate li) with alternating brown

and hyaline areas; cell Sc subhyaline, without distinct

spots. Remainder of wings with large, rounded spots,

more or less coalesced into preapical, medial and basal

bands; few independent spots. Spur at basal angle of

cell R4 usually connected to vein R2+3 to form a sectoral
NUMBER 4 3 23

crossvein. Bases of cells narrowly margined with subhyaline. Basal part of vein Cux one-fourth to one-third

length of vein.

Femora, tibiae, and basal tarsal segments orange.

Femora with black, brown, and white scales; middle

femur rarely with bristle on anterior side above

anteroventral row.

Sides of first abdominal tergum with dense, white or

yellowish-white pile and few black setae; posterior

margin fringed with golden-brown or mixed goldenbrown and white scales. Sides of posterior terga with

black setae and black scales anteriorly, with white

scales posteriorly, black scales less extensive on posterior

terga. Discs of second and following terga with bands of

golden-brown and black scales; posterior margins with

submedial and lateral patches of white and goldenbrown scales, expanding on posterior terga. Venter of

abdomen with golden-brown and white scales, sometimes with black scales medially on posterior sterna.

FEMALE.—Similar to male.

MALE GENITALIA (Figure 11).—Apical part of gonocoxites about 2.5 times longer than basal part; distal

segment of gonostylus about as long as wide. Apex of

epiphallus atrophied; aedeagus trilobed apically;

medial carina flattened dorsally, lateral carina not

produced basad of medial carina. Epandrium acuminate apically in dorsal view, extending past the cercus.

FEMALE GENITALIA (Figure 48)—Tenth tergum

with seven spines apically on each side; ventral extensions from base tapering apically. Apex of sclerite on

each side of gonopore broadly produced above lateral

lobe, tapering to broadly rounded apex; lateral lobe

small, truncate apically; ventral extension broad on

upper two-thirds, tapering and strongly curving mesad

on lower third. Spermathecal ducts emptying together

into genital chamber; ducts with two convolutions before expanding the bulbs; neck of bulbs curved; remainder slightly sinuous, narrow basally, widest postmedially, and narrowing abruptly to apex.

BODY LENGTH.—7.6 to 10.8 mm.

WING LENGTH.—7.4 to 9.7 mm.

DISTRIBUTION.—Anthrax inordinatus occurs on both

sides of the Andes Mountains in central Chile and

western Argentina (Map 5). Its distribution seems to

be the same as that of cordillerensis.

TYPES.—Painter and Painter (unpublished notes)

found the type male of Anthrax inordinatus Rondani

(1863) in the Museo dell' Institute di Zoologia dell'

Universita di Napoli in Naples, Italy. The type is

headless, has a thin film of fungus, and has one wing

MAP 5.—Distribution of Anthrax inordinatus Rondani.

broken, but is otherwise in good condition. It carries

the labels "M° . Zoo.° N°, 11125" and Anthrax inornatus (sic) Rond. Chili."

Painter and Painter found the male and female types

of Argyramoeba poecilophora Schiner in the Naturhistorisches Museum in Vienna, Austria. The male was

in good condition, although a little dusty, and was

designated the lectotype. The female had a Villa head

glued onto it. The lectotype carried the labels "Chile

Novara Reise" and "poecilophora Alta Sammlung,"

but no type label. Photographs of the lectotype show it

to be a typical specimen of inordinatus Rondani.

DISCUSSION.—Anthrax inordinatus differs from

cordillerensis in having a complete sectoral crossvein,

in having the femora orange instead of black, and in

having large patches of golden-brown scales on the

abdomen. Anthrax inordinatus is superficially quite

similar to peruvianus, although the latter is more closely

related structurally to oedipus. Anthrax peruvianus

may be distinguished readily by the presence of a distinctive bristle or bristles on the anterior surface of the

middle femur above the anteroventral row, and by the

absence of a complete sectoral crossvein. Anthrax inordinatus may be distinguished from the other members of the oedipus subgroup by the absence of welldefined pigmented areas in the cell Sc of the wing.

Anthrax pluto pluto Wiedemann

Anthrax pluto Wiedemann, 1828, p. 261.—Osten Sacken,

1858, p. 41 [part].—Painter and Painter, 1962, p. 80.—

Painter and Painter, 1965, p. 432.
24

Argyramoeba pluto.—Osten Sacken, 1877, p. 244 [partj.—

Osten Sacken, 1878, p. 90 [part].—Coquillett, 1894, p. 95

[part].— Kertesz, 1909, p. 66 [part].

Spongostylum pluto.—Aldrich, 1905, p. 223 [part]

[Spogostylum].

MALE.—Scales on head black and white. Basal

antennal segments black; second segment saucer

shaped, produced as a sharp flange apically.

Scales on mesonotum and scutellum mostly black;

some white scales present, especially along margins.

Pile on mesopleuron and pteropleuron mixed black,

white, and brown; scales on pleura black and white.

Prosternum, propleuron, and anterior margin of mesonotum with mixed black and white pile; sometimes

with a few brown hairs, especially on humeral calli;

postalar tuft of pile mixed black and white. Scales on

coxae black and white.

Cells C and Sc of wing (Plate 2e-f) entirely infuscated or with small subhyaline spots before and

after humeral cross vein in costal cell. Cells 1M, 2M,

and R entirely infuscated except for a subhyaline or

hyaline area submedially in cell R and sometimes a

subhyaline area medially or postmedially in cell 2M.

Bases of cells 1A and 2A completely infuscated or

with small subhyaline areas basally. Spots at bases of

cells R2+3, Rsj 2M2, and Cux broadly coalesced with

basal infuscated area; basal third of cell 1M2 infuscated. Discrete or coalesced spots present at bases of

cells Rj and Mi, at apices of veins R2+3, M2j Cul3 and

Cu2, at the medial angle of vein R4 and m crossvein,

subapically on vein R2+3, and below tip of vein Rx;

broad spot extending from vein Ri to vein Mi+2 halfway between bases of cells R2+3 and R4.

Fore and middle femora with black scales anteriorly

and white or mixed black and white scales posteriorly;

scales on hind femur black; a few white scales sometimes posteriorly.

Sides of first abdominal tergum with white pile;

some black hairs usually present posteriorly; posterior

margin with white scales laterally and black scales

medially. Lateral margins of terga two through five

with black pile, linear and lanceolate scales, and black

setae, few obovate white scales present posteriorly on

third; sixth and seventh terga with white scales present

submedially on posterior margins of two through five,

laterally on five and sublaterally on two and three;

terga six and seven with black scales medially and

white scales laterally. Sterna one, two, and three with

threadlike, mostly white scales; black scales sometimes

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

present on two and three; sterna four through seven

with black or mixed black and white scales.

FEMALE.—Similar to male.

MALE GENITALIA (Figure 34).—Apical part of gonocoxites narrow and slightly tapering in lateral view,

apex acute. Distal segment of gonostylus rectangular

basally with broad apical hook, ventral margin

slightly undulating. Apex of epiphallus curled upward

and outward dorsally and laterally, with short ventrolateral extensions; basal recurved process sharp, relatively large. Tip of aedeagus not flared, gonopore distoventral; base of aedeagus large, bulbous, tapering

gradually to junction with ventral bands; lateral and

medial apodemes large in relation to base of aedeagus.

FEMALE GENITALIA (Figure 41).—Tenth tergum

with 11 spines apically on each side; ventral extensions

at base broadly rounded apically. Apical part of sclerite on each side of apices of spermathecal ducts narrow and clavate laterally, sharply produced dorsally

toward the meson; ventral extension very long and

narrow, parallel sides and slightly curved mesad.

Ducts of spermathecae with one convolution before

expanding to bulbs; neck of bulbs broadly and evenly

recurved; remainder slightly curved and broadest

postmedially.

BODY LENGTH.—7.4 to 12.0 mm.

WING LENGTH.—6.7 to 11.0 mm.

DISTRIBUTION.—The small number of specimens

available of pluto pluto indicates that it occurs from

the Appalachian Mountains to the Great Plains as far

north as Michigan and probably as far south as Oklahoma and Tennessee (Map 6). Wiedemann designated "Kentucky" as the type-locality. Anthrax pluto

pluto is allopatric to pluto nigriventris on the south.

TYPES.—Painter and Painter (1962) stated that the

three females comprising the type series in the Vienna

Museum are well preserved. The largest female, carrying the labels "Pluto det. Wiedemann," "Col. Winthem" and "Pluto Wied. Kentucky," was designated

lectotype.

DISCUSSION.—Anthrax pluto pluto differs from

pluto nigriventris in having numerous white scales on

the abdominal sterna and in having the pile on the

lower half of the lateral margin of the first abdominal

tergum entirely white or with only a few black hairs

posteriorly. It may be distinguished from stellans by

the absence of golden-brown scales on the lower margin of the hind femur and the abdominal sterna.
NUMBER 43 25

pluto pluto Wiedeman

pluto nigriventri»

MAP 6.—Distribution of Anthrax pluto pluto and A. pluto

nigriventris.

Anthrax pluto nigriventris, new subspecies

Anthrax pluto.—Osten Sacken, 1858, p. 41 [part].—Brimley,

1938, p. 341.

Argyramoeba pluto.—Osten Sacken, 1877, p. 244 [part].—

Osten Sacken, 1878, p. 90 [part].—Wulp, 1881, p. 85.—

Coquillett, 1894, p. 95 [part].—Kertesz, 1909, p. 66 [part].

Spongostylum pluto.—Aldrich, 1905, p. 223 [part] [Spogostylurri].—Johnson, 1913, p. 55 [Spogostylum].

MALE.—Scales on head black and white. Antennae

black; second segment saucer shaped, produced apically as a sharp flange.

Scales on mesonotum mostly black; few white ones

on disc, and brown and white ones along margins.

Scales on scutellum black and white. Pile on mesopleuron and pteropleuron mixed black, brown, and

white. Scales on pleura sparse, black. Prosternum,

propleuron, and anterior margin of mesonotum with

mixed black, brown, and white pile, black predominating on former, white predominating on latter; postalar

tuft of pile mixed black and white. Scales on coxae

black and white.

Cells C and Sc of wing (Plate 2g) infuscated. Base

of wing out to r-m crossvein and base of cell Cux mostly

infuscated, hyaline and subhyaline areas present postmedially in cell 2M, submedially and subapically in

cell R, and basally in cell R2+3; submedial spot on vein

2A large, poorly defined from base; spot at base of

cell 2M2 narrowly connected to basal pigmented area.

Independent spots present at apices of vein R2+3, M2,

Cul5 and Cu2, in cell R below base of vein Ri, on vein

R2+3 below apex of vein R1} at medial angle and base of

vein R4; at base of cell Mx and at medial angle of m

crossvein; broad spot present midway between r-m

crossvein and base of Cell R* extending from vein Mi+2

to vein Ri.

Femora red orange apically, black basally; tibiae

red orange. Scales on femora and tibiae black.

Pile on sides of first abdominal tergum white dorsally, black and white ventrally; posterior margin with

few white scales laterally. Sides of terga two through

five with dense black pile and lanceolate scales; sides of

sixth tergum with black pile and black and white scales;

sides of seventh tergum with white scales. Discs of

terga two through six with black scales, small patches

of white scales present on posterior margins submedially on two through five, sublaterally on two, three, and

five, and laterally on six; seventh tergum with black

scales medially and white scales laterally. Scales on

venter mostly black, few threadlike white scales present

on first sternum.

FEMALE.—Similar to male. No brown hairs on anterior collar. Mesopleuron with numerous threadlike

white scales ventrally on mesopleuron. Cell 2M of

wing (Plate 2h) with medial longitudinal subhyaline

area only; subhyaline areas basally in cells 1A and 2A

reduced. Fore and middle femora with some white

scales posteroventrally.

VARIATION.—The scales on the mesopleuron and

pteropleuron vary from entirely white to entirely black;

a few white scales may occur on the sternopleuron. The

relative amounts of black, white, and brown pile on

the prosternum, propleuron, and the anterior margin

of the mesonotum vary and the brown pile may sometimes be absent. Cell 2M of the wings may be entirely

infuscated or have a small hyaline area postmedially;

cells 1A and 2A may or may not have subhyaline areas

at their bases. There may be a small subhyaline area in

the costal cell after the humeral crossvein. The size of

the spots and the degree of coalescense in the outer

part of the wing is variable, the females being darker.

The scales on the fore and middle femora may be entirely black or a few may be white. The ventral half of

the lateral margins of the first abdominal tergum may
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

be entirely black pilose or a few white hairs may be

intermixed. A few white scales may occur on the anterior sterna.

MALE GENITALIA (Figure 35).—Similar to pluto

pluto. Distal segment of gonostylus broader basally,

dorsobasal angle shorter and less acute. Apex of epiphallus produced apically above, ventrolateral parts

curving basally at a sharp angle; apex of aedeagus

curved downward, gonopore ventral.

FEMALE GENITALIA.—Similar to pluto pluto. Tenth

tergum with 10 spines on each side.

BODY LENGTH. —6.7 to 12.2 mm.

WING LENGTH.—6.3 to 11.3 mm.

DISTRIBUTION.—Anthrax pluto nigriventris has been

collected in south Florida, central Texas, and eastern

and south-central Mexico. It probably also occurs along

the Gulf coast between Texas and Florida and possibly

north along the Atlantic coast (Map 6).

HOLOTYPE.—cT, Brown wood, Brown Co., Texas,

VI-7 (R. H. Painter).

ALLOTYPE.— 9, Brownwood, Brown Co., Texas,

VI-6-1936 (R. H. Painter).

PARATYPES.—Texas: Brown Co., $, Brownwood,

IX-12-1922 (R. H. Painter); d, Brownwood, VIII11-1932 (R. H. Painter); o", Brownwood, VI-7

(R. H. Painter). Cameron Co., 9, Brownsville, June;

9, Brownsville, July; d, 8 mi. E Brownsville, VTI-18-

1962 (N. Marston-4). Colorado Co., d, VI-7-1922

(Mrs. Grace Wiley). Comal Co., 9, Spring Branch,

VII-4-1946 (S. Camras). Dallas Co., d, Dallas, VII30-1939 (R. H. Painter). Dimmit Co., d, Carrizo

Springs, VII-7-1938 (Jean Russell). Edwards Co.,

d, Rock Springs, VI-14-1962 (R. R. Grable). Hidalgo Co., ? , 25 mi. SW Harlingen, IV-10-1945

(D. E. Hardy and V. L. Wooley). Karnes Co., $,

Hobson, VIII-14-1931. Kleberg Co., d, Kingsville,

VI-7-1921; ? , Kingsville, VI-18-1921 (F. M. Hull);

d, Kingsville, VII-2-1921 (F. M. Hull); 9, Kingsville, VII-17-1921. McLennan Co., 9, Waco (Belfrage). Mason Co., d, Mason, VIII-24-1926 (R. H.

Painter). San Patricio Co., d, 9, FV-1-1938 (W.

Benedict). Travis Co., 9, Austin, VII-15-1923

(R. H. Painter); d, Austin, V-31-1958 (C. S. Lin);

d, summer, 1931 (J. K. G. Silvey). Uvalde Co., ? ,

Sabinal, V-1910 (F. C. Pratt); 9, Sabinal, VII-6-

1936 (M. B. Jackson); 9, Uvalde, VIII-28-1933

(A. W. Linquist). Val Verde Co., d, Del Rio, VII16/17-1946 (H.E.Evans).

Mexico: Guerrero, d, 33 mi. S Iguala, 1450 feet,

VII-5-1954 (J. G. Chillcott). Morelos, 9, 11 mi.

E Cuernavaca, Lobo Canyon, 3900 feet, VIII-15-1962

(R. H. and E. M. Painter). Tamaulipas, d, 15 mi.

S Matamoras, VI-6-1961 (Univ. Kans. Exped.).

Eleven additional specimens have been examined

from Brown, Cameron, Kleberg, Medina, Montague,

and Val Verde counties in Texas; Lee and Monroe

counties in Florida; and "Georgia" (Osten Sacken).

TYPES.—The holotype and allotype are in the collection of R. H. Painter. Paratypes may be found in

the collections of the author, R. H. Painter, Cornell

University, the University of California, the University

of Kansas, the University of Michigan, the U.S. National Museum, Iowa State University, the Florida

State Plant Board, and the Chicago Museum of Natural History.

DISCUSSION.—A specimen from Sanibel Island, Lee

County, and another from Key Largo Key, Monroe

County, Florida, are tentatively referred to this subspecies. They differ from typical forms in having the

infuscation of the wings greatly increased. Cells 1A

and 2A are almost completely filled and the spots in

the outer part of the wing are completely coalesced

except for those at the tips of veins R2+3, R4, Mi and

at the medial angle of vein R«. Otherwise the specimens agree with those from Texas.

One specimen from "Georgia" (Osten Sacken) is

intermediate between pluto pluto and pluto nigriventris. It has white scales on the posterior margins of

the fore and middle femora and has mixed black and

white scales on the abdominal sterna.

Anthrax pluto nigriventris differs from pluto pluto

in having the scales on abdominal sterna two through

seven entirely black or with only a few white ones

posterolaterally. It differs from both pluto pluto and

atriplex in having the pile on the lower half of the

sides of the first abdominal tergum black rather than

white.

Anthrax insulanus, new species

Spongostylum sp., nr. pluto.—Wolcott, 1951, p. 450 [Spogostylum]

MALE.—Scales on head black, white, and yellow.

Basal antennal segments black, second segment saucer

shaped, apical margin produced as sharp flange.

Scales on mesonotum mostly black; some white

scales present, dense along lateral margins. Scutellum

with black and white scales; pile on mesopleuron dense,

snowy white; bristles on pleura black and gold. Prosternum, propleuron, and anterior margin of mesono-
NUMBER 43 27

turn with mixed black and white pile, black predominating on prosternum; postalar tuft of pile mixed black

and white. Scales on coxae mixed black and white.

Cells C and Sc of wing (Plate 2t) evenly infuscated

with brown except for large hyaline spot after and

small one before humeral crossvein in cell C and hyaline spot after humeral crossvein in cell Sc. Remainder

of wing with brown, largely coalesced spots. Cell 1M

with small hyaline spot apically; cells R and 2M with

hyaline spots medially; submedial spot on vein 2A connected with basal infuscation in cell 1A, discrete in

cell 2A; spot at base of cell Cux broadly connected with

spots at bases of cells R1} R2+3, and R8; large elongate

spot extending from before apex of vein Sc in cell Ri

to vein M1+2, narrowly connected with spot at base of

cell R4; spots at base of cell Mi, apex of vein Cux and

apical angle of m crossvein narrowly connected; independent spots present apically on veins R2+3j R4, M2,

and Cu!, at base of cell 2M2, at medial angle of vein

R4, below tip of vein Ri in cell Rl3 and on vein R2+3

above base of cell R4.

Scales on femora mostly black; few white scales anteriorly on fore and middle femora.

Sides of first abdominal tergum with dense, snowwhite pile; posterior margin with white scales laterally

and black scales medially. Sides of terga two through

five with dense black pile, linear scales and setae, few

white scales posteriorly on third; terga six and seven

with white scales, some black pile and scales anteriorly

on sixth. Discs of terga two through seven with black

scales, posterior margins with white scales submedially

and sublaterally on two, sublaterally on three, submedially on four and five, and laterally on six and

seven; few white scales submedially on three, and sublaterally on four and five. Scales on venter black except

for few threadlike white scales on first sternum and

sides of second sternum.

FEMALE.—Similar to male. Wing (Plate 2;) with

spots somewhat larger and more coalesced; independent spots only at apices of veins R2+3, R4, Cu1} at medial

angle of vein R4, and in cell Ri below tip of vein R^

VARIATION.—There is little variation in the specimens examined from the Bahama Islands. The second

antennal segment may be black or red and the tibiae

vary from reddish yellow to dark red. The size and

degree of coalescence of the spots on the wings are

variable but are encompassed approximately by the

limits of the holotype and allotype.

A series of nine specimens from Haiti exhibit some

consistent variations from the typical species. The

white pile in front of the base of the wing is more

sparse and has black setae intermixed; the scales on

the lower part of the thoracic pleura are gold rather

than black, and some scales are usually white. The

posterior margins of the femora have predominantly

gold scales. The infuscation of the wing is similar, but

the area distad to the humeral crossvein in cell C

is sometimes subhyaline and poorly defined from the

infuscated portion. Two specimens from Puerto Rico

are intermediate between those from the Bahamas

and those from Haiti.

MALE GENITALIA (Figure 33).—Similar to pluto

pluto. Distal segment of gonostylus shorter, exterior

side concave at base.

FEMALE GENITALIA (Figure 44).—Similar to pluto

pluto. Tenth tergum with eight spines on each side.

Sclerite on each side of spermathecal ducts shorter and

thicker above; lateral arm not clavate, broadened

basally; dorsal process broad, acuminate, and laterally

curved. Spermathecal ducts with two convolutions before widening into bulbs.

BODY LENGTHS.—9.6 to 11.8 mm.

WING LENGTH.—8.0 to 10.1 mm.

DISTRIBUTION.—Anthrax insulanus probably occurs

throughout the West Indies, although specimens have

been seen only from Haiti, Puerto Rico, and the Bahama Islands (Map 7).

1 \

MAP 7.—Distribution of Anthrax insulanus, new species.

HOLOTYPE.—<$, South Bimini Island, Bahamas,

British West Indies, VI-1951 (M. Cazier, C. and P.

Vaurie).

ALLOTYPE.— ° , South Bimini Island, Bahamas,

British West Indies, VII-1951 (C. and P. Vaurie).

PARATYPES.—British West Indies: Bahamas, 26",

East Bimini Island, VI-1951 (P. and C. Vaurie); 4d \
28

Eleuthera Island, New Portsmouth (Rock Sound),

111-28-1953 (E. B. Hayden and L. Giovannoli); d1,

New Providence Island, 4 mi. SW Nassau, IV-8-1953

(E. B. Hayden and L. Giovannoli); 3d1, South Bimini

Island, VI-iy51 (M. Cazier, C. and P. Vaurie); 2d\

South Bimini Island, VII-1951 (C. and P. Vaurie) ;

d, South Bimini Island, VIII-10/20-1951 (C. and P

Vaurie).

Puerto Rico: d, Ensenada, VII-14/19-1915.

OTHER MATERIAL EXAMINED.—British West Indies:

Bahamas, c?, $ , Cat Island, Arthur's Town, VIIVIII-1935 (W. J. Clench); d1

, New Providence Island, 4 mi SW Nassau, III-8-1953 (E. B. Hayden and

L. Giovannoli); ? , South Bimini Island, VI-1951

(Mont Cazier, C. and P. Vaurie); $ , South Bimini

Island, VIII-10/20-1951 (C. and P. Vaurie).

Haiti: d1, 4$ , Bois Caradeux, VIII-10/11-1934

(E. M. Ducasse); $ , Port-au-Prince ("Student Collection") ; $ , Port-au-Prince, 1-15/23-1922; d1

, Portau-Prince, IH-3-1923 (R. C. Smith); ? , St. Marc,

III-3O/IV-2-1922 (C. H. Curran).

Puerto Rico, d\ Ensenada, VI-14/19-1915 (C. H.

Curran); d1, Mamayes, 111-31-1925; d1, "Monals,"

VIII-1939 (L. F. Martorell).

TYPES.—The holotype and allotype are in the collection of the American Museum of Natural History.

Paratypes may be found in the collections of the author,

R. H. Painter, the American Museum of Natural History, and the Florida State Plant Board.

DISCUSSION.—Anthrax insulanus may be separated

from other species in the pluto subgroup by the absence

of brown or black hairs on the upper half of the

mesopleuron.

A female from 12 mi. SE of Tololapan, Oaxaca,

Mexico, collected by R. H. and E. M. Painter, is very

similar to specimens of insulanus. It has less white pile

on the upper half of the mesopleuron and above the

wing, and the wing is slightly darker. If this is a specimen of insulanus, the distribution of the species probably includes the Gulf coast of Central and South

America, as far north as central Mexico.

Anthrax stellans (Loew)

Argyramoeba stellans Loew, 1869, p. 28 [Argyromoeba].—

Coquillett, 1894, p. 95.—Kertesz, 1909, p. 67.

Spongostylum stellans.—Aldrich, 1905, p. 223 [Spogostylum].—Cole and Lovett, 1921, p. 244 [part] [Spogostylum].

Anthrax stellans.—Painter and Painter, 1965, p. 432.

Argyramoeba pluto.—Osten Sacken, 1877, p. 244 [part].—

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Osten Sacken, 1878, p. 90 [part] [not Wiedemann, 1828;

misidentification].

Anthrax pluto.—Johnson, 1925, p. 108.—Curran, 1927, p.

85 [not Wiedemann, 1828; misidentification].

MALE.—Scales on head black, white, and sometimes

brown. Second antennal segment saucer shaped with

apical margin produced as sharp flange.

Scales on mesonotum mostly black, few white and

brown ones present, especially along margins. Scutellum with white, brown, and black scales. Pile on mesopleuron and pteropleuron black, white, and brown.

Scales on pleura white and brown. Prosternum, propleuron, and anterior margin of mesonotum with

mixed brown, white, and black pile. Postalar tuft of

pile black, few white hairs sometimes present, some

white scales basally. Scales on coxae white, few brown

ones sometimes present.

Cell C of wing (Plate 2a) entirely infuscated or

partly subhyaline beyond humeral crossvein; cell Sc

infuscated except apex and stigmatic area. Remainder

of wing with coalesced spots at bases of cells Cul5 1M2

and Ri, R2+3, and R5, R, and 2M, and extending

from vein Mi+2 to vein Rx midway between bases of

cells R2+3 and R4. Isolated spots at bases of cells R4,

Mi, and 2M2, below tip of Rx in cell Ri, apically and

subapically on vein R2+3, medially and apically on vein

R4, apically on vein M2, medially on m crossvein, and

apically on vein Cu2. A large, poorly defined spot

medially on vein 2A extending across cell 1A and

halfway across cell 2A. Cell 2M hyaline to partly

infuscated medially. A small spot often present apically

on vein Cui.

Fore and middle femora with white scales posteroventrally and black scales anterodorsally, some goldenbrown scales present between black and white ones.

Hind femur with golden-brown scales ventrally and

black scales elsewhere. Sometimes a few white scales

basally.

Pile on sides of first abdominal tergum white, few

black hairs posteriorly; posterior margin with white

scales laterally and black scales medially. Sides of terga

two through six with black pile and scales, with some

white and brown scales, especially posteriorly; white

scales on sides of seventh tergum. Discs of terga two

through five with black scales; posterior margins with

submedial and sublateral patches of white scales,

patches on two and three often enlarged and coalesced,

sublateral patches on four and submedial patches of

five reduced; some brown scales sometimes mixed with

white. Terga six and seven with black scales medially
NUMBER 43 29

and white scales laterally. Scales on first sternum

threadlike, white; posterior sterna with white scales

apically and golden-brown scales preapically and

sometimes basally.

FEMALE.—Similar to male. Infuscation of wing

(Plate 2b) more extensive; cell 2M and apical portion of cell R often entirely infuscated; spots in distal

portion of wing larger. White scales more extensive

on femora, sometimes occupying as much as one-third

of ventral margin of hind pair.

MALE GENITALIA (Figure 30).—Similar to pluto

pluto. Apex of epiphallus flared; apex of aedeagus enlarged, slightly flared, gonopore apical.

FEMALE GENITALIA—Similar to seriepunctatus. Ducts

of spermathecae with two convolutions before expanding to bulbs.

BODY LENGTH.—8.5 to 11.5 mm.

WING LENGTH.—7.6 to 10.6 mm.

DISTRIBUTION.—Anthrax stellans occurs in coniferous forests in Canada, south along the Coast Range

and Sierra Nevada into southern California, in the

Rocky Mountains into Colorado and central Utah, and

at higher elevations along the Canadian border in

northeastern United States (Map 8). It may occur

as far north as the extent of the spruce forests in

northern Canada.

A series of specimens from Tifton County, Georgia,

appear to belong to this species. If so, they are probably a relict population surviving from the last

glaciation.

TYPES.—The type, which is in the Museum of Comparative Zoology at Harvard University, is badly rubbed and molded. Contrary to Loew's statement, the

type is a female. The type-locality is "Oregon."

DISCUSSION.—Anthrax stellans differs from all other

species in the pluto subgroup in having golden-brown

scales ventrally on the hind femur and on the abdominal sterna.

Anthrax seriepunctatus (Osten Sacken)

Argyramoeba seriepunctata Osten Sacken, 1886, p. 103.—

Kertesz, 1909, p. 67.

Spongostylum seriepunctatum.—Aldrich, 1905, p. 223

[Spogostylum].

Anthrax seriepunctatus.—Painter and Painter, 1962, p. 81,

1965, p. 432.

MALE.—Scales on head black and white. Second

antennal segment saucer shaped, distal margin produced as sharp flange.

Scales on mesonotum black and white, few gold

scales present, especially on margins. Scutellum with

white and black scales, few gold scales intermixed

with white. Pile on mesopleuron and pteropleuron

black and white; scales on pleura threadlike, white.

Pile on prosternum, propleuron, and anterior margin

of mesonotum mixed black and white, few gold hairs

sometimes present; postalar tuft of pile white or mixed

black and white. Scales on coxae white.

Cells C and Sc of wing (Plate 2c) subhyaline or

hyaline. Remainder of wing largely hyaline, isolated

spots or clouds at bases of all cells in outer part, on vein

R5 halfway between bases of cells R2+3 and R*, and

in cell R below base of cell Rx; sometimes a faint cloud

submedially on vein 2A and medially on m crossvein.

Fore and middle femora with black scales anteriorly,

white scales posteriorly; hind femur with black scales

anteriorly, mixed black and white scales posteriorly.

Sides of first abdominal tergum with white pile;

posterior margin with long, threadlike white scales.

Sides of second tergum with black pile, few white

scales posteriorly; sides of terga three, four, and five

with black pile and scales anteriorly, and white scales

posteriorly, white scales sometimes reduced on four;

sixth and seventh terga with white scales laterally,

sometimes with a few black scales anteriorly on fifth.

Discs of terga two through five with black scales anteriorly and linear white scales across posterior margins,

few black scales sometimes interrupting white medially

on two and three, and submedially on four and five;

sixth and seventh terga with white scales laterally and

black scales medially. First sternum with sparse linear

white scales; second and third sterna with black scales

anterolaterally and white scales elsewhere; scales on

posterior sterna predominantly black, few white ones

along posterior margins.

FEMALE.—Similar to male. White scales on anterior

margin of scutellum reduced or replaced by gold scales.

Scales on posterior femur often entirely black. Spots

on wing (Plate 2d), often somewhat larger.

MALE GENITALIA (Figure 28).—Similar to pluto

pluto. Distal segment of gonostylus short, broad at

base. Apex of epiphallus shortened and broadened,

ventrolateral parts almost vertical. Apex of aedeagus

slightly flared, gonopore apical. Ventral bands swollen

dorsally at junction with aedeagus.

FEMALE GENITALIA (Figure 42).—Tenth tergum

with nine spines apically on each side; ventral extensions at base sharply pointed apically. Sclerite on each

side of gonopore hatchet shaped, lateral extension
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

stellans (Loew)

seritpunctotus COsten Sacken)

MAP 8.—Distribution of Anthrax stellans and A. sereipunctatus.

broad, dorsal margin angled upward; dorsal extension

short, broadly angled, ventral extension narrow, curved

mesally. Ducts of spermathecae with three convolutions before expanding to bulbs; bulbs bent before

middle, narrow basally, broadened after bend and

constricted at apex.

BODY LENGTH.—9.2 to 12.3 mm.

WING LENGTH.—3.0 to 10.3 mm.

DISTRIBUTION.—The few specimens of seriepunctatus available indicate that the species occurs in the

desert shrub area of the southwestern United States

and central and western Mexico, and eastward into the

mesquite and oak-hickory areas in central Texas (Map

8).
NUMBER 4 3 31

N EW MATERIAL EXAMINED.—Arizona: Gila Co., $ ,

Globe, IX-23-1933 (Parker).

New Mexico: Torrance Co., <$, VI-24, 7,000 feet

(R. H. Painter).

Texas: Brewster Co., cf, Big Bend National Park,

VI-23-1947 (L. D. Beamer); ? , Big Bend National

Park, VII-30-1956 (E. G. Matthews); ? , Glen

Spring, VI-20-1928 (F. M. Gaige); ? , Marathon,

VI-25-1947 (A. C. Michener). Brown Co., tf,

Brownwood, VI-6-1936 (R. H. Painter). Pecos Co.,

<$, ? , Sheffield, VII-5-1917 (J. Bequaert). San Saba

Co., d1, San Saba, VII-23-1921. Sutton Co., <S,

Sonora, V-4-1954 (L. D. Beamer). Travis Co., $ ,

Austin, IV-27-1902. Val Verde Co., <?, 16 mi. SE Del

Rio, IV-10-1950 (Beamers, Stephen, Michener.

Rozens).

Mexico: Baja California Sur, $ , San Ignacio,

IX-29-1941 (Ross and Bohart). Sonora, $ , Guaymas,

IV-11-1921 (E. P. Van Duzee).

TYPES.—Anthrax sereipunctatus was described by

Osten Sacken (1886) from a male and female from

northern Sonora and two females from "Tehuacan."

Painter and Painter (1962) report that the male and

female are in the British Museum and the two females

from "Tehuacan" are in the Turin Museum. The male

was designated as lectotype. The specimens from

Tehuacan are far out of the range postulated for this

species. Both Osten Sacken and Painter and Painter

indicated that they are similar to the specimens from

northern Sonora so they probably are specimens of

either seriepunctatus or atriplex. In either case, an extension of the postulated range of these species down

the eastern coast of Mexico would be indicated.

DISCUSSION.—Anthrax seriepunctatus differs from

atriplex in lacking brown or black scales on the

thoracic pleura, in having long, narrow white scales

predominating on abdominal sterna one through three,

and in having linear white scales across the posterior

margins of abdominal terga two through five. It differs

from other species in the pluto subgroup in that cell

2M is completely hyaline except for small spots at the

extreme base and apex, and in having at most only

a small cloud submedially on vein 2A.

Anthrax atriplex, new species

Argyramoeba pluto.—Osten Sacken, 1886, p. 102 [not Wiedemann, 1828; misidentification].

Anthrax atriplex Marston.—Krombein, 1967, p. 402.

MALE.—Scales on head black and white. Second

369-673 O—70 3

antennal segment saucer shaped, apical margin produced as a sharp flange.

Scales on mesonotum mostly black, some brown and

white ones, especially along lateral margins. Scales on

scutellum black and white, few brown scales sometimes present. Pile on mesopleuron and pteropleuron

white and brown. Scales on pleura brown and white.

Prosternum, propleuron, and anterior margin of

mesonotum with mixed black and white pile, some

brown hairs on first two. Postalar tuft of pile mixed

black and white. Scales on fore coxa brown and white.

Cell Sc of wing (Plate 2k) entirely infuscated except

for stigmatic area and apex; cell C infuscated before

humeral crossvein except for distal hyaline spot, subhyaline to hyaline beyond humeral crossvein. Remainder of wing with spots at bases of cells R4, Mi,

and 2M2, coalesced spots at bases of cells R2+3 and R8,

Gul5 1M2 and R1} and R and 2M; isolated spots in cell

Ri, below tip of vein Rl5 apically and subapically on

vein R2+3, on medial angle of vein R4, on medial angle

of m crossvein, subapically on vein Cu2, and submedially on vein 2A (extending broadly into both cells

1A and 2A); transverse spot present medially on vein

R4+5 extending across cell R2+3, and longitudinal spot

present postmedially in cell Ri.

Scales on fore and middle femora and tibiae black

anteriorly, narrowly white posteriorly. Scales on hind

femur and tibia entirely black.

Sides of first abdominal tergum with white pile, with

few brown hairs posteriorly; posterior margin with

white scales laterally and black scales medially. Sides

of terga two through five with black pile and scales;

sides of terga six and seven with few black scales and

numerous white scales. Discs of terga two through five

with black scales except for submedial and larger sublateral patches of lanceolate white scales on posterior

margins; spots of white scales reduced laterally on

segment four, coalesced medially on segment five;

terga six and seven with black scales medially and

white scales laterally. Scales on venter black and

brown, few white scales present posterolaterally on two,

five, six, and seven.

FEMALE.—Similar to male. Brown hairs less numerous on thoracic pleura, absent on prosternum and

propleuron. Postalar tuft of pile almost entirely white.

Spots on wing (Plate 2/) larger; cells C and 2M partially infuscated medially. Some white scales posteriorly

on sides of abdominal terga; spots of white scales more

extensive on discs of abdominal terga, few gold scales

submedially on posterior margins of two and three.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Scales on first abdominal sternum curly, linear, white;

dark scales on remainder of sterna entirely black.

VARIATION.—The color of the linear scales on the

posterodorsal part of the sternopleuron varies from

white or yellow in the west to brown in the east. The

remaining scales vary from white to yellowish white.

The postalar tuft of pile varies from entirely white in

some specimens from Texas to predominantly black

in a few specimens from southern California. The

brown hairs on the thoracic pleura are often absent in

specimens from the western part of the range. The

pigmentation of the wing may be slightly darker or

lighter than that described for the types. There may

be some yellowish or white scales along the posterior

margins of the abdominal sterna, but the scales on the

anterior parts of the sterna are always black.

MALE GENITALIA (Figure 31).—Similar to pluto

pluto. Apex of epiphallus flared outward; apex of

aedeagus enlarged, slightly flared, gonopore apical.

FEMALE GENITALIA (Figure 45).—Similar to pluto

pluto. Tenth tergum with 12 spines on each side.

Lateral arm of sclerite on each side of spermathecal

ducts broad; ventral arm twisted. Ducts of spermathecae with two convolutions before expansion to bulbs;

bulbs broadest postmedially.

BODY LENGTH.—6.9 to 11.0 mm.

WING LENGTH.—6.1 to 10.8 mm.

DISTRIBUTION.—Anthrax atriplex occurs in the

desert scrub zone in the southwestern United States

from southern Texas to southern California, south into

central and western Mexico and north in the Great

Basin (Map 9). It is allopatric to melanopogon.

HOLOTYPE.—o", 10 mi. NW Nogales, Santa Cruz

Co., Arizona, IV-27-1961, 3600 feet (R. H. and E. M.

Painter).

ALLOTYPE.— ? , same data as holotype.

PARATYPES.—Arizona: Cochise Co., Bowie, VI-23-

1954 (G. D. Bulter); ? , 2 mi. NE Portal, V-18-1961

(R. H. and E. M. Painter); $, 3 mi. W Rodeo, N. M.,

VII-9-1956 (H. and A. Howden); d\ VII-29-1927

(R. H. Beamer). Gila Co., 4c?, 4? , Globe, VII-27-

1932 (R. H. Painter) 2d", ? , Globe, V-9-23-1935

(Parker); d1, Wheatfields, near Globe, VIII-19 (D. K.

Duncan); 8o*, Miami, VII-22-1932 (R. H. Beamer);

d, 2$ , Tonto National Monument, VII-26-1932

(R. H. Painter). Graham Co., ?, Thatcher, IX-6-

1947 (E. J. Taylor). Pima Co., d\ Baboquivari Mtns.,

IV-29-1935 (F. H. Parker); 2dl, 9, Baboquivari

Mtns., VII-24-1941 (R. H. Beamer and E. L. Todd);

d

1

, Baboquivari Mtns., VI-19-1950 (L. D. Beamer);

MAP 9.—Distribution of Anthrax atriplex.

$, Organ Pipe National Monument, IV-13-1947

(A. L. Melander); d1, Sabino Canyon, V-26-1962

(F. D. Parker and L. A. Stange); 2d1, Baboquivari

Canyon, IV-9-1963 (R. C. Dickson); 2o", $, Saguaro

National Monument, V-3, 9-1960 (G. Butler); d1,

Santa Catalina Mtns., V-l 1-1961 (J. Bequaert) ; d1,

Santa Rita Mtns., VIII-9-1930 (R. H. Painter); $,

Santa Rita Mtns., VIII-18-1935 (E. I. Beamer); $,

Santa Rita Mtns., VII-20-1937 (Ruggles); 3 ? , Sells,

VII-31-1932 (R. H. Painter); ? , 25 mi. SE Sells,

VIII-2-1932 (R. H. Painter); d1, Tucson, VIII-3-

1930 (R. H. Painter); $, Tucson, IX-3 (F. M. Carpenter); d1

, 9, 36 mi. SW Tucson, VIII-3-1932

(R. H. Painter); d1, Tucson, IV-27-1948 (A. L. Melander); d1

, Cat Pass, Tucson Mtns., VIII-29-1913

(W. D. Pierce). Final Co., $, 20 mi. S Florence, VIII3-1949 (F. Werner and W. Nutting); ? , Sacaton,

VII-24-1932 (R. H. Painter); d, Rimrock, IV-29-

1925. Yavapai Co., 4dl, 3$ , 5 mi. N Wickenberg,

IV-30-V-14-1937 (L. K. Gloyd).

California: Los Angeles Co., °., Attadena, VI-8-

1914; 5$ , Bautista Canyon, V-5-1931 (C. H. and

D. Martin); $, Tanbark Flat, VII-4-1950 (F. X.
NUMBER 4 3 33

Williams); d, Topango Canyon, VIII-5-1938 (Jean

Russell). Riverside Co., ? , Anza, VIII-6-1935 (E. I.

Beamer); ? , McCoy Spring, IV-9-1963 (E. I.

Schlinger). San Bernardino Co., ? , Wildwood Canyon, VII-13-1957 (H. R. Moffitt); ? , 4 mi. N Cajon

Jet., VII-4-1958 (J. C. Hall). Santa Barbara Co., $,

Bluff Camp, San Rafael Mtns., VI-29-1959.

New Mexico: Dona Ana Co., $, Las Cruces, IV25-1954 (R. H. Beamer). Hidalgo Co., d, Steins,

VIII-8-1932 (R. H. Painter). Otero Co., d, Alamagordo, IV-30-1902. $, "Hot Springs," VII-22-1950.

Texas: Brewster Co., d, 20 mi. S Marathon, IV12-1949 (Michener and Beamer). Dimmit Co., $,

Carrizo Springs, IV-14-1949 (Michener and Beamer).

Maverick Co., d, Quemado, IV-14-1949 (Michener

and Beamer). Tom Green Co., d, San Angelo, VIII29-1924 (R. H. Painter). Uvalde Co., d, VII-26-

1933 (A. W. Linquist). Webb Co., d, Laredo, V20/24-1948 (W. Nutting). Zavala Co., d, Crystal

City, IV-14-1952 (Michener, Beamer, Wille, and

LaBerge).

Utah: Iron Co., d, Parowan, VII-25-1919.

Mexico: Baja California Sur, ? , 10 mi. SW San Jose

del Cabo, VII-9-1938 (Michelbacher and Ross); $,

"Canipole," X-2-1941 (Ross and Bohart). Durango,

$, 15 mi. SW Pasaja, VIII-28-1929 (H. Smith).

Sonora, ? , Guaymas, VII-2-1952 (S. C. Dorman);

d, HermosiUo, VIII-12-1959 (W. L. Nutting and

F. C. Werner); ? , 35 mi. N HermosiUo, IV-26-1961

(R. H. and E. M. Painter); d, ?, 75 mi. S HermosiUo, IV-25-1961 (R. H. and E. M. Painter). Tamaulipas, d, 35 mi. S Matamoras, VIII-26-1954 (J. G.

Chillcott) j ? , Santander Jimenez, VII-19-1962 (N.

Marston-6).

Additional specimens have been examined from

Gila, Maricopa, Pima, Pinal, and Santa Cruz counties, Arizona; Los Angeles, Riverside, and San Diego

counties, California; Hidalgo County, New Mexico;

Lake County, Oregon; Dimmit and Sutton counties,

Texas; and Baja California Sur and Durango, Mexico.

TYPES.—The holotype and allotype are deposited

in the collection of R. H. Painter. Paratypes may be

found in the coUections of the author, R. H. Painter,

Jack C. Hall, the University of Arizona, the Canadian

National Collection, the University of Kansas, Utah

State University, the Museum of Comparative Zoology,

the University of California, the Academy of Natural

Sciences of Philadelphia, the University of Michigan,

Oregon State University, the California Academy of

Sciences, and the U.S. National Museum.

BIOLOGY.—Two specimens were reared by Dr. K. V.

Krombein (1967) from the nests of Megachile gentilis

Cresson (Hymenoptera: MegachiUdae).

DISCUSSION.—Anthrax atriplex differs from seriepunctatus in having some black scales on the thoracic

pleura, in having few, if any, white scales on the anterior three abdominal sterna, and in having the white

scales on the posterior margins of abdominal terga two

through five separated by patches of black scales. It

differs from melanopogon in having the scales on the

underside of the abdomen mostly black rather than

mostly white, and in having the postalar tuft of pile

mostly white rather than mostly black.

Anthrax melanopogon (Bigot)

Argyramoeba melanopogon Bigot, 1892, p. 348 [Argyromoeba].—Coquillett, 1894, p. 95.—Kertesz, 1909, p. 65.

Spongostylum melanopogon.—Aldrich, 1905, p. 223 [Spogostylum].

Anthrax melanopogon.—Painter and Painter, 1962, p. 77;

1965, p. 432.

Spongostylum stellans.—Cole and Lovett, 1921, p. 244 [part]

[Spogostylum] [not Loew, 1869; misidentification].

Anthrax seriepunctatus.—Maughan, 1935, p. 33 [seriepunctata] [not Osten Sacken, 1886; misidentification].

MALE.—Scales on head black and white. Basal antennal segments black; second segment saucer shaped,

apical margin produced as sharp flange.

Scales on mesonotum mostly black, some white and

gold scales present, especially along margins. Scales on

scutellum black and white. Pile on mesopleuron and

pteropleuron black and white, few brown hairs present.

Scales on pleura mostly white, black scales sometimes

present. Pile on prosternum, propleuron, and anterior

margin of mesonotum mixed black and white, sometimes few brown hairs on last two; postalar tuft of pile

mixed black and white, usually predominantly black.

Scales on coxae white.

Cell C of wing (Plate 2m) usually entirely infuscated, sometimes hyaline along anterior margin beyond

humeral crossvein. Base of wing infuscated out to bases

of cells R, 2M, 1A and 2A; broad infuscated areas submedially on vein 2A, from base of cell Cux to base of

cell R, and from vein Rx to vein Mi+2 between bases

of cells R2+3 and R4; medial portion of cell 2M sometimes partly infuscated; isolated spots at bases of cells

R2+3 and R5, R4, Mi, and 2M2, at medial angles of vein

R4 and m crossvein, apically on veins R2+3, M2, Cul5

and sometimes Cu2, subapically on vein R2+3, and below

tip of vein Rx; basal third of cell 1M2 sometimes partially infuscated.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Femora dark red to black, apices sometimes lighter.

Scales on fore and middle femora black anteriorly,

white posteriorly; hind femur with black scales, sometimes with few white scales posteroventrally.

Pile on sides of first abdominal tergum mostly white,

some black hairs posteriorly; posterior margin with

white scales laterally and black scales medially; sides

of terga two through five with black setae, pile, and

erect linear and lanceolate scales, some white scales

posteriorly on three and five; sides of sixth and seventh

terga with white scales. Discs of terga two through five

with black scales; patches of white scales submedially

on two through five, and sublaterally on four and five;

patches of larger white scales sublaterally on two and

three; terga six and seven with small black scales anteriorly on six, and medially on six and seven, remainder with white scales. Venter with sparse, threadlike

white scales, usually some black scales medially on

sterna three through seven.

FEMALE. Wing (Plate 2n) usually more extensively

infuscated, cell 2M and bases of cells 1A and 2 A entirely infuscated; spots on remainder of wings larger

and sometimes coalesced.

MALE GENITALIA (Figure 32).—Similar to pluto

pluto. Distal segment of gonostylus longer.

FEMALE GENITALIA.—Similar to seriepunctatus.

Tenth tergum with nine spines. Ducts of spermathecae

with two convolutions before widening to bulb.

BODY LENGTH.—9.2 to 12.2 mm.

WING LENGTH.—8.1 to 10.4 mm.

DISTRIBUTION.—Anthrax melanopogon occurs in

mountain forests from southern California north to

British Columbia and south in the Rocky Mountains

to southern' Colorado, northern New Mexico, and

northern and eastern Arizona (Map 10). The typelocality is "Washington."

TYPES.—Painter and Painter (1962) designated as

lectotype for this species a male specimen in the Bigot

collection in the British Museum. They noted that the

specimen was headless but otherwise in good condition

and agreed well with Bigot's description.

DISCUSSION.—Anthrax melanopogon differs from

atriplex in having the scales on the underside of the

abdomen mostly white. It differs from stellans in having the scales on the ventral part of the hind femur

black rather than golden brown and white, and in

lacking gold scales on the underside of the abdomen.

It differs from seriepunctatus in having cell 2M partially infuscated medially.

Anthrax cybele (Coquillett)

Argyramoeba cybele Coquillett, 1894, p. 96.—Kert6sz, 1909,

p. 62.

Spongostylum cybele.—Aldrich, 1905, p. 222 [Spogostylum].

Anthrax cybele.—Painter and Painter, 1965, p. 431.

MALE.—Scales on head black, and white or yellow.

Second antennal segment lenticular, apical margin not

produced as a sharp flange.

Scales on mesonotum mixed yellowish white and

black, margins with white scales, few yellow ones intermixed and some black ones on postalar callus. Scales

on scutellum black, white, and yellow. Pile on mesopleuron and pteropleuron black. Scales on pleura gold

and yellow. Prosternum with black pile; propleuron

and anterior margin of mesonotum with mixed black

and white pile, few yellow hairs on former. Postalar

tuft of pile white. Scales on coxae gold and black.

Cells C and Sc of wing (Plate 2o) entirely infuscated. Cells R, 1M, and 2M entirely infuscated

except for subapical and submedial hyaline spots in

former, and apical stigmatic area in latter. Cell 1A

with postbasal subhyaline spot, and postmedial and

apical hyaline areas. Cell 2A with broad infuscated

area submedially on vein 2A not reaching posterior

margin, and small subapical spot on vein 2A. Cell Ri

infuscated out to tip of vein Sc except for basal hyaline

spot; cell 1M2 infuscated on basal third except for

small hyaline spot basally on vein Cu^ Bases of cells

R2+3J RSJ and Cul3 infuscated. Independent spots at

bases of cells R4 and Mi, in cell Rx below tip of Rl5 on

vein R4+5 between bases of cells R2+3 and R4, and

medially on m crossvein.

Fore and middle femora and tibiae with black scales

posteriorly and white scales anteriorly; hind femur

and tibia with black scales.

Sides of first abdominal tergum with dense white

pile, few yellow hairs posteriorly; hind margin with

black scales medially and white scales laterally. Sides

of terga two through seven with black setae and few

linear black scales. Discs of terga with mostly black

scales; white scales laterally and medially on posterior

margin of second tergum, laterally on terga six and

seven, and medially on third tergum. Venter with

linear black scales, a few yellowish-white scales laterally on posterior margins of second and following

sterna.

FEMALE.—Similar to male. Sternopleuron with

some black scales. Cell 2M of wing with hyaline area

basally. Venter of abdomen without light scales.
NUMBER 43 35

MALE GENITALIA (Figure 29).—Distal part of gonocoxites parallel sided in lateral view, rounded apically;

apices sharply rounded in ventral view, with mesal

ridges defining shallow medial sulcus. Distal segment

of gonostylus inverted-trapezoidal basally, with narrow

apical hook. Apex of epiphallus with sharp lateral

angles, proximal process large, sharply recurved.

Aedeagus narrow and sharp apically, gonopore ventral. Basal part of aedeagus enlarged, bulbous; lateral

and proximal apodemes small in relation to base of

aedeagus.

FEMALE GENITALIA (Figure 40).—Similar to pluto

pluto. Tenth tergum with 12 spines on each side.

Sclerite on each side of gonopore very narrow dorsally,

with spurlike mesal extension and narrow, spatulate

outer lobe.

BODY LENGTH.—6.7 to 7.4 mm.

WING LENGTH.—6.1 to 6.7 mm.

DISTRIBUTION.—Anthrax cybele probably occurs

from southern California to southwestern Arizona and

into northwestern Mexico (Map 10).

NEW MATERIAL EXAMINED.—Arizona: Maricopa

Co., d, Phoenix, IV-19-1931 (E. M. Painter); ?,

Gila Bend, IV-23-1935 (F. H. Parker). Pima Co.,

2 ? , 5 mi. W Tucson, VIII-5-1961, 2300 feet (R. H.

and E. M. Painter).

California: Riverside Co., $, 20 mi. E Indio, IV26-1963 (E. I.Schlinger).

DISCUSSION.—Anthrax cybele is a very distinctive

species. It may be distinguished from all the species in

the oedipus group except irroratus Say by the lenticular form of the second antennal segment. It differs

from irroratus in that cells C and Sc are entirely infuscated rather than with alternating hyaline and

pigmented spots.

Anthrax cephus Group

The cephus group is the largest and most diverse group

in the genus Anthrax in the New World. Thirty names

have been proposed to date in the group. Of these,

thirteen are synonyms, one is a homonym, and one is

relegated to subspecific rank. In addition, twelve new

species and one new subspecies are described here,

making a total of 29 taxa known from North and

South America.

Specimens of the cephus group may be separated

from species of other groups most readily by the setae

on the face, which extend almost to the level of the

antennae rather than restricted to the lower and lateral

melanopogon CBigot)

cybele CCoquillett)

MAP 10.—Distribution of Anthrax melanopogon

and A. cybele.

margins. Anthrax plurinotus of the trimaculatus group

has the setae distributed in this way, but it has distinct,

isolated spots at the bases of the apical cells.

Approximately 1400 specimens in this group have

been studied. These represent the majority of specimens in North and South American museums. All except about 20 have been assigned with relative certainty to the various taxa recognized. The few unassigned specimens may represent individual variants,

intermediates between species or subspecies, or possibly

distinct taxa too poorly represented to be recognized.

These specimens are discussed under the species to

which they seem most closely related.

Group Description

Body generally black; tibiae and male genitalia

usually red or yellow, femora and parts of thoracic
36

pleura, basal antennal segment, and parts of abdominal

sterna sometimes red or yellow. Pruinosity generally

gray and brown, silver along margins of eyes, sometimes velvet black above on thorax and abdomen. Head

globoid, eyes separated at vertex by 1.0 to 1.5 times

width of ocellar tubercle in male, 2.0 to 2.5 times its

width in female. Antennal sockets separated by 0.25 to

1.0 of their diameters, separated from eye margins by

0.5 to 0.75 of their diameters. Face projecting slightly

above level of eyes, oral margin evenly rounded or

obtusely angular. Front covered with light or dark

setae and sparse lanceolate scales of variable color. Face

with dense setae extending from oral cavity almost to

antennal sockets; sparse scales along eye margins and

sometimes over entire face. Setae and scales on occiput

and fringe of pile on posterior margin unicolorous or

light below and dark above. First antennal segment

narrow basally, more or less enlarged mesad apically,

0.75 to 2.0 as long as apical width and entirely covered

with setae. Second segment lenticular or globoid, somewhat narrower than apex of first segment and with

setae dorsally and laterally. Third segment bulbous or

hemispherical basally, tapering abruptly or gradually

to a styliform apex; base sometimes flattened mesolaterally, usually as wide as second segment in mesal

view and slightly narrower in dorsal view; styliform

part 0.5 to 1.5 times as long as base, 1.0 to 2.5 times as

long as style; style 1.0 to 4.0 times as long as wide, with

a distinct apical tuft of setae.

Mesonotum with fine setae scattered over disc and

with bristles and coarse setae along lateral margins;

recumbent linear scales present on disc, becoming

curly and semierect along lateral margins; anterior

margin with long hairs and setae. Scutellum with fine

setae and linear scales on disc, and long setae and

broader scales posteriorly and laterally. Mesopleuron

with coarse setae and pile above, with fine setae, pile

and erect scales below. Pteropleuron with coarse setae

and fine pile and scales on anterior half, posterior part

usually bare, rarely with long linear, curly scales.

Sternopleuron with fine setae and linear or lanceolate,

recumbent scales. Hypopleuron bare or with a patch of

short setae or scales at anterior corner. Metapleuron

sometimes with a patch of linear or lanceolate scales

behind or below spiracle. Propleuron and prosternum

with long, erect pile. Postalar tuft of pile often with

a few linear scales at base. Coxae with bristles and

lanceolate semierect scales.

Wing pigmented anteriorly and basally, hyaline

apicoposteriorly, or entirely pigmented, rarely nearly

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

hyaline; no isolated spots present in apical part of

wing. Posterobasal part of wing often greatly narrowed with posterior margin of cell 2A concave, or

broadened with cell 2A twice as wide as cell 1A;

alula well developed with posterior margin rounded,

or reduced with posterior margin straight or concave.

Base of cell R4 rounded, angled with a spur, or connected to vein R2+3 forming a sectoral crossvein; vein

R4 without a medial spur. Base of cell R2+3 rounded or

with a short or long spur. R-m crossvein located at

basal fifth or two-fifths of cell 1M2. M crossvein sometimes supporting a spurious vein extending from near

its apex to posterior margin near apex of vein Cux. Contact of cells 1M2 and Cux 0.5 to 3 times as long as

base of cell Cu,. Calypteres lightly pigmented; knob of

halteres more or less pigmented basally, usually

yellowish apically.

Fore and middle femora with or without a more

or less extensive anteroventral row of macrochaetae;

middle femur sometimes with one or more bristles

postmedially on the anterior side; hind femur with a

more or less extensive row of bristles anteroventrally

and sometimes with a few bristles apically on the posterodorsal and posteroventral sides. Fore tibia with rows

of macrochaetae posteroventrally, posterodorsally and

sometimes anterodorsally; middle and hind tibiae with

rows of macrochaetae at all four angles; anterodorsal

row on latter sometimes double. Scales on femora and

tibiae entirely dark or sometimes light posteriorly;

scales on hind tibiae semierect when a double row of

macrochaetae is present anterodorsally.

Sides of first abdominal tergum with dense pile;

sides of posterior terga with fine setae and semirecumbent or semierect linear or lanceolate scales, some pile

often present laterally on terga two and three.

Posterior margin of first tergum and surfaces of following terga with sparse, fine black setae and linear or

lanceolate, dense, dark scales; broader, light scales

often present along posterolateral margins of terga one

through four. Lateral portions of fifth tergum and much

or all of terga six and seven of male often with dense,

overlapping, posteriorly or posterolaterally directed

silver scales. Females often with patches of silver scales

laterally on terga four, five, and six. Venter with fine

long setae, and scattered lanceolate or linear scales of

variable color.

Apices of gonocoxites of male genitalia rounded or

acuminate, often with medial carinae defining a

medial sulcus, rarely with lateral carinae; setae on

gonocoxites sparse and coarse, sometimes more dense
NUMBER 4 3 37

medially toward apex. Basal segment of gonostylus

usually inconspicuous, rarely produced as an acuminate

lobe or a broad, flat plate apically. Distal segment of

gonostylus variable in form, usually extending apicolaterally, but rarely dorsolaterally. Apex of epiphallus

with a dorsal spine, transverse shield, or raised plate,

sometimes flattened with recurved lateral lobes. Basal

part of aedeagus bulbous or tubular. Dorsal bands of

epiphallus bare or with fine setae.

Eighth tergum of female with dense, fine hairs extending apically, basal medial process long, narrow and

laterally flattened, or vestigial. Tenth tergum with one

to thirty stiff spines on each side surrounding the cerci.

Sclerites on ninth sternum usually hatched shaped,

quite variable in form. Ventral arms of ninth tergum

simple and acuminate or broadened apically. Spermathecal tubes of variable length, plastic or stiff, rarely

contorted before bulbs; bulbs tubular to spherical or

awl shaped, usually symmetrical, but sometimes with

apex produced slightly laterad.

Taxonomic Characters

The extent of the wing pigmentation is the most valuable taxonomic character for separating species of the

cephus group. The intensity of pigment is less useful,

however, since it is difficult to define and varies considerably in certain species. A sectoral crossvein between

vein R2+3 and the basal angle of vein R4 is present in

the males of delicatulus and innubilipennis. The species related to pluricellus have an additional vein extending from the m crossvein to the wing margin just

above the apex of vein Cux. Thus, these species may key

to the family Therevidae in some texts. This particular

vein has both origin and insertion different from that

extending from the m crossvein to vein Cux in species

of the tigrinus group.

The form of the wing is a good taxonomic character. The species luctuosus and cathetodaithmos have

the wing broader than is usual in species of Anthrax.

In this case the cells are somewhat shorter and

broader, particularly those of the anal area. Cell 2A is

more than 1.5 times broader than cell 1A which is

itself broadened. At the other extreme, the species of

the argyropygus complex have the wings elongate and

usually narrowed. The anal margin of the wing is

greatly reduced with cell 2A narrower than cell 1A

and with the posterior margin concave. The majority

of the species are intermediate between these two conditions, but there is little intraspecific variation. The

shape of the alula may be an excellent diagnostic

character in that its margin may be straight or

rounded, or it may even be vestigial as in argyropygus.

The alula is defined as the membranous lobe at the

base of the wing before the bases of cells 1A and 2A.

It is clearly shown in Plate 3/.

Under the species analis and gideon several forms

have been discriminated which have wing patterns

grossly different from those of typical specimens.

These forms probably were derived from isolated

populations that were reunited subsequently and

interbred freely with the parent population. Their

origins are discussed under the section on evolution.

As interpreted here, the forms have no taxonomic

status but are briefly described lest ignoring them

might lead to confusion.

The chaetotaxy of the legs is quite variable in the

cephus group in contrast to the albofasciatus and

oedipus groups. In the primitive condition there are

four to six strong anteroventral bristles on the middle

and hind femora. These are progressively reduced in

the species of the analis complex, with analis and

repertus having only one or two weak bristles apically.

In the species of the cephus complex the number of

femoral bristles may be increased. In addition, the

species of the cephus complex, and delicatulus and

innubilipennis, have one or more postmedial bristles

on the anterior side of the middle femur. These are

in the same position as those found on oedipus in the

oedipus group. The species of the cephus complex also

have the anteroventral row of bristles on the hind tibia

doubled with the intermixed scales semierect.

The form of the antennae is quite variable among

the different species. Since the form is very difficult to

describe and varies somewhat due to unequal shrinkage during drying, it is not used extensively in classification. The drawings (Figures 95-120), however, may

be helpful in verifying the identity of species.

The male and female genitalia are distinctive in

almost all of the species. Those of the male are

especially complex and have been most useful in segregating the species of the group. Since the best characters are internal, however, they are not emphasized

in the key and diagnoses. The terms employed are

illustrated in Figures 50 and 75.

Evolution

Since the cephus group is restricted to North and

South America, it is possible to consider more easily
38

the origins and relationships of the species. Similar

patterns of distribution have been observed in other

bombyliid genera, and it is hoped that this discussion

will provide an impetus for the future consideration of

distributional patterns and species relationships in an

evolutionary context. It should be emphasized that the

patterns expressed here are strictly hypothetical. Alternative hypotheses may explain the distribution of the

species, but the patterns described seem to fit best the

available evidence.

The patterns have been determined by considering

the distributions and phenetic relationships of the

species along with their ecological preferences, where

these are known, and, working backward through the

Pleistocene, postulating the effects of climatic and

geographical changes on the changing populations.

Since the relationships of populations become less certain as one goes back through time, the origins of

populations necessarily become more vague.

The evolution and dispersal of the species complexes

of the group are illustrated in Figures 1-6. A hypothetical center of dispersal is indicated by a black circle

and the main lines of dispersal radiate from it. Barriers

which have resulted in speciation (or subspeciation)

are indicated by broken lines. The hypothetical lines

of dispersal leading to the present distributions of the

taxa are indicated by narrow lines branching from the

main trunks.

Several modes of speciation seem to have been

involved in the evolution of the cephus group. The

first is the classical case of separation by water as by

the inundation of portions of the Isthmus of Panama

during the interglacial periods of the Pleistocene, or

by the separation of island faunas. According to Dunbar (1949), if the modern ice sheets of Greenland

and Antarctica melted, as they apparently did during

the interglacial periods, sea level would rise by as

much as 100 feet, inundating low-lying areas in the

Isthmus of Panama. Conversely, if the great continental ice sheets were restored, sea level would fall by

as much as 300 feet, shifting the shore line seaward

to the present 50-fathom line. The separation of populations in northwestern South America and southern

Mexico has occurred repeatedly and apparently has

been one of the primary factors effecting the evolution

of the cephus group. Because only portions of the

Isthmus were inundated, even during the period of

highest sea level, islands would have been formed on

which small populations could have been isolated. This

may have been the origin of the species costaricensis.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Isolation of populations due to climatic change also

has been an important factor in the evolution of the

cephus group. The cool, wet climate during the glacial periods would have resulted in the shifting toward

the equator of biotic zones and the increase in extent

of forests, whereas during the interglacial periods biotic

zones would have expanded toward the poles and

areas of desert and grassland increased in extent (Martin, 1958). Because most of the species of the cephus

group are forest inhabitants, the interruption of contiguous forest belts during dry periods has resulted in

differentiation of populations. This apparently has occurred at intervals in south-central Brazil, northeastern

Mexico, and south-central United States. Here again,

small populations may be isolated on ecological islands

surrounded by an inhospitable area, although no examples have been noted in the cephus group.

If such ecological islands were to change gradually

to a new type of habitat (e.g., from forest to grassland)

the isolates would be under strong selective pressure

to adapt to the new conditions. The result of such a

change, after dispersal in the new habitat, would be

allopatric species or subspecies occupying adjacent

habitats. This type of relationship has been observed

commonly in the Bombyliidae and probably is the way

analis and repertus became adapted to dry habitats.

It would also explain the separation of argentatus and

nitidus.

The species that retains the most generalized characters in the group is argentatus, which is endemic

to the coniferous forests of northwestern United States

and southwestern Canada. Since it has characters in

common with several Palaearctic species of Anthrax,

it seems probable that it was derived from a Eurasian

ancestral population by way of the Bering Strait, and

that it retains some primitive characters because of

the lack of selective pressure in its relatively stable

environment.

The migration of the ancestral population into

North America probably took place during the Pliocene, or perhaps slightly earlier, when floras similar

to those in northwest United States existed in Alaska

(Chaney, 1940). Although the continents of North

America and Asia were connected several times during

the glacial periods of the Pleistocene, the climate was

probably too rigorous to allow the survival of Bombyliidae at that time (Hopkins, 1959).

Cockerell (1914) indicates that no fossil Anthracinae (in his interpretation, Anthracinae and Exopro-
NUMBER 43 39

sopinae) were found in extensive collections of fossil

Bombyliidae from the Miocene Florissant shales of

Colorado. He postulates that this group arose in the

Old World and migrated to North America during

or after the late Miocene. This agrees with my

interpretation.

Although argentatus is a species adapted to northern

climates, as its ancestor presumably was, the greatest

diversification of the group has taken place in tropical

and subtropical areas. The adaptation of the ancestral

stock to tropical and subtropical forests probably took

place during and after the first glacial period of the

Pleistocene. As the climate gradually cooled during

the early Pleistocene, the ancestral population was

driven southward. According to Sears and Clisby

(1955), spruce (Picea sp.) grew in the area of Mexico

City during the most recent glacial period, which

indicates that more temperate coniferous forests (such

as those to which argentatus is presently adapted)

may then have extended south into Central America,

and probably during the previous periods. If this were

the case, the ancestral population probably migrated

as far as southern Mexico or Central America during

the first glacial period. As the boreal forest retreated

during the first part of the first interglacial period,

small populations of flies would have been isolated on

mountaintops where they would have been under

strong selective pressure to adapt to the warming environment. As the isolates adapted to the new environment they would have been free to disperse throughout

the tropical and subtropical areas from southern Mexico to southeastern Brazil.

After their dispersal during the first part of the first

interglacial period, the population would have been

split into three segments by the inundation of the

Isthmus of Panama and the separation of the tropical forest in southeast Brazil from that around the

Amazon Basin at the height of the interglacial period

(Figure 1).

One of these segments, probably that in northwestern South America, may have evolved to the luctuosus

complex. Anthrax luctuosus and cathetodaithmos are

well-defined species with a strong resemblance in the

structure of the male genitalia to argentatus, but the

broad wings with a distinctive pattern and the structure of the female genitalia indicate that there is no

close relationship with any other species of the cephus

group. The two species probably were derived from

populations isolated by an inundation of the Isthmus

of Panama and since have become partially sympatric

in Central America and northwest South America

(Figure 3). At present, luctuosus has expanded over

a wide range of tropical habitats.

The segment of the ancestral population isolated in

southeast Brazil may have evolved to the present species hylaios. Like the luctuosus complex, hylaios

shows no close relationship to other species of the

cephus group, which may indicate that it was isolated

early in the history of the group. This species has not

expanded its range beyond the Serra do Mar of southeast Brazil.

The third segment of the ancestral population, probably that in southern Mexico, apparently gave rise to

most of the remaining species of the cephus group.

After expansion of its range during the second glacial

period, this population would have been split at the

Isthmus of Panama and south-central Brazil during

the second interglacial period into three populations,

one giving rise to the analis complex in southeast Brazil,

the second to the argyropygus complex in northwest

South America, and the other to the cephus complex in

southern Mexico.

The third glacial period with the accompanying

cool, wet climate and low sea level probably allowed

the ancestral population of the cephus complex to disperse throughout southern Mexico and northwestern

South America, and into southern Brazil. During the

first part of the following interglacial period the ancestral population also migrated into eastern North

America. At the middle of the period, the four populations were isolated in these areas and developed into

distinct species (Figure 5). The relationship of these

four taxa has not changed greatly since they were isolated, although the expansion of forests during the

last glacial period has made possible the expansion of

the ranges of the Neotropical species to their present

locations. Although cephus and aterrimus are superficially similar, their basic characters show no closer

relationship than with the other species of the complex.

During the third glacial period, the pre-argyropygus population probably migrated north into Mexico

and southeast to the coast of Brazil, and one part apparently moved up the Lesser and Greater Antilles to

the island of Jamaica where it has evolved into the

species delicatulus. With the isolation of the Mexican

population during the third interglacial period, and the

expansion and contraction of the range of the popula-
40

tion during the fourth glacial period and up to the present, the species costaricensis, snowi, and pluricellus have

been disassociated. The species laticellus may have

reached Arizona during the first part of the third interglacial period, after which it was isolated (Figure 4).

Toward the end of the third interglacial period, the

pre-argyropygus ancester again expanded into Mexico,

Arizona, and eastern North America. The recent

warmer and drier climate has resulted in the isolation

of argyropygus painteri in Arizona and the semiisolation of argyropygus albosparsus in southern Mexico, as well as the restriction of the range of argyropygus

argyropygus to include eastern United States, and the

Gulf coast of Texas and eastern Mexico. Anthrax

angustipennis remains in northwestern South America

and has expanded into southeastern Brazil (Figure 6).

The exact relationship between angustipennis, macquarti, and repertus in South America is open to question. Whereas the structure of both the male and female

genitalia indicate a close relationship between macquarti and repertus, the reduced anal margin of the

wing indicates a close relationship between macquarti

and angustipennis. Also, a specimen has been studied

which appears to be an intermediate between the last

two species. A possible explanation is that macquarti

was isolated from angustipennis in southeast

Brazil during the third interglacial period. Anthrax

austrinus, a close relative of macquarti, occurs in south

Brazil. It may have been separated from macquarti by

the cold uplands in southern Parana during the last

glacial period (Figure 5).

The analis complex probably evolved from an ancestral form during the second interglacial period in

southeastern Brazil. During the third glacial period it

expanded into Central America and, during the third

interglacial period, into North America. Also, it apparently has migrated by way of the Bahama Islands

from Florida to the West Indies, probably during the

fourth glacial period. During the third interglacial period, the North American population was isolated from

that in northwestern South America and evolved to the

present species, analis. Apparently, the population in

Florida was also isolated for a short time during the

third interglacial period but was reunited with resulting

hybridization giving rise to the forms now recognized

in the southeastern United States. The West Indian

population has evolved to funebris. In South America,

the northwestern population was apparently isolated

from that in eastern Brazil by the restriction of forests

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in south-central Brazil. The northwestern population

gave rise to gideon, the southeastern population to

repertus. It is not clear how clinopictus and repertus

were separated. Perhaps the pre-repertus form became

adapted to a dry habitat, as it is now, and was semiisolated in northeastern Brazil, whereas clinopictus remained in the forest area of south-central Brazil. At

any rate, the two species are distinct because they are

now at least partly sympatric. From the last glacial

period up to the present, gideon has expanded its range

into southern Mexico and the coastal forest of southeast Brazil, while repertus has expanded into grassy or

semidesert areas from southern Brazil through the

Guianas to Central America and along the Pacific

coast southward into Peru. Anthrax analis has expanded its range throughout the United States and

northward far into Canada. In southern Mexico, it is

sympatric with gideon and intermediates have been

seen from one locality (Figure 2).

The separation of argentatus and nitidus probably

took place during the third interglacial period. When

the pre-argentatus population retreated from Arizona

during this time, a small isolated population probably

became adapted to the desert habitat and has expanded

to become allopatric to argentatus as it is at present.

In addition to the above species, four anomalous

species are known from only one locality. Anthrax

inaquosum is known from Mossor6, Rio Grande do

Norte, Brazil. It apparently is not closely related to any

other species in the group. It may have had an origin

separate from that of the cephus group in South

America.

Anthrax innubilipennis is known from only three

males collected at Iguala, Guerrero, Mexico. It is most

closely related to delicatulus from Jamaica. The origin

that would best fit into the evolutionary scheme given

above is from Jamaica into southern Mexico by way

of Costa Rica or Nicaragua during the fourth glacial

period (Figure 4).

Anthrax koebelei is known from only three complete

reared specimens labeled "Arizona." The male genitalia are most similar to the analis complex, and koebelei may have been derived from an "off-shoot" of the

pre-analis population during the third interglacial

period.

Anthrax xanthomeros is known from only one specimen collected at Benque Viejo, British Honduras. It

seems to be a rather primitive species with no close

relationship to other taxa.
NUMBER 43 41

Key to the Species of the Anthrax cephus Group

in North and South America

1. Cell 2M2 divided by a crossvein extending from m crossvein obliquely to wing margin

just above apex of vein Cui (Plate 4a) 2

Cell 2M2 not divided by such a crossvein 4

2(1). Femora black. Anterior terga and scutellum with only black scales 3

Femora yellow. Golden-brown scales along posterior margins of terga and on scutellum;

lateral margins of abdomen with tufts of golden-brown hairs posteriorly. Eastern

Mexico pluricellus Williston

3(2). Cells 1A and 2A of wing entirely pigmented; cell Ri pigmented apically only along the

anterior margin. Pigment of wing dark brown (Plate 4b). Costa Rica

costaricensis, new species

Cells 1A and 2A hyaline apically; cell Ri completely pigmented apically. Pigment of

wing light brown (Plate 4a). Arizona snowi, new species

4(1). Posterior margin of alula straight or slightly concave (Plate 5/); wing usually elongate

and narrow with anal margin reduced 5

Posterior margin of alula distinctly convex (Plate 4;); wing not particularly elongate

and narrow, the anal margin not reduced 20

5(4). Vein Rt joined to vein R2+3 by an evenly curved sectoral crossvein (Plate 5a, c). Middle

femur with a postmedial bristle anteriorly 6

Vein K* not joined to vein R1+i (in rare instances the veins may be joined by an

adventitious crossvein, in which case it is angled and may bear short spurs). Middle

femur usually without a postmedial bristle anteriorly 7

6(5). Anterior margin of wing pigmented out to apex of cell Rij cell M mostly pigmented

(Plate 5a). Jamaica and Dominican Republic delicatulus Walker, $

Wing mostly hyaline; cells C and Sc and extreme base of wing pigmented (Plate 5c).

South-central Mexico innubilipennis, new species, $

7(5). Males 8

Females 14

8(7). Cell M entirely pigmented 9

Cell M hyaline medially (Plate 5g). Arizona (Map 20)

argyropygus painteri, new subspecies, $

9(8). Posterior margin of cell 2A convex 10

Posterior margin of cell 2A concave 13

10(9). Pigment extending from cell Rj+i into cell Rs beyond r-m crossvein (Plate 5/, k, I).

Cell 2A not narrowed, as broad as cell 1A 11

Cell Rs hyaline beyond r-m crossvein. Cell 2A narrower than cell 1A (Plate 5d).

Amazon Basin, northwestern South America and southeastern Brazil (Map 19)

angustipennis Macquart, S

11(10). Cell 2A more than one-half pigmented; no spots of darker pigment present at bases

of cells. South America 12

Cell 2A entirely hyaline or only lightly pigmented at extreme base; bases of cells R«,

R2+s, and Ri, and Cui with spots of darker pigment (Plate 5/). Arizona

hticellus, new species, $

12(11). Wing pigment extending along vein Cuj almost to wing margin, leaving a large, semiisolated hyaline area apically in cells 1A and 2A; pigment in cell Ri+» extending

more than three-fourths of way to base of cell R« (Plate 5/). Dorsum of thorax and

terga with some orange-brown linear scales. Santa Catarina and Rio Grande do

Sul in Brazil austrinus, new species, $

Wing pigment not projecting along vein Cuj forming a semi-isolated hyaline area

apically in cells 1A and 2A; pigment in cell Rj+S extending less than two-thirds of

way to base of cell R* (Plate 5 k). Dorsum of thorax and abdomen without orangebrown scales. Southeast Brazil to the Andes Mountains (Map 21)

macquarti d'Andretta and Carrera
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

13(9). Mesonotum, scutellum, and anterior tcrga without patches of gold scales. Ventrolateral

part of apex of epiphallus projecting apically creating a deep medial emargination

(Figure 67). South-central Mexico (Map 20). argyropygus albosparsus (Bigot), 3

Mesonotum, scutellum, and anterior terga with patches of gold scales (except in

specimens from Florida). Ventrolateral part of apex of epiphallus not projecting

apically, medial emargination shallow (Figure 66). Eastern United States, extreme

southeastern Canada, and eastern Mexico (Map 20)

argyropygus argyropygus Wiedemann, $

14(7). Cell 2A partially pigmented basally; more than half of cell 1A pigmented. Eastern

United States to South America 15

Cell 2A hyaline; less than half of cell 1A pigmented (Plate 5n). Arizona (Map 20).

argyropygus painteri, new subspecies, 9

15(14). Pigment in cell Ri extending past tip of vein Ri to apex of cell. Eastern North America,

Mexico, and West Indies 18

Pigment in cell Ri not extending past tip of vein Ri. Northern and eastern South

America (if specimens from Arizona key out here, see laticellus) 16

16(15). Pigment extending from cell RM into cell Ri beyond r-m crossvein leaving a basal

hyaline area in cell Rs (spot of pigment sometimes very faint) (Plate 5k, I) 17

Pigment not extending into cell Rs beyond r-m crossvein (Plate 5e). Northwestern

South America, Amazon Basin and southeastern Brazil (Map 19)

angustipcnnis Macquart, 9

17(12). Wing pigment extending along vein Cu» almost to wing margin, leaving a large, semiisolated hyaline area apically in cells 1A and 2A; pigment in cell R>+i extending

more than three-fourths of way to base of cell R» (Plate 51). Dorsum of thorax and

abdomen with some orange-brown linear scales. Santa Catarina and Rio Grande do

Sul in Brazil austrinus, new species, 9

Wing pigment not projecting along vein Cu» forming a semi-isolated hyaline area

apically in cells 1A and 2A; pigment in cell Ra+i extending less than two-thirds of

way to base of cell R* (Plate 5k). Dorsum of thorax and abdomen without orangebrown scales. Southeastern Brazil to the Andes Mountains in northwestern South

America (Map 21) macquarti d'Andretta and Carrera, 9

18(15). Middle femur without a bristle postmedially on anterior side (above anteroventral

row). Eastern North America and Mexico 19

Middle femur with a distinctive bristle or bristles postmedially on anterior side. Jamaica

and Dominican Republic. (If specimens from Mexico key to this species, see A.

innubilipennis) delicatulus Walker, 9

19 (18). Mesonotum, scutellum, and anterior terga with patches of gold scales (except specimens

from Florida). Pile on mesopleuron and sternopleuron largely black or gold. Extreme

southeastern Canada, eastern United States, and eastern Mexico (Map 20)

argyropygus argyropygus Wiedemann, 9

Mesonotum, scutellum, and anterior terga without patches of gold scales. Pile on

mesopleuron and sternopleuron largely pure white. South-central Mexico (Map 20).

argyropygus albosparsus (Bigot), 9

20(4). Anterior corner of hypopleuron with a tuft of setae and/or scales. Middle femur with

a postmedial bristle or bristles above the anteroventral row 21

Anterior comer of hypopleuron bare. Middle femur without bristles above anteroventral

row 24

21 (20). Wing entirely pigmented or with only a narrow hyaline area apically 22

Apical half of wing hyaline 23

22(21). Discs of cells subhyaline, pigment darker along veins (Plate Ad). Southeastern United

State (Map 18) aterrimus (Bigot)

Pigment in discs of apical cells as dark as that along veins (Plate Ac), apical margin

of wing sometimes hyaline. Central America, Amazon Basin, and southern and

eastern Brazil (Map 17) cephus Fabricius
NUMBER 43 43

23(21). Pile on lateral margins of first tergum predominantly white. Pile on pleura white,

brown, or mixed. Pigment on wing light brown (Plate 4«). Southern Mexico

to southern Brazil (Map 15) midas Fabricius

Pile on lateral margins of first tergum black. Pile on pleura black. Pigment on

wing dark brown (Plate 4/). Southern Panama, Amazon Basin, and southern

and eastern Brazil (Map 16) hyalacrus Wiedemann

24(20). Pigment filling base of wing at least out to bases of cells Rs, 1M», and Gui 25

Wing nearly hyaline, small yellowish-brown spots present at bases of cells RM and Rs,

in cell R below base of cell Ri and at bases of cells 1M* and Cui (Plate 3Jt). Northeastern Brazil inaquosum, new species

25(24). Body densely pruinose; if integument is shining, then pigment of wing dark brown or

more extensive than in 25b 26

Body mostly shining black with sparse pruinosity. Pigment of wing light brown, translucent, extending out to bases of cells R*+s and Rs, 2M» and Cut, and leaving apices

of cells 1A and 2A broadly hyaline (Plate 3;). Arizona and southern California

(Map 1J) nitidus, new species

26(25). Apical margin of wing pattern irregular or wing entirely pigmented; apical part of

cells C and Sc pigmented 28

Apical margin of wing pattern straight, perpendicular to wing axis; apical part of

cells C and Sc not pigmented (Plate 41) 27

27(26). Posterior part of pteropleuron bare; setae, pile, and scales on mesopleuron and sternopleuron black. Fringe of hairs on posterior margin of occiput black. South-central

Mexico to Peru (Map 13) cathetodaithmos, new species

Posterior part of pteropleuron with linear, woolly, gold or yellowish-white scales; pile,

setae, and scales on mesopleuron and stemopleuron white or yellowish white, a

few black setae sometimes present. Fringe of hairs on posterior margin of occiput

white or yellowish white. Southern Mexico to southeast Brazil (Map 12)

luctuosus Macquart

28(26). Pigment on wing not extending almost to apex of cell Ri or, if so, then not extending

across cell RM to vein R*, not leaving a hyaline area before the base of cell R«.

First antennal segment dark red or black; femora, tibiae, and pleura black (femora

rarely yellow). Base of third antennal segment usually constricted abruptly to

styliform part in mesal view 29

Pigment on wing extending apically almost to apex of cell Ri and thence across cell

Rut to vein R«, leaving a hyaline area before base of cell R« (Plate 4g). First antennal

segment yellow; femora, tibiae, and lower pleura yellow. Base of third antennal

segment tapering gradually to styliform part in mesal view (Figure 103). Tropical

forests in southern and eastern Brazil (Map 14) hylaios, new species

29 (28). Femora black or dark red or, if yellow, then cell 2A more than one-half pigmented.. 30

Femora orange, with five to seven strong macrochaetae on middle and hind pairs.

Wing pattern as in Plate 3g, cell 2A less than one-half pigmented. Margins of

mesonotum, scutellum, and posterior margins of abdominal terga with rusty brown

scales. British Honduras xanthomeros, new species

30(29). Wing at least slightly hyaline apically 32

Wing entirely pigmented 31

31(30). Third antennal segment with styliform part at least 1.5 times as long as base (Figure

116). Southeast United States analis Say, f. grossbecki (Johnson)

Third antennal segment with styliform part about equal in length to base. Arizona

koebeUi, new species

32(30). Posterolateral margins of terga two and three without white scales, linear yellow scales

sometimes present; silver scales on posterior terga of male parallel to axis of body.

Styliform part of third antennal segment more than 1.5 times longer than basal part

in dorsal view (Figure 116) (analis complex) 33

Posterolateral margins of terga two and three with patches of lanceolate, truncate,

white scales; silver scales on posterior terga of male diagonal to axis of body. Styliform

part of third antennal segment about equal in length to basal part (Figure 102).

Mountainous areas from Colorado north to British Columbia and south into southern

California (Map 11) argentatus (Cole)
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

33(32). Cells 1A and 2A of wing completely pigmented or with only extreme tip of cell 1A

subhyaline. Nearctic and Neotropical species 34

Cells 1A and 2A of wing hyaline apically for a distance equal to length of r-m crossvein. (If from United States, see analis Say). Neotropical species 39

34(33). Cell 1M» of wing more than three-fourths pigmented; cell Re pigmented beyond base

of cell Mi 35

Cell 1M» of wing less than three-fourths pigmented; cell Rs not pigmented beyond base

of cell Mi, or, if so, then cell 1M2 less than half pigmented 37

35(34). Apex of cell Mi more or less hyaline; apical third or fourth of cell Cui hyaline 36

Wing of female entirely pigmented (Plate 3c); wing of male with a narrow hyaline

area along apical margin from apex of vein Rs+s to apex of vein Cui; cell Cui entirely

pigmented or with a small subhyaline area at anteroapical margin (Plate 3b).

Florida analis, f. grossbecki (Johnson)

36(35). Hyaline area apically in cell lMa of wing more or less surrounded by pigment extending

from cell Rs across cell Mi into cell 2Mi. Pigment extending from apex of cell Ri

across tip of cell R!+s leaving a preapical hyaline spot in cell Rs+»; base of cell R«

narrowly hyaline or subhyaline (Plate 4*). West Indies (Map 24)

funebris Macquart

Hyaline area apically in cell lMa open posteriorly and usually open apically, if pigment

extends from Ri into cell Mi, then pigment not extending from tip of cell Ri across

apex of cell Rjtj; base of cell R« not narrowly hyaline or subhyaline (Plate 3d-e).

(Males from West Indies, see funebris Macquart). Coastal plain of southeastern

United States, north as far as New York, and west as far as Alabama

analis, f. cedens Walker

37(34). Pigment on wing extending into cell 2M» less than length of r-m crossvein; usually

one-half or less of cell lMi pigmented; cell Rs usually pigmented as far as base of

cell Mi; pigment dark brown to velvet black (Plate 4i). Mesonotum, scutellum, and

abdominal terga usually velvet black pollinose. Abdominal terga six and seven of

male without silver scales. Tropical forest in rocky hills and mountainous areas from

southern Mexico into northwestern South America, and in the Serra do Mar along

the southeast coast of Brazil (Map 23) 38

Pigment on wing extending into cell 2M» more than length of r-m crossvein; usually

three-fifths or more of cell 1M2 pigmented; cell Rs not pigmented as far as base of

cell Mi or, if so, then cell 1M» more than two-thirds pigmented; pigment light brown

to dark brown (Plate 3a). Mesonotum, scutellum, and terga brownish pollinose.

Terga six and seven usually with overlapping silver scales. Sandy areas from southern

Canada to Central America (Map 24) analis Say (typical)

38(37). Cell Rs of wing pigmented along posterior margin for twice length of r-m crossvein

beyond base of cell Mi. Colombia and Venezuela... .gideon, i. propinquus (Schiner)

Cell Rs of wing pigmented along posterior margin for length of r-m crossvein or less

beyond base of cell M, (Plate 4») gideon Fabricius (typical)

39(33). Cell Rx of wing entirely pigmented (Plate 4fc). Body with numerous gold and dark

yellow scales and pile. Femora orange. Sao Paulo, southern Minas Gerais and northwestern Parana in Brazil to northern Argentina clinopictus, new species

Cell Ri of wing hyaline apically (Plate 4;) or, if pigmented, then body without yellow

or gold scales, or femora black or dark red. Rio Grande do Sul in Brazil into

Central America and south along west side of Andes into Ecuador and Peru

22) repertus Walker
NUMBER 4 3 45

Anthrax argentatus (Cole)

Spongostylum argentatum Cole.—Cole and Lovett, 1919,

p. 227 [Spogostylum].—Cole and Lovett, 1921, p. 244

[Spogostylum].

Anthrax argentatus.—Maughan, 1935, p. 32 [argentatum].

MALE.—Integument generally black; brown to gray

pruinose; tibiae, proximal tarsal segments, and genitalia reddish. Front covered with coarse black setae and

linear black scales; a few white scales usually present

ventrolaterally. Face bare just below antennae, with

coarse black setae and a few linear white scales on

lower two-thirds; oral margin with linear and lanceolate yellowish-white scales laterally. Occiput with

sparse, short black setae and sparse, linear black scales.

Fringe of pile on posterior margin of occiput black.

Second antennal segment rounded or with a sharp

ridge apically; base of third segment hemispherical,

slightly flattened; styliform portion and style slightly

shorter than first two segments (Figure 102).

Scales on mesonotum linear, mostly black; some

curly, white or yellowish-brown scales laterally and on

humeral and postalar calli. Scales on pleura black, a

few yellow and white scales sometimes on sternopleuron; scales on coxae black, yellow, and white. Postalar tuft of pile and that on prosternum, propleuron,

and anterior margin of mesonotum black; a few white

hairs intermixed, especially on mesonotum. Scutellum

with linear and lanceolate, mostly black scales; white

scales on posterior margin and laterally on anterior

margin.

Wing (Plate 3/) translucent brown basally, hyaline

apically, the color filling cell C, all except the apex of

Sc, the basal half of R, the extreme bases of R2+3 and

R8, the basal third of 1M2, the extreme base of 2M2

and Cux and all but the extreme apices of 1A and 2A.

Stigmatic area anterodistally in cell 2M not pigmented. Calypter not pigmented, fringe of pile black.

Alula well developed, posterior margin rounded.

Scales on femora lanceolate and ovate-truncate,

black; a few white ones sometimes posteriorly on fore

and middle pairs; scales on tibiae linear, black. Middle

and hind femora with four to six setae anteroventrally.

Pile on lateral margins of first abdominal tergum

white, a few brown or black hairs posteriorly; lateral

margins of second, third, fourth, and fifth segments

with dense black hairs, setae and erect linear scales.

Terga two, three, four, and anterior half of five with

linear and lanceolate black scales except for small submedial patches of lanceolate white scales and large

lateral patches of ovate-truncate, silvery-white scales

on posterior margins of two and three; posterior half

of fifth, and sixth and seventh sloping ventrally on each

side of meson at a 45° angle, covered with dense, overlapping, elongate-truncate, laterally projecting silver

scales except for a few lanceolate black scales and black

setae medially on six and seven, and on medial half of

five. Abdominal sterna with lanceolate scales except

for some lanceolate-truncate, white or yellow scales

along posterior margins.

MALE GENITALIA (Figure 50).—Gonocoxites short

and broad, upper margin rounding to lower margin

apically in lateral view; apices rounded and slightly

emarginate medially in ventral view, with sparse,

evenly distributed setae extending two-thirds of way to

base. Basal segment of gonostylus convex dorsally, extending to base of distal segment; distal segment with a

flat footlike part slightly emarginate apically, from

which a curved, styliform part arises dorsomedially;

footlike part with fine setae. Apex of epiphallus with a

triangular, acuminate dorsal process extending slightly

basad; apically rounded ventrolateral parts. Dorsal

bands of epiphallus joined medially, without setae. Base

of aedeagus large, bulbous, tapering to junction with

ventral bands slightly before junction of dorsal bands.

FEMALE.—Similar to male. Only a few white scales

laterally on six and seventh abdominal terga. Middle

and hind femora sometimes with only three macrochaetae anteroventrally. Fore and middle tibiae with

white scales posteriorly.

FEMALE GENITALIA (Figure 84).—Tenth tergum

with 10 spines on each side. Ventral arm of ninth tergum narrow, slightly expanded and unequally bilobed

apically. Dorsomedial angle of sclerite on each side

of gonopore not greatly produced, bent interiorly;

lateral arm formed in same plane as dorsomedial angle,

slightly bent upward and truncate apically; ventral

arm narrow, parallel sided and bent sharply mesad ventrally. Spermathecal ducts about two-thirds as long as

bulbs, strongly contorted just before joining bulbs;

middle section about twice as long as basal section,

slightly shorter than apical section before contortions;

bulbs of spermathecae elongate, tubular, tapering at

base, slightly narrowed to apical "nipples" which are

one-fifth as long as base.

DISTRIBUTION.—Anthrax argentatus occurs in

mountainous areas from southern California north

into British Columbia and south in the Rocky Mountains into Colorado (Map 11). It is generally allopat-
46

MAP 11.—Distribution of Anthrax argentatus (horizontal

lines) and A. nitidus (vertical lines).

ric to A. nitidus Marston in southern California,

although the two species were collected together at Surprise Canyon in the Panamint Mountains, Inyo Co.,

California.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TYPES.—The holotype and allotype are in the collection of the California Academy of Sciences. The

type-locality is Hood River, Oregon. Two paratypes

have been seen by the author.

DISCUSSION.—Anthrax argentatus appears to be

most closely related to nitidus, to which it is allied by

the structure of the antennae and male genitalia, and

the chaetotaxy of the legs. The two may be readily

distinguished by the wing pattern. In the past, argentatus often has been confused with analis. Besides the

characters listed in the key, analis differs in having

only one or two weak macrochaetae anteroventrally on

the middle and hind femora, rather than four to six

strong ones.

Anthrax nitidus, new species

MALE.—Integument black, shining under thin gray

and brown pruinosity; margins of eyes and lower part

of occiput white pruinose. Front and face with black

setae, extending almost to antennae on latter. Occiput

with black setae; fringe of pile on posterior margin

black, a few white hairs behind vertex. First antennal

segment much shorter than apical width, surrounding

base of second segment; second segment lenticular;

base of third segment flattened laterally, narrowing

abruptly to styliform part; styliform part about 1.5

times longer than base^ about 3 times longer than very

short style. (Figure 101).

Mesonotum and scutellum with fine black setae and

linear black scales; some linear white scales laterally

on mesonotum. Sternopleuron, mesopleuron, and anterior half of pteropleuron with black setae and sparse,

linear white scales, some black scales on last two.

Prosternum and propleuron with black pile; anterior

margin of mesonotum with mixed black and white

pile; postalar tuft of pile black, Hypopleuron and metapleuron bare. Coxae with black bristles and mixed

black and white, linear scales.

Wing (Plate 3;) translucent, light brown basally,

hyaline apically; pigment filling cells C and Sc (subhyaline apically) and basal part of wing from vein Ri

above r-m crossvein across bases of cells R2+3, RB, 1M2,

and Cui; pigment fading apically and posteriorly in

cells 1A and 2A. Base of cell R4 angled, not spurred;

r-m crossvein at basal fourth of cell 1M2; vein R2+s

arising slightly basad of r-m crossvein; contact of cells

1M2 and Cux about 1.5 times width of base of cell Cux.

Cell 1A punctiform at margin. Posterobasal part of

wing enlarged, cell 2A distinctly wider than cell 1A;
NUMBER 43

alula rounded posteriorly. Stigmatic area unpigmented.

Calypter unpigmented, fringe of hairs white. Halter

yellow, knob yellow.

Scales on legs black. Middle femur with three macrochaetae anteroventrally; hind femur with five or six

macrochaetae anteroventrally. Fore tibia without

macrochaetae anterodorsally; hind tibia with single

row of macrochaetae anterodorsally.

First abdominal tergum with white pile laterally;

posterior terga with sparse black setae and hairs laterally. Posterior margin of first tergum and discs of two

through four with linear black scales; posterior margin

of second tergum and lateral part of posterior margin

of third tergum with linear white scales. Fifth tergum

mostly with linear black scales, some lanceolate white

scales posterolaterally; terga six and seven with linear

black scales medially and lanceolate white scales laterally. Venter with sparse, fine, black setae.

MALE GENITALIA (Figure 51).—Gonocoxites very

broad, dorsal margin falling almost at a right angle to

ventral margin in lateral view; apices broadly rounded

in ventral view; mesal margins bluntly angled, medial

sulcus narrow; setae sparse, evenly distributed threefourths of way to base. Basal segment of gonostylus

slightly convex in lateral view, with a few fine setae

apically. Distal segment of gonostylus oval in cross section basally, somewhat flattened dorsoventrally; styliform part arising from dorsomedial margin and extending upward and outward, apex slightly curved

dorsally; base with sparse, short setae. Apex of epiphallus with a sharp dorsal projection and a sharp,

slightly recurved flange on each side. Dorsal bands of

epiphallus not joined medially, without setae. Base

of aedeagus'broad, bulbous with upper margin curving downward and apically to junction with ventral

bands before approximation of dorsal bands; ventral

bands with a pouchlike protuberance halfway to apex.

FEMALE.—Similar to male. Seventh abdominal tergum without white scales.

VARIATION.—The contact of cells 1M2 and Cux may

be as much as twice the width of the base of cell Cui.

Cell 1A may be slightly open or closed and short petiolate. The postalar tuft of pile may be black or yellow.

DISTRIBUTION.—The few specimens available of A.

nitidus indicate that the species occurs in desert areas in

southern California and western Arizona. Its range

probably also includes Sonora and Baja California

(Map 11).

369-673 O—70 A

HOLOTYPE.—cfj Surprise Canyon, Panamint Mts.,

Inyo Co., California, IV-24-1957 (P. D. Hurd)

(UCAL).

ALLOTYPE.— $, 10 mi. W Blythe, Riverside Co.,

California, X-2-1954 (J. C. Hall) (UCAL).

PARATYPES.—Arizona: o*; "Palm Canal," V-29-

1955 (G. D. Butler and D. Tuttle) (ARIZ).

California: Inyo Co., cf, Surprise Canyon, Panamit

Mts., IV-24-1957 (P. D. Hurd) (NLM).

DISCUSSION.—Structure of the male genitalia indicates that the closest relative to nitidus is argentatus.

These two species seem to have no other close relatives

in North and South America. Anthrax nitidus is unique

among the species of Anthrax in North and South

America in having the integument shining black. In

this respect it closely resembles certain species of

Aphoebantus occurring in southwestern United States.

Anthrax luctuosus Macquart

Anthrax luctuosus Macquart, 1840, p. 70 [luctuosa].—Paramonov, 1947, p. 95.

Argyramoeba luctuosa.—Schiner, 1868, p. 123 [Argyromoeba].—Kert&z, 1909, p. 65.

MALE.—Integument mostly black; femora, trochanters, tibiae, metapleura, lateral margins of

abdominal terga, and abdominal sterna yellow; discs

of mesonotum, scutellum, and abdominal terga

velvety black pruinose; face, lateral margins of front,

and lower part of occiput silver pruinose; remainder of

integument grayish or brownish pruinose. Front with

fine, black setae and sparse linear black scales; face

with dense, white setae extending to antennae and a

few black setae. Occiput with short setae, white below,

black above, and a few linear, recumbent black scales;

fringe of pile on posterior margin white on lower twothirds, yellowish white on upper third. First antennal

segment about as long as wide; second segment globular, subequal in diameter to first segment and basal

part of third segment; basal part of third segment

flattened laterally, tapering gradually to styliform part;

styliform part about three times as long as style

(Figure 99).

Discs of mesonotum and scutellum with sparse, fine

black setae and linear, recumbent black scales; lateral

margins with linear, woolly scales, white anteriorly,

black posteriorly, and brown medially. Pile, scales, and

setae on mesopleuron and sternopleuron white; black

setae and hairs sometimes present. Pile on propleuron,

prosternum, and anterior margin of mesonotum white.
48

Pteropleuron with white or mixed black and white

setae and scales anteriorly, with woolly, linear, gold or

yellowish-white scales posteriorly. Fore coxa with white

setae and scales; middle and hind coxae with coarse,

white or mixed black and white setae and scales.

Postalar tuft of pile black. A tuft of white hairs behind

spiracle on metapleuron. Hypopleuron bare.

Wing (Plate 4/) hyaline apically, velvet black

basally out to a line perpendicular to wing axis running from tips of cells 1A and 2A through extreme

bases of cells R2+3 and R5 to vein Rx and slightly further apically in cells Sc and C. Cells C and Sc yellowish beyond margin of black. R-m crossvein at basal

one-fifth to one-third of vein Mi+2. Contact of cells

1M2 and Cux less than half of width of base of cell

Cui. Cell 1A closed at wing margin. Cell 2A enlarged

posteriorly, more than 1.5 times as wide as cell 1A.

Stigmatic areas in cell M pigmented. Alula well developed, posterior margin slightly convex. Calypter

pigmented, fringe of hairs light brown. Stem of halter

reddish brown, knob yellow.

Scales on femora and tibiae black. Middle femur

without distinct bristles anteroventrally; hind femur

with three to five short bristles anteroventrally. Hind

tibia with a single row of macrochaetae anterodorsally.

Lateral margins of first abdominal tergum with

dense, silvery-white pile; pile on sides of second to

fourth terga brown, black, or mixed. Posterior margin

of first tergum with a dense patch of lanceolate,

truncate, medially-directed, silvery-white scales laterally. Discs of terga two, three, and four, mesal half of

posterior margin of first tergum, and mesal half of fifth

tergum with sparse, fine black setae and linear, recumbent black scales. Lateral parts of fifth tergum and all

of terga six and seven with dense, overlapping, truncate,

posterolaterally directed, silvery-white scales. Venter

with sparse, fine white setae and linear scales.

MALE OENITALIA (Figure 54).—Gonocoxites broad,

dorsal margin attenuating abruptly to sharply rounded

tips in lateral view, mesal margins curving outward

and recurving inward to widely separated, sharply

rounded apices in ventral view; a few setae toward

the base and some scattered setae apically. Basal segment of gonostylus convex dorsally, not extending past

distal segment in ventral view; distal segment with an

acuminate ventral lobe from which the structure curves

upward and curls over dorsally. Apex of epiphallus

narrow, with an acuminate, nearly vertical dorsal spine

and ventrolateral lobes rounded apically in lateral

view; dorsal bands narrowly separated mesally. Base

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of aedeagus narrow, gradually tapering to junction with

ventral bands below approximation of dorsal bands.

FEMALE.—Similar to male. Posterolateral margins of

terga five and six with overlapping, truncate, silver

scales, and white setae; remainder of posterior terga

with black setae and scales. Sterna with black, white,

or mixed black and white setae and scales.

FEMALE GENITALIA (Figure 81).—Tenth tergum

with about 14 spines. Ventral arm of ninth tergum

narrow, parallel sided, not enlarged apically. Dorsomedial angle of sclerite on each side of gonopore extending mesally as a short, broadly rounded lobe as

wide as lateral arm; lateral arm parallel sided, slightly

enlarged at obtusely angular apex; ventral arm broadened medially parallel to meson, straight and narrow

in dorsointerior view. Spermathecal ducts about twice

as long as bulb; first and second sections short, about

equal in length; third section about as long as basal

two; second divisional collar enlarged, about twice as

wide as first; bulb globoid, symmetrical, slightly longer

than wide.

DISTRIBUTION.—Specimens of A. luctuosus have

been collected from southern Mexico through the

Amazon Basin into southeastern Brazil (Map 12).

l\ 11 V\

MAP 12.—Distribution of Anthrax luctuosus.

This is one of the few species that seems to be adapted

to areas of tropical or equatorial forest as well as desert

areas.

TYPE.—According to Painter (personal notes) the

type female of A. luctuosus is in the Paris Museum.

It is headless, mostly denuded, and has only part of one
NUMBER 43 49

hind leg present. It carries the green museum label "A.

luctuosa Macqt. Cayenne" and "16103" and pin labels

"No. 898 Anthrax luctuosa" and "128,38."

Although Macquart wrote "Patrie inconnue," in his

personal copy in the Paris Museum this has been

crossed out with "Cayenne" written in. Thus, the typelocality is probably Cayenne, French Guiana.

DISCUSSION.—Only one male has been found

among the 27 specimens of this species available for

study. A female from Pambal, Paraiba, Brazil, differs from the normal type in having the scales and

hairs on the body almost entirely light yellow.

Characters for the separation of luctuosus from

cathetodaithmos and other species in the cephus group

are given under cathetodaithmos.

Anthrax cathetodaithmos, new species

FEMALE.—Body black, legs reddish; integument

brown and velvet-black pruinose. Setae on front black;

scales lanceolate, black above, mixed black and yellowish white below. Setae on face extending almost to

antennae, yellowish white below, mixed yellowish white

and black above. Setae on occiput black, fringe of

hairs on posterior margin black. First antennal segment about as long as apical width; second segment

globular, about as wide as first segment; base of third

segment compressed laterally, narrowing abruptly to

styliform part, styliform part slightly longer than base,

about twice as long as style (Figure 99).

Scales, setae, and pile on mesonotum and scutellum

black. Mesopleuron and sternopleuron with black hairs,

setae, and scales; anterior half of pteropleuron with

mixed black and yellowish-white setae and hairs, posterior half bare; hypopleuron and metapleuron bare.

Pile on prosternum, propleuron, and anterior margin of

mesonotum black. Postalar tuft of pile black. Setae and

scales on coxae black.

Wing (Plate 4/) dark brown basally, hyaline apically,

pigment margin running perpendicularly to wing axis

from tip of vein 2A to anterior margin slightly beyond

r-m crossvein; apices of cells C and Sc not pigmented,

disc of cell 2A hyaline. Stigmatic area not pigmented.

Base of cell R4 without a spur, base of cell R2+3 with a

short spur. R-m crossvein at basal fourth of cell 1M2;

contact of cells M1+2 and Cux less than half as long as

base of cell Cux. Cell 1A as broad as cell M; cell 2A

greatly broadened, about 1.5 times wider than cell 1A

medially; alula well developed, posterior margin slightly

convex. Calypter pigmented dark brown, fringe of hair

brown. Stem of halter light brown, extreme base of

knob brown, remainder yellow.

Scales on fore and middle femora black, on hind

femur and tibia mixed black and white. Middle femur

without macrochaetae; hind femur with four weak

macrochaetae anteroventrally. Hind tibia with a single

row of macrochaetae anterodorsally.

First abdominal tergum with black pile laterally;

lateral margins of terga two through four with mixed

black and yellowish-white pile and setae, the black

predominating anteriorly; lateral margins of terga five

and six with yellowish-white pile and setae; seventh

tergum with black setae. Linear scales on posterior

margin of first tergum and on second tergum black;

scales on posterior terga black medially and anteriorly,

yellowish white posteriorly and laterally, the area of

black scales decreasing posteriorly. Setae and scales on

venter mostly white, some black setae present medially

on first tergum.

FEMALE GENITALIA (Figure 80).—Tenth tergum

with 16 spines on each side. Ventral arm of ninth

tergum broad basally, narrow apically, apex not enlarged. Dorsomedial angle of sclerite on each side of

gonopore projecting dorsomedially as a short, sharply

rounded lobe; lateral arm widened to middle, falling

obliquely to sharp ventroapical angle; ventral arm narrow, parallel sided, bending slightly inward at apex.

Ducts of spermathecae about 3 times longer than bulbs;

first section very short; second section about two-thirds

as long as third which is about 1.5 times longer than

bulb and gradually expanded apically; bulb globoid,

symmetrical, but distinctly broader than long.

MALE.—Similar to female. Setae on face entirely

black. Pile and scales on thorax entirely black. Pile

and scales on abdomen entirely black.

MALE GENITALIA (Figure 53).—Gonocoxites broad

basally, dorsal margins attenuating abruptly to sharply

rounded tips in lateral view; mesal margins curving

outward and recurving slightly inward to broadly separated, sharply rounded tips in ventral view; fine

setae along mesal margins apically, coarse setae scattered over remainder. Basal segment of gonostylus not

evident in lateral view, tip sharply rounded in ventral

view; distal segment of gonostylus with a sharp, laterally produced spine arising from a knoblike apex above

a constricted medial portion. Apex of epiphallus narrow in dorsal view, with a sharp, nearly vertical dorsal

spine; ventrolateral part rounded apically, not projecting beyond apex of aedeagus; base of aedeagus

broad, tapering to junction with ventral bands below
50

approximation of dorsal bands; dorsal bands narrowly

separated almost to dorsal spine.

VARIATION.—The pile on the face, prosternum,

propleuron, anterior margin of mesonotum, and lateral

margins of the first abdominal tergum may be entirely

black or predominantly white. The scales on the hind

tibia and femur may be entirely black or mixed black

and white. The scales on the posterior abdominal

terga may be entirely black, partly yellowish white, or

in patches of silver laterally on terga five and six. The

scales on the abdominal sterna may be black or predominantly white.

DISTRIBUTION.—Anthrax cathetodaithmos apparently occurs in mountainous areas covered with tropical forest from Peru to southwestern Mexico (Map

13). It is sympatric to luctuosus, however, which

MAP 13.—Distribution of Anthrax cathetodaithmos.

occurs eastward into the Amazon Basin and southeastern Brazil. Further collecting may show that

cathetodaithmos occurs in more rocky terrain and perhaps at greater altitude.

HOLOTYPE.— $ , 3 mi. E Colima, Colima, Mexico,

VIII-27-1962, 1100 feet (R. H. and E. M. Painter),

"hovering over Bembex nesting place near Punte el

Saldo" (RHP).

ALLOTYPE.—cf, 75 mi. E Tingo Maria, Huanuco,

Peru, X-4^1954 (E. I. Schlinger and E. S. Ross)

(CAS).

PARATYPES.—Colombia: Boyaca, 2$, Muzo, VI,

VII-1936, 900 m (J. Bequaert) (NLM, MCZ).

Panama: $ , Bella Vista, 111-23-1926 (C. T.

Greene) (AMNH); cf, "Sabanas," XI-17-1923,

F4860 (AMNH).

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Peru: Pasco, $ , Puerto Bermudez, Rio Pichis, VII18-1920 (AMNH).

DISCUSSION.—Anthrax cathetodaithmos, although

very closely related to luctuosus, may be readily distinguished by the absence of woolly, linear scales on

the posterior part of the pteropleuron. It also differs

in having the pile and scales on the anterior pleura

black rather than white, in having the setae and scales

on the anterior coxa black rather than white, in lacking silver scales laterally on the first abdominal tergum, and in having the fringe of pile on the posterior

margin of the occiput black rather than white.

The variation shown among the few specimens of

cathetodaithmos and luctuosus may indicate that additional species are included in this complex.

Anthrax luctuosus and cathetodaithmos may be

readily separated from the other species in the cephus

group by their distinctive wing pattern in which the

apical margin is perpendicular to the axis of the wing

and the apices of cells C and Sc are not pigmented.

Anthrax hylaios, new species

MALE.—Legs, coxae, lower half of pleura except

lower half of sternopleuron, underturned edges of abdominal terga, abdominal sterna, genitalia, and first

antennal segment orange, remainder of body black;

dark area of integument grayish or brownish pruinose,

margins of eyes silver pruinose. Front with black setae

and lanceolate black scales, some white scales laterally

below; face with black setae extending almost to antennae, some white hairs and setae below. Occiput

with black setae and sparse, linear black scales;

fringe of hairs on posterior margin black exteriorly

and yellow interiorly above, becoming white below.

First antennal segment about as long as apical width;

second segment globoid, somewhat flattened distally

and somewhat inserted in apex of first segment; basal

part of third segment slightly broader than second segment, flattened mesolaterally, tapering gradually to

styliform part which is about as long as base and

slightly longer than style (Figure 103).

Anterior part of mesonotum and posterior edge with

fine black setae and mixed golden-brown and black,

linear scales; posterior half of mesonotum and disc of

scutellum with linear black scales; lateral margins of

mesonotum and posterior margin of scutellum with

golden-brown scales. Sternopleuron and anterior half

of pteropleuron with fine black setae and linear yellow

scales; mesopleuron with fine black setae, white pile,
NUMBER 4 3 51

and linear scales, some golden-brown hairs posterodorsally; prosternum, propleuron, and anterior margin

of mesonotum with white pile, some black setae on

latter; postalar tuft of pile black. Fore coxa with black

setae and white, linear and lanceolate scales; middle

and hind coxae with black setae and mixed black and

white scales. Hypopleuron and metapleuron bare.

Pigment of wing (Plate 4g) dark brown, filling all

except extreme tip of cell Ri and extending from apex

of cell Ri across cell R2+3 to medial angle of vein R4;

cell R2+3 and cell R5 pigmented basally two-thirds of

distance to base of cell R4, a narrow hyaline area

along posterior margin of cell Rs; basal one-fourth of

cell 1M2 and extreme base of cell Cut pigmented; cells

1A and 2A hyaline apically for twice width of r-m

crossvein. Base of vein R4 angular; r-m crossvein located at basal third of cell 1M2, base of vein R2+3

located opposite r-m crossvein; contact of cells 2M 2

and Cuj about equal to width of base of cell Cux;

apex of cell 1A punctiform. Posterior margin of alula

rounded. Stigmatic area pigmented; calypter pigmented, fringe of hairs yellowish white. Halter light

brown, knob light brown basally and yellow apically

above, yellow below.

Fore and middle femora with light-brown scales

anterodorsally and light-yellow scales posteroventrally;

fore and middle tibiae with light-yellow scales posteriorly and light-brown scales anteriorly; scales on hind

femur and tibia mostly brown. Middle femur with one

or two macrochaetae anteroventrally; posterior femur

with an incomplete row of macrochaetae anteroventrally; anterodorsal surface of fore tibia with macrochaetae apically; hind tibia with single row of

macrochaetae anterodorsally.

First abdominal tergum with white setae laterally;

lateral margins of terga two through four with dense

black pile and erect, linear scales above with sparse

yellow hairs below; fifth, sixth, and seventh terga with

long, overlapping, lanceolate-truncate, yellowish-white

scales above and sparse yellow hairs below, some black

hairs present above on anterior part of five. Posterior

margin of first tergum and discs of other terga with

fine black setae and linear black scales, some linear

gold scales on posterior margin of terga one, two, five,

and six. Abdominal sterna with sparse black and yellow setae and sparse, linear yellow scales.

MALE GENITALIA (Figure 52).—Gonocoxites broad,

undulating ventrally in lateral view, dorsal margins

straight, tapering to sharply rounded apices; apices

with sharp lateral angles, with sharp mesal ridges and

a broad medial sulcus in ventral view; setae present

on apical third, densest in a basomedial tuft. Basal

segment of gonostylus convex dorsally in lateral

view, with a narrow, truncate basal lobe dorsally; apex

angled, with fringe of setae. Distal segment formed

upright, with broad distobasal lobe covered with dense,

coarse setae; apex with distal flattened area with sharp

tooth interiorly, and with sharp ridge basally. Apex of

epiphallus flattened, with narrow, apically truncate

medial part with sharp, basally directed medial spine

basally; lateral part curved downward laterally and

apically, apex broadly rounded in lateral view, sharply

rounded in dorsal view; apex of epiphallus connected

to aedeagus by transverse, flexible band. Dorsal bands

narrow, united medially, without setae. Aedeagus narrow basally, tapering irregularly to junction with ventral bands beyond junction of dorsal bands.

FEMALE.—Similar to male. Setae on lower half of

face mixed black and white. Base of third antennal

segment distinctly wider than second segment in

lateral view. Hairs and setae on sternopleuron and

pteropleuron mostly white. Pigment of wing filling base

of cell Cux and all of cells 1A and 2A except extreme

apices. Silver scales on abdominal terga restricted to

sides of sixth tergum. Abdominal sterna with black

setae and brown scales.

FEMALE OENITALIA (Figure 79).—Tenth tergum

with 19 spines on each side. Ventral arm of ninth

tergum enlarged apically and bent mesointeriorly at

almost a right angle. Dorsomedial angle of sclerite on

each side of gonopore curved upward and mesally as

a broad, rounded lobe; lateral arm curved upward

and rounded apically in dorsointerior view, forming a

broad plate emarginate apically and rounded laterally

in ventral view; ventral arm expanded parallel to

meson, acuminate apically. Ducts of spermathecae

about 3 times as long as bulbs; first section short, about

half as long as third section; second section about 4

times as long as third section which is about two-thirds

as long as bulb and slightly expanded at extreme apex;

bulb globoid, about 1.5 times longer than wide and

tapered to junction with duct.

VARIATION.—The specimens examined show little

variation from the holotype and allotype. The setae on

the face may be white, black, or mixed. The first

antennal segment may be orange or dark brown. The

setae on the lower pleura may be either white or

black in the male.
52

DISTRIBUTION.—Anthrax hylaios occurs in tropical

forests in the Serra do Mar of eastern Brazil, and in

lowland areas in southern Brazil and southeastern Paraguay (Map 14).

MAP 14.—Distribution of Anthrax hylaios.

HOLOTYPE.—cf, Itatiaya, Estado de Rio de Janeiro,

Brazil, X-16-1949, 700 m (W. Zikan) (IOC).

ALLOTYPE.— 9) Itatiaya, Estado de Rio de Janeiro,

Brazil, VII-3-1935, 700 m (J. F. Zikan) (IOC).

PARATYPES.—Brazil: Guanabara, $ , Rio de Janeiro,

XI-28-1933 (Dr. Zikan) (IOC); ? , Rio de Janeiro,

XI-10-1936 (Dr. Zikan) (NLM). Mato Grosso, tf,

Maracaju, VII-1937 (Servicio Febre Amarela, M.E.S.,

Brazil) (SASP). Minos Gerais, $ , Calado, Rio Doce,

11-12-15-1939 (Martins and Lopes) (IOC). Rio de

Janeiro, $ , Grajahu, VIII-20-1939 (S. Lopes)

(IOC); $ , Itatiaya, 700 m, 11-19-1945 (J. F. Zikan)

(IOC); $ , Itatiaya, 700 m, 111-18-1945 (J. F.

Zikan) (IOC); $ , Itatiaya, 700 m, II-7-1947 (J. F.

Zikan) (IOC). Santa Catarina, d, Nova Teutonia,

27° 11' S, 52°23' E, XI-29-1938 (Fritz Plaumann)

(BM). Sao Paulo, $ , Porto Cabral, Rio Parana, X-6 -

15-1941 (L. Travassos Filho) (SASP); $ , Porto

Cabral, Rio Parana, XI-1-10-1941 (L. Travassos

Filho) (NLM).

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

OTHER MATERIAL EXAMINED.—Brazil: Rio de Janeiro, $ , Itatiaya, 700 m, XI-27-1953 (J. F. Zikan).

DISCUSSION.—There appear to be no close relatives

to hylaios in either North or South America. The distinctive wing pattern will serve to separate it from

other species.

Anthrax xanthomeros, new species

FEMALE.—Body mostly black, femora, tibiae, apices

of abdominal sterna, and lateral margins of terga

orange; integument mostly brown and gray pruinose,

margins of eyes and lower two-thirds of occiput silver

pruinose. Front with black setae and lanceolate black

scales; face with white setae extending almost to antennae. Setae on occiput black on upper third, white

below; fringe of hairs on posterior margin black exteriorly and white interiorly on upper third, white below. First antennal segment about as long as apical

width; second segment lenticular; base of third segment flattened mesolaterally, distinctly wider than second segment, tapering gradually to styliform part which

is shorter than basal part and slightly longer than style.

Mesonotum with linear, rusty-brown scales laterally

and anteriorly, extending inward from in front of base

of wings, extending backward submedially from anterior margin, and extending forward in a triangle

from posterior margin; scutellum with linear rustybrown scales on posterior margin, some lanceolate yellow scales medially; remainder of mesonotum and scutellum with linear black scales. Sternopleuron with fine

white setae and linear white and yellow scales; ventral

and posterior parts of mesopleuron and anterior half

of pteropleuron with fine white setae and linear gold

scales, some coarse black setae above. Prosternum, propleuron, anterodorsal part of mesopleuron, and anterior margin of mesonotum with white pile; postalar

tuft of pile mixed yellow and brown. Metapleuron with

several white scales just above coxa; hypopleuron bare.

Fore coxa with black and white setae and linear white

scales; middle and hind coxae with black setae and

mixed gold and white linear scales.

Wing (Plate 3g) light brown basally, hyaline apically; pigment filling most of cells C and Sc (apices subhyaline) and extending out to a line running from

vein Ri just beyond base of cell R2+3 to base of cell Cux

and thence basally, filling basal two-thirds of cell 1A

and anterobasal fourth of cell 2A. Base of cell R4

angled, with rudiment of spur; r-m crossvein arising at

basal two-sevenths of cell 1M2; vein R2+3 arising slightly
NUMBER 43 53

basad of r-m crossvein; contact of cells 1M2 and Cux

about one-fourth longer than width of base of cell Cui;

cell 2A broadly open. Stigmatic area broadly pigmented. Alula not narrowed, rounded posteriorly.

Galypter lightly pigmented, fringe of hairs yellow and

brown. Halter yellow, knob brown anterobasally, yellow elsewhere.

Scales on femora and fore and middle tibiae yellow,

some black scales anteriorly at apex of fore and middle

femora and apically on hind femur; hind tibia with

black scales. Middle femur with five macrochaetae

anteroventrally; hind femur with anteroventral row

of macrochaetae incomplete basally; fore tibia with

complete row of macrochaetae anterodorsally; hind

tibia with single row of macrochaetae anterodorsally.

First abdominal tergum with yellowish-white pile

laterally; posterior half of second tergum, and third

and fourth terga with black pile laterally, anterior half

of second tergum with linear rusty-brown scales. Posterior margin of first tergum with short, lanceolate

white scales laterally and linear rusty-brown scales

medially; discs of remaining terga mostly covered with

linear black scales, posterior margins with linear rustybrown scales. Venter with fine setae, yellow basally and

black apically. First sternum, and anterior and lateral

parts of second sternum with lanceolate white scales;

middle of posterior margin of second sternum and remaining sterna with short, lanceolate-truncate, yellow

scales.

DISTRIBUTION.—Anthrax xanthomeros is known

from only the holotype, although it probably also occurs throughout northern Central America and into

southern Mexico.

HOLOTYPE.— °., Benque Vie jo, British Honduras

(Father Stanton) (MCZ). The type lacks one middle

and one hind leg and the third segment of the left

antenna. The head is covered with fungus.

DISCUSSION.—Anthrax xanthomeros has no close

relatives in North and South America. It has in

common with hylaios its orange femora, strong femoral

macrochaetae, and antennal structure, but the distinctive wing pattern of hylaios clearly sets them apart.

Species with a wing pattern similar to xanthomeros

may be distinguished readily by their reddish-brown or

black femora and by the lack of rusty-brown scales on

the margins of the mesonotum and on abdominal terga

two through seven.

Anthrax midas Fabricius

Anthrax midas Fabricius, 1805, p. 124.—Kertdsz, 1909, p. 46.

Anthrax guianicus Curran, 1934, p. 362 [guianica].

Anthrax mexicanus Cole, 1957, p. 202.

MALE.—Integument generally black; femora, tibiae,

and sometimes pleura reddish brown; eye margins

silver pruinose, remainder of body brown to gray

pruinose. Front with black setae and scales; setae on

face mostly black, a few white setae below. Occiput

with short black setae and linear scales, black above,

white below; fringe of pile on posterior margin black

above, white below. First antennal segment about 1.5

times longer than its apical width; second segment

globular; third segment flattened mesolaterally, tapering gradually from base to styliform part, styliform part

about equal to base; style about two-thirds as long as

styliform part (Figure 95).

Discs of mesonotum and scutellum with fine black

setae and linear black scales; lateral margins with

woolly black scales and coarse black setae. Sternopleuron, mesopleuron, and anterior half of pteropleuron with fine black setae and brown, white, or mixed,

linear scales and hairs. Prosternum and propleuron

with white or mixed brown and white pile; anterior

margin of mesonotum with mixed black setae and

white or brown and white pile; postalar tuft of pile

black. Anteroventral corner of hypopleuron with some

fine, black or white hairs; metapleuron sometimes with

fine brown or white hairs extending from behind

spiracle to coxa. Anterior coxa with fine, brown and

white hairs and scales, and black setae. Middle and

hind coxae with linear brown scales and black setae, a

few white scales sometimes present.

Wing (Plate 4e) brown basally, hyaline apically;

pigment fading apically. Pigment filling all of cells

C and Sc, and extending out to a line curving from

vein Ri before tip of vein Sc through bases of cells

R2+3, Rs, and Cui to tip of vein 2A. Stigmatic area

lightly pigmented. Basal angle of vein R2+3 with short

spur; basal angle of cell R4 with long spur. R-m crossvein located at basal third of cell 1M2; contact of cells

1M2 and Cux slightly longer than width of base of cell

Cui. Wing not narrowed basally; cell 2A slightly wider

than cell 1A. Calypter pigmented, fringe of hairs

brown. Alula rounded posteriorly. Halter brownish

black, knob yellow apically.

Scales on femora and tibiae entirely black, or posterior margins of fore and middle pairs with white

scales; scales on hind tibia and basitarsus sometimes
54

semierect. Middle femur with four to seven macrochaetae anteroventrally; anterior side with one or more

macrochaetae postmedially. Hind femur with a complete row of macrochaetae anteroventrally and partial

rows posteroventrally and anterodorsally.

Lateral margins of first abdominal tergum with

dense white pile, a few black setae and recumbent

white scales sometimes present posteriorly; lateral

margins of terga two through five with dense black

pile and setae. Posterior margin of first tergum and

discs of terga two through five with sparse, fine black

setae and linear black scales. Terga six and seven

and sometimes lateral margins of five with dense, overlapping, elongate, posteriorly projecting, silver scales.

Venter with sparse white scales and fine setae, white

anteriorly, black posteriorly.

MALE GENITALIA (Figure 58).—Gonocoxites long

and narrow, dorsal margins tapering gradually to

ventral margins apically; apices narrowed in ventral

view, infolded mesally to form flat lobelike structures;

setae on gonocoxites evenly distributed basally beyond

medial fold, apices bare. Basal segment of gonostylus

oblong in lateral view, with dense tuft of setae internally

at apex. Distal segment of gonostylus pyriform in ventral view, ovoid in lateral view, apex extending outward and downward; lateral and ventral surfaces

with fine setae. Apex of epiphallus flattened dorsoventrally, apex broad and bluntly angled in dorsal view,

short, sharp, recurved lobes present on each side midway between junction of dorsal bands and apex. Dorsal

bands proximate mesally, with some setae before junction. Base of aedeagus narrow, tapering to junction

with ventral bands before lateral lobes of epiphallus.

FEMALE.—Similar to male. White scales more extensive on pleura and fore and middle femora. Fringe of

hairs on calypter often white. Scales and setae on

anterior sterna white, on posterior sterna black. Pigmentation of wing slightly more extensive, reaching

base of cell 2M2. White scales on abdominal terga

restricted to sides of five and six.

FEMALE GENITALIA (Figure 76).—Tenth tergum

with 21 spines on each side. Ventral arm of ninth tergum more or less parallel basally, irregularly margined,

and slightly expanded distally. Dorsomedial angle of

sclerite on each side of gonopore produced dorsally as

short, sharply rounded lobe; lateral arm broad, parallel

sided, and truncate distally, ventral arm broad, twisted

parallel to meson, and slightly curved outward. Spermathecal ducts about as long as bulbs; first section very

short; middle section about two-thirds as long as apical

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

section which is about two-thirds as long as bulb and

broadened apically; bulb elongate tubular, about 2.5

times longer than wide, tapering basally and with a

slightly enlarged, darker apical part.

DISTRIBUTION.—Anthrax midas occurs in equatorial and tropical forests from southern Mexico to southeastern Brazil (Map 15).

MAP 15.—Distribution of Anthrax midas.

TYPES.—The holotype female of Anthrax midas

Fabricius is in the Universitets Zoologiske Museum,

Copenhagen, collection. According to Painter and

Painter (unpublished notes) it is in fairly good condition although the antennae are missing. It carries the

red type label and "A. midas, ex Am. Merid. Schmidt."

A homotype has been seen by the author.

The types of Anthrax guianicus Curran are in the

American Museum of Natural History. The holotype is

lacking its head; the allotype lacks one wing, four legs

and its abdomen, and has its head glued on. The paratype female is in excellent condition. Curran reversed

the sexes in his description.

The types of Anthrax mexicanus Cole are in the collection of the University of California at Berkeley. A

paratype male in excellent condition has been seen by

the author.

DISCUSSION.—The specimen which d'Andretta and

Carrera (1952) referred to as Anthrax leucopygus

Macquart, 1855 (not Macquart, 1840), and on which

they based their new name, macquarti, has been seen

by the author. The specimen belongs to midas rather

than leucopygus.
NUMBER 43 55

Anthrax midas differs from hyalacrus in having the

pile on the lateral margins of the first abdominal tergum white rather than black and in having the pile on

the pleura white or brown rather than black. It differs

from other species in the cephus complex in having the

wings only one-half pigmented.

Anthrax hyalacrus Wiedemann

Anthrax hyalacrus Wiedemann, 1828, p. 315.

Anthrax acroleuca.—Curran, 1934, p. 363, [not Wiedemann, 1828].

MALE.—Integument black; velvety black pruinose

on discs of mesonotum, scutellum, and abdominal

terga; silvery pruinose along margins of eyes, grayish

or brownish pruinose elsewhere. Setae and scales on

front black; face with black setae above extending to

bases of antennae, with black or mixed black and white

setae below. Setae and scales on occiput and fringe of

hairs on posterior margin black. First antenna! segment about as long as apical width, second segment

globular; third segment flattened mesolaterally, tapering gradually to styliform part; style about half as long

as styliform part (Figure 98).

Setae, pile, and scales on thorax and coxae black. A

few black hairs usually behind spiracle. Anterior corner

of hypopleuron with a small patch of hairs.

Wing (Plate 4/) velvet black basally, hyaline apically. Pigment extending out to a line running from

tip of cell Sc along vein Rx to a point about halfway between bases of cells R2+3 and R4 and thence posteriorly

through base of cell 2M2 to tip of cell 1 A. Veins in pigmented portion black, in hyaline portion light brown.

Cell 2A slightly wider than cell 1 A. Stigmatic area pigmented. Calypter pigmented, fringe of hairs black.

Halter black with light-tipped knob.

Femora and tibiae with dense, black scales; hind

femur, tibia, and tarsus sometimes with dense, overlapping, cuneate white scales. Middle femur with one or

two large postmedial bristles anteriorly and up to three

bristles anteroventrally; posterior femur with five to

seven short bristles anteroventrally.

Pile and scales on first five abdominal terga black;

pile dense laterally. Sixth and seventh terga and posterolateral margins of fifth with laterally produced,

dense, overlapping, truncate silver scales, or silver

scales restricted to posterolateral margins of sixth and

seventh terga.

MALE GENITALIA (Figure 57).—Gonocoxites very

narrow apically, tapering to acute apices in lateral

view, apices sharply rounded and separated mesally by

a distinct sulcus; setae on gonocoxites evenly distributed, extending beyond medial fold. Basal segment of

gonostylus long and thin in lateral view, with a number

of coarse setae apically; distal segment elongate, oval,

basal part bent slightly outward dorsally, with fine

setae laterally on lower two-thirds. Apex of epiphallus

flattened dorsoventrally, with a recurved, sharp lobe on

each side and a short, triangulate apical plate above

apex of aedeagus. Dorsal bands narrow, widely separated medially, without setae. Aedeagus very broad

and bulbous basally, tapering apically to junction with

ventral bands before lobes of apex of epiphallus.

FEMALE.—Similar to male. Apical margin of pigment on wing sometimes running apically along vein

R4+5 for a distance of half width of cell R5. Posterior

abdominal terga without silvery scales.

FEMALE OENITALIA (Figure 78).—Tenth tergum

with about 31 spines on each side. Ventral arm of ninth

tergum undulating, not enlarged apically. Dorsomedial

angle of sclerite on each side of gonopore not produced, right angled; lateral arm narrow, not tapering,

sharply rounded distally; ventral arm broadened parallel to meson and twisted. Spermathecal ducts about

half as long as bulbs; first section very short; middle

section about two-thirds as long as apical section which

is about one-third as long as bulb, expanded apically

and bent at almost a right angle medially; bulb elongate, tapering from apex to base, about three times

longer than maximum width and with an enlarged,

darker apical part

DISTRIBUTION.—Anthrax hyalacrus occurs in equatorial and tropical forests from the Amazon Basin north

into Panama and south into southern and eastern Brazil

(Map 16).

TYPES.—Painter (unpublished notes) found two females of Anthrax hyalacrus at the Vienna Museum,

one carrying the labels "Brasilia," "hyalacra Coll.

Winthem" and "hyalacra Wied. Brasilia" in Wiedemann's handwriting, the other carrying the labels hyalacra Coll. Wiedem." and "A. hyalacra in Amer. mer.

Brasil." Both specimens are abraded and the first lacks

most of the antennae while the second is covered with

fungus but has the left antenna complete. Because of

the condition of the syntypes, no lectotype was chosen.

A homotype compared by Painter leaves little doubt as

to the identity of this species.

BIOLOGY.—Two specimens of hyalacrus were reared

by Dr. Carl W. Rettenmeyer from nests of a Trypoxy-
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MAP 16.—Distribution of Anthrax hyalacrus.

lonidwasp (Hymenoptera: Sphecidae) on Barro Colorado Island, Canal Zone. The wasps were nesting in a

cardboard carton in a clearing at the Biological Station.

DISCUSSION.—One male from Rio Caiary-Uaupes,

Amazonas, Brazil, has the lower two-thirds of the front

covered with linear, semierect, white scales and white

setae. The setae on the face are mixed black and white.

This specimen also has the anterodorsal part of the

hind legs covered with overlapping white scales. Other

males with white scales on the hind legs are from Porto

Cabral, Sao Paulo, Brazil, and Azupizu to Miriantiriafii, Cam. del Pichis, Peru. The specimens from Sao

Paulo were collected with a male with only black scales

on the legs.

Anthrax hyalacrus may be readily distinguished from

midas by the entirely black pile on the pleura and first

abdominal segment. It differs from cephus and aterrimus in having the apical half of the wing hyaline.
NUMBER 43 57

Anthrax cephus Fabricius

Anthrax cephus Fabricius, 1805, p. 124.—Wiedemann, 1828,

p. 297.—Macquart, 1840, p. 59.—Walker 1849, p. 266.

Argyramoeba cephus.—KertSsz, 1909, p. 62.

MALE.—Integument generally black; legs, pleura,

and genitalia reddish brown; integument mostly grayish or brownish pruinose, margins of eyes and lower

part of occiput silvery pruinose. Front with black setae

and a few black scales below; face with black setae extending upward to antennae, a few white hairs and

scales below. Scales and setae on occiput black; fringe

of pile on posterior margin black above, white or mixed

brown and white below. First antennal segment about

as long as apical width; second segment globular;

base of third segment flattened mesolaterally, tapering

gradually to styliform part which is about equal to base;

style one-half to three-fourths as long as styliform part.

Mesonotum and scutellum with fine black setae and

linear black scales, sometimes brown or white scales in

front of base of wings and white scales rarely present

at apex of scutellum. Sternopleuron, mesopleuron, and

anterior part of pteropleuron with fine black setae and

black and/or white, threadlike scales; upper half of

mesopleuron and pteropleuron with some coarse black

setae and black and/or white pile. Pile on prosternum,

propleuron, and anterior margin of mesonotum black,

white, or mixed; postalar tuft of pile black. A few

black or white hairs behind spiracle; anteroventral

corner of hypopleuron with a few black or white hairs

or threadlike scales. Fore coxa with black setae and

black and/or white pile and scales; middle and hind

coxae with black setae and black or mixed black and

white scales.

Wing (Plate 4c) entirely black pigmented or subhyaline apically. Stigmatic area lightly pigmented;

calypter pigmented, fringe of hairs brown. R-m crossvein located one-third or two-fifths of way from origin

to bifurcation of vein Mi+2; contact of cells 1M2 and

Cui one to two times as long as base of cell Cu^ Vein

R4 with long spur at base. Cell 2A as wide as cell 1A.

Scales on legs black. Middle femur with one to four

macrochaetae anteroventrally and one to several macrochaetae postmedially on anterior side; hind femur

with four to seven macrochaetae anteroventrally. Hind

tibia with double row of macrochaetae anterodorsally;

scales mostly recumbent.

Pile on lateral margins of first abdominal tergum

white, sometimes a few black hairs posteriorly; lateral

margins of terga two through four, and sometimes

five and six with dense black pile. Posterior margins

of first tergum, discs of terga two through four and

sometimes five and six with sparse black setae and

linear black scales. Seventh tergum and sometimes all

or lateral parts of terga five and six with dense, overlapping, posteriorly-produced, silver-white scales. Setae

and scales on sterna entirely black, or white (or yellow) anteriorly and black posteriorly.

MALE GENITALIA (Figure 59).—Gonocoxites long

and narrow, narrowing abruptly before very narrow

tips in lateral view, apices infolded mesally, forming

narrow, acuminate, incurved lobes in ventral view,

setae evenly distributed basally beyond medial fold,

extreme apices bare. Basal segment of gonostylus oblong, with a few setae apically; distal segment bulbous

in ventral view, with a diagonally truncate tip, dorsal

margin tapering to ventral margin in lateral view; fine

setae present on basal part. Apex of epiphallus flattened

dorsoventrally, apex rounded, sharp, recurved lateral

lobes present preapically. Dorsal bands narrow, proximate medially, with a few setae before junction. Base

of aedeagus narrow, but bulbous, tapering to junction

with ventral bands after junction of dorsal bands, with

enlarged area before junction of dorsal bands.

FEMALE.—Similar to male. White pile and scales

sometimes more extensive on pleura. Fore and middle

femora and tibiae sometimes with white scales posteriorly. Hind femur sometimes with complete row of

macrochaetae anteroventrally. Apex of wing sometimes hyaline as far back as a line running from tip

of subcosta to tip of M2 in specimens from Central

America. Silver scales on apex of abdomen restricted

to lateral parts of fifth tergum and sometimes to the

sixth and seventh.

FEMALE GENITALIA (Figure 77).—Tenth tergum

with about 21 spines on each side. Ventral arm of

ninth tergum undulate, not expanded apically. Dorscmedial angle of sclerite on each side of gonopore produced dorsally as a short, rounded lobe about as long

as broad; lateral arm short, broad, parallel sided and

truncate apically; ventral arm broadened medially

parallel to meson, narrowing below and slightly curved

toward meson. Spermathecal ducts about as long as

bulbs; first section very short; second section slightly

shorter than third which is slightly expanded and bent

at a right angle apically; bulb elongate elliptical, broadest submedially, slightly constricted postmedially, without differentiated distal section.
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—Anthrax cephus occurs in tropical

forests from southeastern Brazil through the Amazon

Basin (Map 17).

TYPE.—According to notes supplied by R. H. and

E. M. Painter, the type male of Anthrax cephus is in

the Copenhagen Museum. Tips of the wings are

broken off, the antennae are missing, and the body is

greasy. It carries the label "A. cephus ex Am. Mer.

Schmidt." The description of the type agrees with

typical specimens of A. cephus. Fabricius gave "America Meridionali" as the type-locality.

DISCUSSION.—Two specimens from Chauguinois

District, Panama, and La Suiza, Costa Rica, differ

from specimens from Brazil in having a vertically defined hyaline area at the apex of the wing. Since

specimens from northern and western Brazil tend to

have the tip of the wing hyaline or subhyaline, it seems

probable that the Central American specimens rep17.—Distribution of Anthrax cephus.
NUMBER 4 3 59

resent only an extreme manifestation of this characteristic and that they are the same species. Since the

tops of the wings of the type are broken off, it is not

possible to ascertain whether it showed this condition.

A specimen from Porto Cabral, Rio Parana, Sao

Paulo, Brazil, differs from other specimens in the same

series in having the tip of the wing hyaline and areas

in the preapical and posterior cells subhyaline. It seems

probable that this specimen is a hybrid between cephus

and midas, since a specimen of the latter was collected

at the same locality.

For characters separating cephus from aterrimus

see the latter species. Anthrax cephus differs from

other species in the cephus complex in having the wing

entirely pigmented or with only a small area apically

hyaline, rather than with the apicoposterior half

hyaline.

Anthrax aterrimus (Bigot)

Argyramoeba aterrima Bigot, 1892, p. 349 [Argyromoeba].

Anthrax aterrimus.—Painter and Painter, 1962, p. 72.—

Krombein, 1967, p. 400.

Argyramoeba cephus.—Osten Sacken, 1877, p. 243.—Coquillett, 1894, p. 95.—Johnson, 1895, p. 325 [part] [not Fabricius, 1805; misidentification].

Spongostylum cephus.—Aldrich, 1905, p. 222 [Spogostylum]

[not Fabricius, 1805; misidentification].

Spongostylum slossonae Johnson, 1913, 55.—Cole, Malloch,

and McAtee, 1924, p. 186 [Spogostylum].

Anthrax slossonae.—Brimley, 1938, p. 341.

MALE.—Integument entirely black or legs and genitalia reddish brown; integument generally grayish and

brownish pruinose, margins of eyes and lower part of

occiput silvery pruinose. Setae and scales on front

black; face with black setae extending up to antennae,

few white setae along oral margin. Setae on occiput

black, scales black above, white below; fringe of pile

on posterior margin black exteriorly and white interiorly above, white below. First antennal segment

slightly longer than apical width; second segment

globular; base of third segment flattened mesolaterally,

tapering gradually to styliform part which is about

equal to base in length; style one-half to three-fourths

as long as styliform part of third segment (Figure 96).

Setae and scales on mesonotum and scutellum mostly

black; a few white or brown, threadlike scales in front

of bases of wings and apically on scutellum. Sternopleuron, mesopleuron, and anterior half of pteropleuron with fine black setae and threadlike white

scales; some white pile and some brown and/or black

scales, setae, and hairs dorsally on mesopleuron and

pteropleuron. Pile on prosternum, propleuron, and

anterior margin of mesonotum white or with some

brown and black hairs intermixed; postalar tuft of pile

black with some hairlike white scales basally. A few

white hairs behind spiracle; a few black setae and hairlike white scales anteroventrally on hypopleuron. Fore

coxa with white scales and hairs and black setae; middle and hind coxae with mixed black and white scales

and black setae.

Wing (Plate 4rf) entirely pigmented with brown

or black, discs of apical cells sometimes subhyaline,

apical part of wing rarely entirely subhyaline. Stigmatic area lightly pigmented, calypter lightly pigmented, fringe of hairs brown. R-m crossvein located

one-third to two-fifths of way from base to bifurcation

of vein Mi+2; contact of cells 1M2 and Cui slightly

shorter to slightly longer than base of cell Cux. Vein R*

with long spur at its basal angle.

Scales on legs entirely black; scales on hind tibia often

semirecumbent. Middle femur with complete row of

macrochaetae anteroventrally and one to several postmedial bristles on anterior side; posterior femur with

complete rows of macrochaetae anteroventrally, posteroventrally, and anterodorsally. Posterior tibia with

double row of macrochaetae anterodorsally.

Pile on lateral margins of first abdominal tergum

white; terga two through four with dense black pile

laterally, a few white hairs posteriorly on fourth tergum. Posterior margin of first tergum, discs of terga

two through four, and medial part of five and sometimes six with sparse black setae and linear black scales.

Remainder of posterior terga with dense, overlapping,

posteriorly produced, silvery-white scales. Hairs and

scales on venter sparse, black, white, or mixed, the

white usually predominating anteriorly, the black

posteriorly.

MALE GENITALIA (Figure 56).—Gonocoxites long,

apical part narrow, dorsal margins tapering to ventral

margins apically; apices rounded and slightly curved

mesally in ventral view, folded inward mesally to form

deep mesal sulcus; long setae covering entire surface

almost to base. Basal segment of gonostylus oblong in

lateral view, extending to base of distal segment, with

dense tuft of setae apically; distal segment small, ovoid

in ventral view, with short, truncate process extending outward near the apex and twisted apically, entire

segment except styliform part with fine short setae.

Apex of epiphallus flattened dorsoventrally, apex

bluntly angled in dorsal view; a short, sharp recurved
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lobe present on each side midway between junction of

dorsal bands and apex; dorsal bands proximate mesally,

with some setae before junction. Base of aedeagus small

and narrow, almost tubular, tapering slightly to junction with ventral bands below lateral lobes of

epiphallus.

FEMALE.—Similar to male. Pile and scales on thoracic

pleura usually more predominantly white. Fore and

middle femora and tibiae usually with white scales

posteriorly; hind femur sometimes with a few white

scales posteriorly at base. White scales and hairs on

abdominal venter usually restricted to first three sterna.

Silvery-white scales apically on abdominal terga

restricted to posterolateral margins of five and six.

FEMALE GENITALIA (Figure 75).—Tenth tergum

with about 27 spines on each side. Ventral arm of ninth

tergum enlarged apically, irregularly margined.

Dorsomedial corner of sclerite on each side of gonopore

greatly produced dorsally as a rounded lobe; lateral

arm short, broad and emarginate apically, extending

posterolaterally; ventral arm broad evenly margined

and twisted mesally on lower half. Each duct of

spermathecae greatly elongated, about 6 times longer

than bulb; first section very short; second section about

two-thirds as long as apical section which is about

3.5 times longer than bulb and gradually expanded

from base to apex; bulb slightly longer than wide,

rounded, not well defined from duct.

DISTRIBUTION.—Anthrax aterrimus occurs in forested areas in southeastern United States as far north

as Connecticut and the southern tip of Illinois, and as

far west as central Texas (Map 18).

TYPES.—According to Painter and Painter (1962)

the type of Argyramoeba aterrima, a female rather than

a male as stated by Bigot, is in the Bigot collection of

the British Museum. It is heavily draped with fungus

but otherwise in good condition. It carries the label

"Argyromoeba aterrima cf n. sp. Inedit. Quincy Juin

1886 J. Bigot Baltimore." Bigot's description agrees

well with typical specimens of the species, including

the paratype series of slossonae (Johnson).

The types of Spongostylum slossonae are in the

Museum of Comparative Zoology at Harvard University. The entire series from which the species was

described, with the exception of the types, has been

examined by the author. The type-locality is Cumberland Gap, Kentucky.

BIOLOGY.—Anthrax aterrimus was recorded (as

cephus Fabricius) as being "bred from the nest of a

Mud-wasp in Texas, forming tubes of clay five or six

MAP 18.—Distribution of Anthrax aterrimus.

inches long, pasted together like organ-pipes," very

probably Trypoxylon politum Say (Hymenoptera:

Sphecidae), by Osten Sacken (1877). An additional

specimen in the collection of R. H. Painter carries the

label "bred from mud dauber nest." Krombein (1967)

reared this species from nests of Trypargilum striatum

(Provancher), T. collinum rubrocinctum (Packard),

T. clavatum, Isodontia auripes (Fernald) (Hymenoptera: Sphecidae), Euodynerus megaera (Lepeletier), Monobia quadridens (Linnaeus), and Ancistrocerus spinolae (Saussure) (Hymenoptera: Vespidae).

DISCUSSION.—Anthrax aterrimus often has been

confused with cephus, a South American species to

which it is closely related. Specimens of cephus usually

are darker, without subhyaline areas in the posterior

cells of the wing, and with numerous black hairs and

scales on the mesopleuron and sternopleuron. There is

much variation, however, especially among specimens

of cephus, so that the only certain diagnostic character

is the male genitalia. In aterrimus the distal segment

of the gonostylus is ovoid in ventral view with a short,

twisted process extending outward near the apex (Figure 56). In cephus this structure is slightly larger,

bulbous basally in ventral view, with a longer styliform process with a diagonally truncate tip (Figure

59). Anthrax aterrimus may be separated readily from

other members of the cephus complex by the presence

of pigment in the outer part of wing.

Anthrax aterrimus has also been confused with
NUMBER 4 3 61

analis, f. grossbecki. It may be distinguished readily

by the presence of a small patch of hairs at the anterior

corner of the hypopleuron and by the presence of a

postmedial bristle on the anterior side of the middle

femur.

Anthrax pluricellus Williston

Anthrax pluricellus Williston, 1901, p. 277 [pluricella].—

Kertesz, 1909, p. 51 [pluricella].—Painter and Painter,

1962, p. 79.

Exoprosopa capucina.—Rau, 1940, p. 594 [not Fabricius,

1805, misidentification].

MALE.—Body mostly black, tibiae and femora yellow; integument blue gray and brown pruinose; eye

margins, face, and lower half of occiput white pruinose.

Front with black setae and linear black and yellow

scales; fringe of hairs on posterior margin black exteriorly, white interiorly above, white below. First

antennal segment about as long as apical width; second segment globoid, slightly flattened apically; base

of third segment flattened mesolaterally, narrower than

second segment in lateral view, tapering abruptly to

styliform part which is slightly longer than base and

about 1.5 times as long as style (Figure 105).

Discs of mesonotum and scutellum with linear black

scales; rust-brown scales broadly over anterior and

lateral margins and in triangular patch posteriorly on

mesonotum, and anteriorly and laterally on scutellum.

Sternopleuron, lower half of mesopleuron, and anterior

half of pteropleuron with black and gold setae, white

and yellow scales, and sometimes a few black scales.

Prosternum, propleuron, dorsal half of mesopleuron,

and anterior margin of mesonotum with white pile,

some black setae and hairs on last two areas. Postalar

tuft of pile black, some white or yellow scales often

present basally. Fore coxa with white bristles and gold

scales. Hypopleuron bare; metapleuron with a few

hairlike white scales behind spiracle.

Wing (Plate 4a) light brown with hyaline areas at

apex and extending inward from posterior margin,

pigment filling cells C, Sc, and R1} extending beyond

base of cell R4 in cells R2+3 and R5, filling base of cell

R*, and extending from cell R5, broadly across base

of cell Mx and apex of cell 1M2 onto spurious vein

in cell 2M2; extreme base of cell 2M2 and basal

fourth of cell Cui pigmented; extreme apex of

cell M and apical spot in cell 2A hyaline. Vein R4

angled basally, with a short spur; r-m crossvein at basal

two-fifths of cell 1M2, vein R2+3 arising slightly basad;

contact of cells 1M2 and Cui slightly shorter than width

of base of cell Cui; cell 1A punctiform apically; cell 2A

not reduced, wider than cell 1A postmedially; alula

reduced, posterior margin straight; a spurious vein

extending from apical angle of m crossvein to posterior

margin. Stigmatic area lightly pigmented. Calypter

lightly pigmented, fringe of hairs white. Halter light

brown, apical margin of knob yellow, base dark brown.

Bases of legs mostly with light-yellow scales, some

black scales dorsally on femora at apex and dorsally

on hind tibia. Middle femur with two or three macrochaetae anteroventrally toward apex; anterodorsal side

of fore femur with a few weak macrochaetae; hind

tibia with single row of macrochaetae anterodorsally.

Sides of first abdominal tergum with white pile, a

large or small area of light or dark brown pile

anteriorly; lateral margins of terga two through four

with black pile, setae, and scales, numerous light brown

hairs mixed with scales anteriorly on second tergum

and posteriorly on terga two through four. Posterior

margin of first tergum with sparse black or rusty-brown

linear scales; discs of terga two through four and central half of five with linear black scales; rusty-brown

scales present anteriorly on two and posteriorly on two

through five; centers of terga six and seven with black

scales anteriorly and rust-brown scales posteriorly. Sides

of terga five through seven with patches of elongate,

posteriorly produced, overlapping, lanceolate-truncate,

white scales and with tufts of black and yellow setae

and pile. Venter with fine white and yellow setae, and

linear white and yellow scales, darkest posteriorly.

MALE GENITALIA (Figure 60).—Gonocoxites broad,

dorsal margins broadly rounded to truncate apically

in lateral view, apices broadly rounded in ventral view,

medial sulcus not defined, setae sparsely distributed

ventrally and along dorsal margin. Basal segment of

gonostylus convex above in lateral view, with a few

fine setae apically. Distal segment of gonostylus oval

basally, somewhat flattened dorsoventrally with a broad

mesal lobe in ventral view; styliform part curved outward, apex flattened apically; fine setae present dorsally and laterally at base. Apex of epiphallus broad,

somewhat flattened dorsoventrally; dorsal part with a

raised, basally projecting central lobe and truncate

apex; lateral part with long, flat lobes projecting basolaterally. Dorsal bands meeting mesally, but not

coalesced, with numerous short setae. Base of aedeagus

large and bulbous, dorsal margin curving downward

to junction with ventral bands below approximation

of dorsal bands.
62

FEMALE.—Similar to male. White scales on posterior

abdominal terga restricted to lateral margins of fifth

segment.

FEMALE GENITALIA (Figure 83).—Tenth tergum

with about 16 spines on each side. Ventral arm of

ninth tergum slightly enlarged and undulate distally.

Dorsomedial corner of sclerite on each side of gonopore

produced dorsally as a dactylate lobe; lateral arm

slightly broader than dorsal lobe and rounded apically;

ventral arm narrow, tapering to apex which is evenly

curved mesad. Each duct of spermathecae about as

long as bulb; first segment very short; second segment

about as long as third which is bent at about a right

angle before expanding apically to bulb; bulb about 2.5

times longer than maximum width, tapering to junction

with duct and with dark, slightly enlarged apical

section.

DISTRIBUTION.—The specimens referred to by Rau

(1940) 2&Exoprosopa capucina (Fabricius) have been

studied by the author and are almost certainly A.

pluricellus, since they agree closely with the description of the type given by Painter and Painter. These

specimens were collected 40 km south of Ciudad Victoria, Tamulipas, Mexico. The type-locality is "Atoyac

in Vera Cruz," Mexico. The species probably occurs

all along the eastern coast of Mexico east of the Sierra

Madre Oriental and perhaps inland on the central

plateau.

TYPE.—Painter and Painter (1962) state that the

type of Anthrax pluricellus, a female rather than a

male as Williston indicated, is in the British Museum.

It is in good condition except that the hind legs, one

middle leg, and the left antenna are missing. Williston

(1901) placed this species in Anthrax (auctt.) (Villa

Lioy) since he believed that the head, which had been

glued on, did not belong to the body.

BIOLOGY.—The specimens referred to by Rau were

reared from the nests of Chalybion zimmermanni'Dahlbom (Hymenoptera: Sphecidae).

DISCUSSION.—Anthrax pluricellus, snowi, and

costaricensis differ from other species in having cell

2M2 divided by a spurious vein extending from the

m crossvein to the wing margin. Only a single series

of each species has been collected (with the exception

of the type of pluricellus) and each appears to be

distinct, although subsequent collecting may show that

the taxa are only subspecies or variants of a polymorphic species. Because the taxa differ in several unrelated characters, and there is no significant variation

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

within the series available, they are regarded here as

distinct species.

Anthrax pluricellus differs from both costaricensis

and snowi in having yellow rather than black femora

and in having rusty-brown scales on the mesonotum,

scutellum, and abdominal terga.

Anthrax snowi, new species

MALE.—Body mostly black, legs reddish black, integument blue green and brown pruinose; eye margins,

face, and lower part of occiput white pruinose. Front

with black setae and sparse black scales, a few white

scales laterally below; face with mixed black and

white setae extending almost to antennae. Occiput with

black setae and a few black scales; fringe of hairs on

posterior margin black exteriorly, white interiorly. First

antennal segment about as long as apical width; second

segment globoid, flattened apically; base of third segment flattened mesolaterally, about as wide as second

segment in lateral view, tapering sharply to styliform

part which is slightly longer than base and about 1.5

times longer than style (Figure 104).

Discs of mesonotum and scutellum with linear black

scales, some white scales laterally in front of base of

wings and along posterior margin of scutellum.

Sternopleuron with white setae and hairlike scales,

lower part of mesopleuron and anterior part of

pteropleuron with black setae and hairlike white scales.

Prosternum, propleuron, dorsal part of mesopleuron,

and anterior margin of mesonotum with white pile;

some black hairs and numerous black setae on last two.

Fore coxa with white bristles and white scales, some

black bristles apically; middle and hind coxae with

black bristles and black and white scales. Hypopleuron

and metapleuron bare.

Wing (Plate 4a) light brown with hyaline areas at

apex and extending inward from posterior margin;

pigment filling cells C, Sc, and R1} extending beyond

base of cell R4, in cells R2+3 and R5, filling base of cell

R4, extending from cell R5 broadly across base of cell

Mi and apex of cell 1M2 onto spurious vein in cell 2M2,

but not quite reaching posterior margin; base of cell

1M2 out to base of cell 2M2 and extreme anterior margin pigmented; extreme base of cell 2M2 and basal

fourth of cell Cu! pigmented; extreme apex of cell M

and large apical spot in cell 2A hyaline. Vein R4

angled basally, with a short spur; r-m crossvein at

basal two-fifths of cell 1M2, vein R2+3 arising slightly

basad; contact of cells 1M2 and Cux slightly shorter
NUMBER 4 3 63

than width of base of cell Cux; cell 1A punctiform

apically; cell 2 A not reduced, wider than cell 1A postmedially; alula reduced, posterior margin straight; a

spurious vein extending from apical angle of m crossvein to posterior margin of wing. Stigmatic area lightly

pigmented. Calypter lightly pigmented, fringe of hairs

white. Halter light brown, apical margin of knob yellow, base dark brown.

Scales on fore and middle femora black anteriorly

and mixed yellow and white posteriorly; hind femur

with mixed yellow and white scales on basal half posteriorly; hind femur with mixed yellow and white scales

on basal half posteriorly and extreme base anteriorly,

remainder with black scales. Fore and middle tibiae

with black scales anteriorly and white scales posteriorly;

hind tibia with black scales. Middle femur with two or

three macrochaetae anteroventrally toward apex; anteroventral row of macrochaetae on hind femur

incomplete basally; anterodorsal surface of fore tibia

with a few weak macrochaetae apically; hind tibia with

a single row of macrochaetae anterodorsally.

Sides of first abdominal tergum with white pile, a

large area of light-brown pile anteriorly; lateral margins of terga two through four with black pile, setae,

and scales. Posterior margin of first tergum with

sparse, black linear scales; discs of terga two through

four and central half of five with linear black scales;

lateral fourth of fifth tergum, and sixth and seventh

with elongate, posteriorly produced, overlapping,

lanceolate-truncate, white scales. Venter with fine

white and yellow setae, and linear white and yellow

scales, darkest posteriorly.

MALE GENITALIA (Figure 61).—Gonocoxites broad,

dorsal margins tapering convexly to blunt ventral

angles in lateral view; apices broadly rounded in ventral view, inner margins rounded to broad, shallow

mesal sulcus; setae evenly distributed, extending almost

to base mesally. Basal segment of gonostylus sinuate

dorsally in lateral view, acutely angled posteriorly, with

short, rounded lobes covered with fine setae apically

in ventral view. Distal segment of gonostylus broad

basally with a short mesal projection, tapering to laterally curved styliform part; apex with oval flattened

area; basal part with fine setae. Apex of epiphallus

flattened dorsoventrally, projecting upward at a low

angle; medial part broadly rounded apically in dorsal

view, with a short, rounded projection basally, with

shallow longitudinal sulci between medial and lateral

parts; lateral parts rounded apically and curved downward, extending basally to middle of medial part.

369-673 O—70—6

Dorsal bands narrow, united mesally, without setae.

Base of aedeagus broad, bulbous, tapering dorsally to

junction with ventral bands below junction of dorsal

bands; ventral bands narrow, at a right angle

preapically.

VARIATION.—The paratype male, the only other

specimen available, is similar to the holotype.

DISTRIBUTION.—This species is known only from

the type-locality.

HOLOTYPE.— $ , Oak Creek Canyon, Cococino Co.,

Arizona, August, 6000 feet (F. H. Snow) (KANS).

PARATYPE.—cT, Oak Creek Canyon, Cococino Co.,

Arizona,—July, 6000 feet (F. H. Snow) (RHP).

DISCUSSION.—Anthrax snowi may be distinguished

from pluricellus by the reddish brown rather than

yellow femora and the absence of rust-brown scales

on the dorsal surface of the thorax and abdomen. It

may be distinguished from costaricensis by the entirely

pigmented apex of cell Rx and the hyaline apices of

cells 1A and 2 A as well as by the lighter pigmentation

of the wings.

Anthrax costaricensis, new species

FEMALE.—Body black, legs dark red; integument

mostly gray and brown pruinose. Front with black setae

and linear scales; face with mixed black and yellow

setae. Occiput with black setae above and white setae

below; fringe of hairs on posterior margin black exteriorly and white interiorly above, white below. First

antennal segment about as long as apical width; second segment globoid, apical margin flattened; base of

third segment flattened mesolaterally, slightly wider

than second segment in lateral view, tapering abruptly

to styliform part which is slightly longer than base

and about 1.5 times longer than style (Figure 106).

Discs and margins of mesonotum and scutellum with

linear black scales. Sternopleuron, lower half of mesopleuron, and anterior half of pteropleuron with fine

black setae and linear white scales, some white setae

on sternopleuron. Prosternum with white pile; propleuron and anterior margin of mesonotum with white

and brown pile, numerous black setae on latter; upper

half of mesopleuron with white pile and black setae.

Postalar tuft of pile black. Fore coxa with white bristles

and scales, some black bristles apically; middle and

hind coxae with black bristles, and black and white

scales. Hypopleuron bare, metapleuron with a few

white scales behind spiracle.
64

Wing (Plate 4b) dark brown basally and anteriorly, and in a preapical band extending toward posterior margin; cells C and Sc filled, cell Ri filled beyond

apex of vein Sc and along anterior margin to apex;

cells R2+3 and R5 filled beyond base of cell R4, cell R*

filled; pigment extending from cell R5 broadly across

apex of cell 1M2 and base of cell Mx almost to posterior

margin on vein M2 and spurious vein on m crossvein;

base of cell 1M2 out to base of cell 2M2 and along

extreme anterior margin, extreme base of cell 2M2

and basal fourth of cell Cui pigmented; cells 1A and 2A

completely pigmented. Base of cell R4 angled, with

short spur; r-m crossvein at basal two-fifths of cell

1M2 vein R2+3 arising opposite; contact of cells 1M2

and Cui slightly longer than width of base of cell Cui;

cell 1A punctiform apically; cell 2A not reduced, wider

than cell 1A postmedially; alula reduced, posterior

margin concave; a spurious vein extending from apical

angle of m crossvein to margin of wing. Stigmatic area

lightly pigmented. Calypter lightly pigmented, fringe

of hairs white. Stem of halter light brown; knob dark

brown basally, yellow at extreme apex.

Femora and hind tibia with black scales; a few white

scales posteriorly on fore and middle femora; fore and

middle tibiae with black scales anteriorly and white

scales posteriorly. Middle femur with three or four

small macrochaetae anteroventrally toward apex;

anteroventral row of macrochaetae on hind femur

incomplete basally; fore tibia with one or two macrochaetae anterodorsally; anterodorsal row of macrochaetae on hind tibia with double row of macrochaetae.

Lateral margins of first abdominal tergum with a

large patch of black pile anteriorly surrounded by white

pile; terga'two through four with black pile and setae

laterally. Posterior margin of first tergum with a few

black scales; discs of terga two through seven with

linear black scales, patch of lanceolate-truncate white

scales laterally on five. Sterna one and two with fine

white setae and linear white scales, a few black setae

laterally on two; remaining sterna with black setae

and scales.

FEMALE GENITALIA (Figure 82).—Tenth tergum

with about 15 spines on each side. Ventral arm of ninth

tergum undulating, expanded parallel to meson apically. Dorsomedial angle of sclerite on each side of

gonopore elongated dorsomedially as a narrow, sharply

rounded lobe; lateral arm short, narrow in dorsointerior view, curled under below; ventral arm narrow,

parallel sided, almost straight. Each spermathecal duct

about two-thirds as long as bulb; basal section very

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

short; middle section about as long as distal section

which is curved and expands distally to bulb; bulb

elongate tubular, about four times longer than wide,

with a lighter, slightly broader apical section about

one-third of entire length.

VARIATION.—The two female para types do not vary

appreciably from the type. The anterodorsal row of

macrochaetae on the hind tibia is single in the

paratypes.

DISTRIBUTION.—Anthrax costaricensis is known only

from the type-locality, although it probably also occurs

throughout much of Central America.

HOLOTYPE.— ° , La Suiza, Costa Rica, III—3 (F.

Schild) (A. L. Melander collection) (USNM).

PARATYPES.—Costa Rica: 2 ? , La Suiza, VII-18,

19 (F. Schild) (A. L. Melander coll.) (USNM,

NLM).

DISCUSSION.—Anthrax costaricensis differs from

pluricellus in having the femora black rather than yellow and in lacking rust-brown scales on the dorsal surface of the body. It differs from both pluricellus and

snowi in having the wings pigmented with dark rather

than light brown, in having the posterior edge of cell

Ri hyaline and in having cells 1A and 2A entirely pigmented rather than with subapical or apical hyaline

areas.

Anthrax laticellus, new species

MALE.—Body black, femora and tibiae red; integument blue gray and brown pruinose. Front with

black setae and linear white scales; face with black

setae extending almost to antennae. Occiput with black

setae and scales on upper three-fifths, with some white

scales below; fringe of hairs on posterior margin black

exteriorly, white interiorly on upper two-thirds, white

below. First antennal segment shorter than apical

width; second segment lenticular; base of third segment flattened mesolaterally, about as wide as second

segment in lateral view, narrowing abruptly to styliform part which is slightly longer than base and about

1.5 times longer than style (Figure 120).

Discs and lateral margins of mesonotum and scutellum with linear black scales, a few white scales in front

of base of wing. Sternopleuron, lower half of mesopleuron, and anterior half of pteropleuron with white

setae, and few black setae present, especially on pteropleuron. Prosternum, propleuron, upper half of mesopleuron, and anterior margin of mesonotum with white

pile, some white setae on mesopleuron and numerous
NUMBER 43 65

black setae on anterior margin of mesonotum. Postalar

tuft of pile black. Coxae with black setae and black

scales, some white scales on anterior pair. Hypopleuron

and metapleuron bare.

Wing (Plate 5/) light brown basally and anteriorly,

hyaline posteriorly. Pigment filling cells C and Sc, cell

Ri to base of cell R4, and extending along anterior

margin almost to apex, a short band extending across

cell R2+3 to base of cell R4; cell R2+3 filled two-thirds of

way to base of cell R4, a broad triangular band extending across cell R5 halfway to base of cell R4; extreme

bases of cells R5 and Cux and anterobasal corner of cell

1M2 pigmented, a small spot present at base of cell

2M2; cell R entirely filled; margin of pigment extending from base of cell Cux to posterior margin at extreme

base of cell 2A. Cell R4 angled at base, with a short

basal spur; r-m crossvein at basal third of cell 1M2,

vein R2+3 arising slightly apicad; contact of cells 1M2

and Cui about 1.5 times longer than base of cell Cux;

cell 1A punctiform apically. Cell 2A not reduced,

wider than cell 1A postmedially; alula vestigial, posterior margin straight. Stigmatic area lightly pigmented. Calypter lightly pigmented, fringe of hairs

brown. Halter light brown, knob dark brown.

Legs mostly with black scales, some white scales posteriorly on fore and middle femora and tibiae. Middle

femur with one anteroventral bristle; row of macrochaetae on hind femur incomplete basally. Anterodorsal surface of fore tibia with a few weak

macrochaetae anterodorsally; hind tibia with a single

row of macrochaetae anterodorsally.

First abdominal tergum with white pile laterally;

lateral margins of terga two through four with black

pile and linear black scales. Posterior margin of first

tergum with dense, linear, white scales. Discs of terga

two through four and a medial spot on five with linear

black scales; remainder of fifth tergum and sixth and

seventh with dense, posteriorly produced, lanceolate,

truncate, overlapping, white scales. Venter with white

and yellow setae anteriorly and black setae posteriorly,

a few white scales anteriorly and a few black scales

posteriorly.

MALE GENITALIA (Figure 72).—Gonocoxites broad

basally, tapering apically; apices with flat digitate areas

ventrally defined by sharp lateral and mesal carinae.

Basal segment of gonostylus rounded dorsally and acute

apically in lateral view, extending little past the bases

of the distal segments in ventral view. Distal segment

of gonostylus triangular basally in ventral view, cylindrical and gradually tapering in lateral view, extending dorsolaterally with a preapical dorsolateral blunt

tooth, and with the distal part angled sharply outward

forming a sharp tooth and a flat apical surface; setae

fine, on ventral and interior surfaces toward base. Dorsal part of apex of epiphallus with a medial sharp

tooth extending upward and backward at a 60° angle,

curving downward laterally, apex deeply emarginate

in dorsal view; ventrolateral part bluntly rounded

apically in lateral view. Dorsal bands narrow, not

united mesally toward apex, without setae. Base of

aedeagus broad but not bulbous, tapering gradually

to junction with ventral bands beyond dorsal bands in

lateral view.

VARIATION.—The only additional specimen of this

species, a male, is similar in almost every respect to the

holotype. The contact of cells 1M2 and Cui is slightly

shorter than the width of cell Cu^

DISTRIBUTION.—Anthrax laticellus is known only

from the holotype and paratype. It may occur throughout the forested areas of northern and eastern Arizona.

HOLOTYPE.—<$, "Gr. Can." (Grand Canyon?),

VII-11 (Adams collection) (RHP).

PARATYPE.—Arizona: Cochise Co., d, Texas Pass,

Dragoon Mts., VII-21-1917 (C.U.Expedit.) (NLM).

DISCUSSION.—The holotype is in rather poor condition. The left antenna is missing, and the front and

face are covered with dirt. Part of the right wing is

missing. The paratype lacks the apical segment of

both antennae and five legs. Despite the poor condition of the types, this species is being described in order to make this revision as complete as possible.

Anthrax laticellus differs from other members of the

argyropygus complex in having cell 2A of the wings

as broad as cell 1A with its posterior margin strongly

convex and having spots of darker pigment at the bases

of cells R4, R2+3, R5, and Cui.

Anthrax angustipennis Macquart

Anthrax angustipennis Macquart, 1840, p. 64.

Argyramoeba angustipennis.—Kert6sz, 1909, p. 60.

Anthrax binotatus Macquart, 1846, p. 113.

MALE.—Body black, femora and tibiae dark red;

integument brown and gray pruinose. Front with black

setae and a few black scales; setae on face black or

mixed yellow and black. Occiput with black setae and

a few black scales; fringe of hairs on posterior margin

black exteriorly and white interiorly above, mostly

white below. First antennal segment about as long as

apical width; second segment globular; base of third
66

segment flattened laterally, tapering abruptly to styliform part, about as broad as second segment in lateral

view; styliform part slightly longer than base, about

twice as long as style (Figure 107).

Discs of mesonotum and scutellum with linear black

scales, gold scales sometimes present laterally. Sternopleuron, lower half of mesopleuron, and anterior part

of pteropleuron with fine black setae and linear yellow,

black, or white scales. Pile on prosternum, propleuron,

dorsal half of mesopleuron, and anterior margin of

mesonotum white, some black hairs and setae on last

two, or with predominantly black pile on last three.

Postalar tuft of pile white. Fore coxa with black or

white bristles and yellow, white, or black scales; middle and hind coxae with black bristles and black and/or

yellow, or white scales. Hypopleuron and metapleuron

bare.

Wing (Plate 5d) pigmented anteriorly and basally,

hyaline posteriorly; pigment filling cells G, Sc, and Ri,

filling cell R2+3 to base of cell R4 and along apical margin to apex; margin of pigment extending posteriorly

from base of cell R4 to base of cell R5; and thence to

posterior margin at basal third of cell 2A. Base of cell

R» angled, with short spur; r-m crossvein at basal twofifths of cell 1M2, base of vein R2+3 slightly basad; contact of cells 1M2 and Cux about 2.5 times longer than

width of base of cell Cui; cell 1A puncuform apically;

cell 2A somewhat narrowed, narrower than cell 1A

postmedially; alula reduced, posterior margin straight.

Stigmatic area heavily pigmented; calypter pigmented,

fringe of hairs black. Stem of halter brown, knob brown

basally, yellow apically.

Fore and middle femora with black scales anteriorly

and a few white scales posteriorly; hind femur with

black scales; fore and middle tibiae with black scales

anteriorly and white scales posteriorly; hind tibia with

black scales. Middle femur with three or four weak

macrochaetae anteroventrally; fore tibia with complete

row of macrochaetae anterodorsally, hind tibia with

single row of macrochaetae anterodorsally.

Lateral margins of first abdominal tergum with yellow or white pile ventrally and black and brown pile

dorsally; lateral margins of terga two through five with

black setae, hairs, and scales. Posterior margin of first

tergum with linear black scales; discs of terga two

through four and sometimes center of five with linear

black scales. Remainder of terga with dense, overlapping, posteriorly produced, lanceolate-truncate, white

scales. Venter with yellow or black setae and white or

black scales.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MALE OENITALIA (Figure 69).—Gonocoxites long

and narrow, slightly folded inward along mesal line

apically; apices obtusely angled in ventral view; upper

margins lined with setae, ventral parts with setae fourfifths of way to base. Basal segment of gonostylus normally developed, evenly rounded dorsally in lateral

view, with a few fine setae mesally in ventral view; distal segment of gonostylus ovoid, with blunt tooth apically, with fine setae ventrally toward base. Dorsal part

of apex of epiphallus bluntly rounded apically in dorsal

view, evenly rounded laterally, with sharp projection

basally; ventrolateral part evenly rounded outward,

downward and inward from dorsal part and apex of

basal bands, rounded basad apically. Dorsal bands with

numerous setae both before and after their junction.

Basal part of aedeagus long and slender, almost tubular,

tapering slightly to junction with ventral bands beyond

junction of dorsal bands.

FEMALE.—Wing (Plate be) less extensively pigmented; pigmented area extending out to a line extending from tip of vein Rt

 across cells R! and R2+3 to a

point on vein R4+5 one-half to four-fifths of way to its

bifurcation, diagonally across cell R5 to a point on vein

M1+2 one-fifth to one-half of way to its bifurcation, and

then basally in a straight line to tip of vein 1 A, or curving across apex of cell 1A to hind margin midway in

cell 2A. Contact of cells 1M2 and Cui only slightly

longer than width of base of cell Gu!.

FEMALE GENITALIA (Figure 85).—Tenth tergum

with about 21 spines on each side. Ventral arm of ninth

tergum relatively broad, apex bilobed and turned inward distally. Dorsomedial corner of sclerite on each

side of gonopore with very small dorsal angle; lateral

arm very short, apically rounded; ventral arm long,

broadened parallel to meson and tapering to sharp,

mesally curved apex. Each duct of spermathecae about

3 times as long as bulb; first section very short, middle

section about 1.5 times longer than third section which

is slightly longer than bulb, expanded distally and with

small expansions postbasally; bulb long and slender,

symmetrical, about 3 times longer than wide and distinctly darker than duct

DISTRIBUTION.—The few specimens available of this

species indicate that it occurs in the tropical rain forest

of northern South America and in the Serra do Mar in

eastern Brazil (Map 19).

TYPES.—The type male of Anthrax angustipennis is

in the Paris Museum. According to notes supplied by

R. H. and E. M. Painter, it carries the labels "No. 905

Anthrax angustipennis" and "Sources de l'oyapok 2896,
NUMBER 4 3 67

jA ' SINUSOIDAL PROJECTION

f t ) > •-•.-...x.-

M AP 19.—Distribution of Anthrax angustipennis.
68

9?." It is badly rubbed, and the third antennal segments are missing. Specimens from the Amazon region

of Brazil agree well with the redescription of the type

given by Painter and Painter. The type-locality given

in Macquart's description is "De la Guyane, aux sources

de l'Oyapock." The river Oiapoque separates French

Guiana from the Territorio do Amapa, Brazil.

According to notes supplied by R. H. and E. M.

Painter, the two female types of Anthrax binotatus

Macquart are in the Bigot collection of the British

Museum under the label "Anthrax binotatus, n. sp." in

Macquart's handwriting, pasted on a larger label with

"Nova Grenada Nomm. par Macq." One of the specimens is headless but otherwise in good condition; the

other has the third segments of the antennae missing,

part of the head eaten, and a dense covering of fungus.

Specimens from northern South America agree well

with a redescription prepared by Painter. The typelocality is probably the island of Grenada in the British

West Indies.

DISCUSSION.—Females of angustipennis may be

readily distinguished from females of related species by

the absence of pigment in cell Rt beyond the apex of

vein Ri and by the absence of a postbasal spot in cell

R5. Males may be distinguished by the ovoid, flattened

distal segment of the gonostylus of the male genitalia

and by the distinctive wing pattern. Anthrax angustipennis may distinguished from other species in South

America by the greatly reduced, straight-margined

alula.

An interesting male specimen from Jatai, Goias,

Brazil, appears to be a hybrid between angustipennis

and macquarti. The wings are like those of angustipennis except that a patch of pigment extends into

cell R5 postbasally as in macquarti. The genitalia are

nearer those of macquarti, the gonocoxites being short

and broad, and the distal segment of the gonostylus

styliform, although broadened in lateral view. The

apex of the epiphallus has a dorsal plate as in macquarti, but the ventrolateral part projects laterally

and posteriorly. The dorsal bands have setae as in

angustipennis. This suggests that macquarti and austrinus are more closely related to angustipennis than

to the analis complex.

Anthrax argyropygus argyropygus Wiedemann

Anthrax argyropygus Wiedemann, 1828, p. 313 [argyroPyg<*]-—Curran, 1927, p. 85 [argyropyga].—Brimley, 1938,

p. 341 [argyropyga].—Painter and Painter, 1962, p. 71.—

Krombein, 1967, p. 397.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Argyramoeba argyropyga.—Osten Sacken, 1877, p. 242.—

Osten Sacken, 1878, p. 89.—jCoquillett, 1894, p. 95.—

Johnson, 1895, p. 325.—Kertesz, 1909, p. 61.

Spongostylum argyropyga.—Aldrich, 1905, p. 222 [Spogogostylum].—Johnson, 1913, p. 54 [Spogostylum].—Cole,

Malloch, and McAtee, 1924, p. 185 [Spogostylum].

Argyramoeba contigua Loew, 1869, p. 30; 1872, p. 144.

MALE.—Body black, tibiae red; integument mostly

gray and brown pruinose, eye margins silver pruinose.

Front with black setae and linear scales; setae on face

mixed black and white, extending almost to antennae.

Setae and scales on occiput entirely black, or black

above and white below; fringe of hairs on posterior

margin black above and mixed black and white below,

or black exteriorly and white interiorly above and white

below. First antennal segment about 1.5 times longer

than apical width; second segment lenticular; base of

third segment flattened laterally, about as wide as second segment, narrowing abruptly to styliform part

which is about equal in length to base and style (Figure

110).

Disc of mesonotum with linear black scales; margins with black scales, or often with linear gold scales

in front of wing or along entire margin, in a posterior

triangular area, and along anterior margin; white

scales sometimes in front of wing when gold scales are

absent. Disc of scutellum with linear black scales;

posterior margin usually with gold scales, often with

white scales medially and sometimes along entire margin; anterior margin often with gold scales. Sternopleuron, lower half of mesopleuron, and anterior half

of pteropleuron with yellow, black, or mixed, fine

setae, and yellow and white linear scales. Dorsal part

of mesopleuron with black, or mixed gold and black,

setae and white, or mixed white and yellow pile. Prosternum with white pile, some black hairs rarely intermixed, propleuron with white pile, or often with yellow

hairs intermixed and rarely with some black hairs;

anterior margin of mesonotum with white and black,

or white, yellow and black pile. Anterior coxa with

black and white, or black, yellow, and white bristles

and yellow, white, or mixed, scales; middle and hind

coxae with black, or black and gold bristles, and gold,

yellow, black, white, or mixed, scales. Metapleuron

sometimes with a few scdles posteroventral to spiracle.

Wing (Plate 5i) pigmented brown anterobasally,

hyaline posteriorly; pigment filling cells C and Sc, cell

Ri entirely pigmented except a crescent-shaped area

along posterior margin preapically, and rarely a linear

hyaline area submedially and basally; pigment usually

extending from apex of cell Rx across cell R2+3; cell
NUMBER 43 69

R2+3 filled to base of cell R4 except for a triangular

area along posterior margin before base of cell R4,

entire posterior part of cell rarely hyaline, a triangular

area of pigment usually extending from cell R2+3 into

cell R5 between bases of cells R2+3 and R4; extreme

base of cell R4 often pigmented; cell R entirely pigmented, or often with subhyaline or hyaline area preapically; cell M entirely pigmented; cell 1M2 usually

filled out to a line from r-m crossvein to near base of

cell Cui, rarely only at extreme base; base of cell Cui

pigmented or rarely entirely hyaline; apical margin

of pigment running from base of cell Cux to posterior

margin, leaving apical one-third to two-thirds of cell

2A hyaline. Base of cell R4 angled, often with short

spur; r-m crossvein at basal two-fifths of cell 1M2, base

of cell R2+3 at or slightly before r-m crossvein. Contact

of cells 1M2 and Cui 2.0 to 2.5 times as long as width

of base of cell Cui; cell 1A narrowly to broadly open

apically. Cell 2A narrower than cell 1A postmedially,

posterior margin incurved; alula vestigial, posterior

margin straight. Stigmatic area moderately pigmented.

Calypter pigmented, fringe of hairs yellow and white.

Halter dark brown or black, knob black or yellow at

extreme apex.

Scales on fore and middle femora black anteriorly,

white, yellow or mixed posteriorly, rarely entirely

black, hind femur with yellow and white scales posterobasally, and black scales anteroapically, scales rarely

entirely black. Scales on fore and middle tibiae black

anteriorly, white posteriorly; scales on hind tibia black.

Middle femur with five or six macrochaetae anteroventrally; hind femur with complete row of macrochaetae anteroventrally. Anterodorsal margin of fore

tibia with weak macrochaetae on apical half; hind tibia

with a single row of macrochaetae anterodorsally.

Sides of first abdominal tergum with white pile,

usually with some yellow hairs and black setae, rarely

predominantly brown and yellow; sides of terga two

through four with black setae, hairs, and linear scales.

Posterior margin of first tergum with long, gold or

white scales, or with white scales laterally and gold

scales medially. Discs of terga two through four and

center of five with linear black scales; linear gold

scales usually in lateral bands posteriorly on two

through four and on anterolateral corners of two, sometimes extending entirely across segments; abdomen

rarely without gold scales. Lateral thirds of fifth tergum, and six and seventh with dense, overlapping,

posteriorly produced, lanceolate, truncate, white

scales. Venter with sparse, fine setae, white anteriorly,

yellow posteriorly and sometimes on first sternum;

scales on sterna usually white; yellow or black scales

sometimes present.

MALE GENITALIA (Figure 66).—Gonocoxites long,

broad basally, tapering apically beyond middle; apices

bluntly angled dorsally in lateral view and apically in

ventral view. Basal segment of gonostylus rounded dorsally in lateral view, with two short setaceous, membranous lobes mesally in ventral view; apical segment

of gonostylus short, extending outward and slightly

apicad, triangular basally in ventral view, with a short

styliform part, apex slightly bent dorsoapically. Apex of

epiphallus with a convex, transverse shieldlike strurture above, with sharp dorsolateral angles, without a

medial angle, and shallowly emarginate apically below;

ventrolateral area rounded below, diagonally truncate

apicoventrally. Dorsal bands united apically, with

numerous setae both before and after junction. Base

of aedeagus narrow and tubular, tapering slightly to

junction with ventral bands well before junction of

dorsal bands.

FEMALE.—Similar to male. Thoracic pleura with

mostly white scales and pile; coxae with white scales

and mostly with white bristles. Pigment on wing (Plate

5;) more extensive, cell Rs filled to base of cell R4, a

crescent-shaped hyaline area sometimes present postbasally; pigment sometimes extending along base of

cell Mi; cell 1M2 pigmented out to base of cell 2M2;

cells 1A and 2A entirely pigmented except for apices;

contact of cells 1M2 and Cui about 1.5 times as long as

width of base of cell CUL Light scales on legs usually

mostly white rather than yellow. Gold scales on abdominal terga usually less extensive, sometimes absent;

white scales on posterior terga restricted to lateral

margins of fifth tergum. Venter usually with numerous

linear black scales posteriorly, sometimes with black

setae posteriorly.

FEMALE OENITALIA (Figure 86).—Tenth tergum

with about 18 spines on each side. Ventral arm of

ninth tergum parallel sided, not enlarged apically.

Dorsomedial corner of sclerite on each side of gonopore

not produced, with a sharp angle dorsally; lateral

arm short and acute; ventral arm narrow and parallel

sided. Each spermathecal duct about 3.5 times longer

than bulb; first section very short; second section about

2.5 times longer than third, which is about as long

as bulb and expanded apically; bulb cylindrical, about

3 times longer than wide, broadest subapically and

not well defined from duct.
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—Anthrax argyropygus argyropygus

occurs in forested areas throughout eastern United

States and along the east coast of Mexico (Map 20).

The few specimens available from Mexico seem to

indicate that it is allopatric to argyropygus albosparsus.

TYPES.—According to Painter and Painter (1962),

the apparent type male of Anthrax argyropygus Wiedemann is in the Vienna Museum. The specimen is in

good condition except that it lacks the third segment

of the right antenna. It carries the labels "America"

and "argyropyga Alte Sammlung" which are not in

Wiedemann's handwriting. Wiedemann recorded the

type-locality as "Vaterland?." The redescription of

the specimen agrees well with typical males from

southeastern United States.

The type female of Argyramoeba contigua Loew is

presumably with the remainder of Loew's types in the

Museum of Comparative Zoology at Harvard University. The type-locality is Virginia. The description

agrees well with typical specimens from that area.

BIOLOGY.—One reared specimen has been seen in

the collections examined by the author. It emerged

from the nest of a "Trypoxylonid" (Hymenoptera:

Sphecidae). Krombein (1967) reared this subspecies

from Trypargilum collinum collinum (Smith), T. c.

rubrocinctum (Packard), T. striatum (Provancher),

T'. tridentatum archboldi (Krombein) (Hymenoptera:

Sphecidae), Ancistrocerus catskill catskill (Saussure),

Stenodynerus fulvipes fulvipes (Saussure), S. pulvinatus surrufus Krombein, S. saecularis rufulus Bohart, S.

beameri Bohart, and Pachodynerus erynnis (Lepeletier) (Hymenoptera: Vespidae).

SCALE

00 ZOO 300 400 5OO 600 700 6O0 900 1000 MILES

200 AOO 800 800 I0O0 1200 I«OO KILOMETERS x -

LAMBERT'S AZIMUTHAL EQUAL-AREA PROJECTION

MAP 20.—Distribution of Anthrax argyropygus argyropygus (vertical lines), A. argyropygus

albosparsus (horizontal lines) and A. argyropygus painteri (diagonal lines).
NUMBER 4 3

DISCUSSION.—Specimens from southeastern United

States are usually darker than those from other regions,

having the gold scales on the thorax and abdomen less

extensive or absent. Two variant specimens in poor

condition from Tehuantepec, Oaxaca, Mexico, appear

to belong to this subspecies. They have more extensive patches of gold scales on the thorax and abdomen

than the other specimens available from Mexico. The

genitalia of the male have the dorsolateral corners of

the apex of the epiphallus acutely angled rather than

bluntly angled as in specimens from the United States.

The female has the pigment in cell R8 of the wing extending broadly along the base of cell M^ Another

male labeled Tehuantepec has genitalia like those of

a. albosparsus (Bigot).

The interrelationships of the subspecies of argyropygus are not well understood. Anthrax a. painteri seems

to be isolated in the mountainous area of eastern and

southern Arizona. Anthrax a. argyropygus and a. albosparsus, on the other hand, seem to be allopatric or

perhaps even partially sympatric in southern Mexico.

Specimens that appear related to each species have

been seen from Tehuantepec, Oaxaca. Many more

specimens from critical areas in southern Mexico will

be required to determine the true relationships of the

taxa.

Both a. argyropygus and a. albosparsus may be distinguished from a. painteri by the presence of pigment

in cell M in the male and in the apical half of cell 1A

in the female. Mexican specimens of a. argyropygus

may be separated from a. albosparsus by the presence

of linear gold scales on the mesonotum, scutellum, and

anterior abdominal terga, and by the numerous black

and gold hairs and setae on the pleura of the females.

Anthrax argyropygus painteri, new subspecies

MALE.—Body black, tibiae red; integument blue

gray and brown pruinose, eye margins silver pruinose.

Front with black setae and a few black and white

scales; face with mixed black and white setae. Occiput

with black setae on upper two-thirds, with white setae

below; scales on occiput mostly white, a few black ones

dorsally; fringe of pile on posterior margin black exteriorly and white interiorly on upper third, white on

lower two-thirds. First antennal segment about 1.5

times longer than apical width; second segment lenticular; base of third segment flattened laterally, narrowing abruptly to styliform part, about as wide as

second segment in lateral view; styliform part slightly

71

longer than base, 1.0 to 1.5 times longer than style

(Figure 112).

Disc of mesonotum with linear black scales; a few

linear gold scales on lateral margin in front of base of

wing, and on anterior half and posterior part of disc.

Scutellum with linear black scales on disc and linear

white scales on posterior margin, a few gold scales

laterally. Sternopleuron, mesopleuron, and anterior

half of pteropleuron with white setae and scales, a few

coarse black and gold setae on mesopleuron and pteropleuron; upper half of mesopleuron with white pile.

Prostemum, propleuron, and anterior margin of mesonotum with white pile; postalar tuft of pile black.

Coxae with coarse white and black bristles, white

scales, and a few yellow bristles. Hypopleuron and

metapleuron bare.

Wing (Plate 5g) mostly hyaline, cells C and Sc

pigmented light brown; bases of cells R2+3 and R5 narrowly surrounded by pigment; vein R2+3 narrowly lined

with pigment as far as base of cell R«; a small spot present apically in cell Ri; spur at base of cell R* narrowly

lined with pigment; small spots present at bases of cells

Cui and 1M2 and in cell R below base of Ri; base of

wing filled out to base of cell M. Base of cell R4 angled,

with long spur; r-m crossvein at basal third of cell 1M2;

base of vein R2+3 slightly basal to r-m crossvein; contact

of cells 1M2 and Cui about four times as long as

width of base of cell Cu*; cell 1A broadly open; cell

2A reduced, narrower than cell 1A; alula vestigial,

posterior margin straight. Calypter lightly pigmented,

fringe of hairs white. Halter brown, knob yellow

posteriorly at tip.

Fore and middle femora with black scales anteriorly

and some white and yellow scales posteriorly, posterior

femur with yellow scales covering basal third posteriorly and extreme base anteriorly, remainder with

black scales. Fore and middle tibiae with black scales

anteriorly and white scales posteriorly, hind tibia with

black scales. Middle femur with three macrochaetae

anteroventrally; hind femur with a complete row of

macrochaetae anteroventrally; fore tibia with a few

weak macrochaetae anterodorsally.

Lateral margins of first abdominal tergum with

white pile; lateral margins of terga two through four

with black setae, hairs and linear scales, some gold

scales and hairs anteriorly on second. Posterior margin

of first tergum with long, lanceolate white scales; discs

of terga two through four and middle of five with

linear black scales, narrow bands of linear gold scales

on anterior and posterior margins of two and posterior
72

margin of three. Lateral thirds of fifth tergum and

sixth and seventh with dense, overlapping, posteriorly

produced, lanceolate, truncate, white scales. Venter

with yellow and white setae and white scales, some

black setae and scales posteriorly.

MALE GENITALIA (Figure 68).—Gonocoxites short,

broad basally, tapering, relatively broad apically;

apices with sharp dorsal angles in lateral view, concave

laterally and bluntly angled apically in ventral view.

Basal segment of gonostylus sharply curved dorsally in

lateral view, with narrow, membranous lobe medially in

ventral view, lobe covered with fine setae; distal segment of gonostylus triangular basally, tapering outward

to a short styliform apex which is slightly expanded and

truncate. Setae on gonocoxites extending back as far as

medial fold, densest apically below; base of distal segment of gonostylus with fine setae. Dorsal part of apex

of epiphallus formed as a transverse convex shield with

sharp lateral angles and a very small medial angle dorsally, with a deep, right-angled apical emargination

below; ventrolateral part extending beyond dorsal

part as narrow truncate lobe. Dorsal bands with numerous setae, united mesally, broadly curved to junction

with apex of epiphallus. Base of aedeagus narrow and

tubular, tapering gradually to junction with ventral

bands well before junction of dorsal bands.

FEMALE.—Similar to male. Wing (Plate 5h) more

extensively pigmented; cell Ri filled except for a crescent-shaped hyaline area preapically along posterior

margin, pigment extending apically into cell R2+3; cell

R2+3 filled to base of cell R4 except for a small hyaline

area before base of cell R4 posteriorly; a triangular

area extending from cell R2+3 into R5 midway between

bases of cells R2+3 and R4; bases of cells R5, 1M2, and

Cut pigmented; cells R and M completely filled, anterobasal part of cell 1A filled.

FEMALE GENITALIA (Figure 88).—Tenth tergum

with about 16 spines on each side. Ventral arm of ninth

tergum narrow basally, broadened apically to form a

flat, mesally projecting flat plate. Dorsomedial angle

of sclerite on each side of gonopore not projecting,

obtusely rounded; lateral arm tapering to sharp,

slightly upcurved apex; ventral arm parallel sided on

basal two-thirds before tapering to sharp apex. Each

duct of spermathecae about 3.5 times longer than bulb;

basal section very short; middle section about 1.75

times longer than apical section which is slightly

longer than bulb and expanded apically; bulb symmetrical, elongate elliptical, slightly wider postbasally.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

VARIATION.—The additional specimens examined do

not vary significantly from the holotype and allotype.

DISTRIBUTION.—The few specimens available of

painteri were collected in mountainous areas in southeastern Arizona (Map 20). It may also occur in adjacent areas of similar habitat in central Arizona,

western Texas and extreme north-central Mexico.

HOLOTYPE.— o*, Southwest Research Station, 5 mi.

W Portal, Pima Co., Arizona, VIII-13-1959, 5400 feet

(H. E. Evans) (RHP).

ALLOTYPE.— °., Washington, Santa Cruz Co., Arizona, VIII-6-1932 (R. H. Painter) (RHP).

PARATYPES.—Arizona: Cochise Co., <?, Chiricahua

Mts., VII-4-1940 (D. E. Hardy) (RHP) ; <J, Southwest Research Station, 5 mi. W Portal, VIII-18-1959,

5400 feet (H. E. Evans) (NLM).

OTHER MATERIAL EXAMINED.—Arizona: Cochise

Co., ? , Canelo (Hyatt's Ranch), VI-30-1958 (A. and

H. Dietrich); <?, 5 mi. W Portal, Chiricahua Mts.,

VIII-7-1958 (R. M. Bohart). Pima Co., d\ Baboquivari Canyon, W Side Baboquivari Mts., VII-25-27-

1952 (H. B. Leech and J. W. Green); d\ Santa Rita

Mts., VI-22-1935 (F. H. Parker).

DISCUSSION.—The male of a. painteri differs from

both a. argyropygus and a. albosparsus in having cell

M hyaline except at the extreme base and apex rather

than completely pigmented. The female may be distinguished by the lack of pigment in the apical half of cell

1A and cell 2A. Anthrax a. painteri differs from laticellus in having cell 2A very narrow with its posterior

margin broadly concave.

Anthrax argyropygus albosparsus (Bigot)

Argyramoeba albosparsa Bigot, 1892, p. 348 [Argyromoeba].—

Kert6sz, 1909, p. 60 [albosparse].

Spongostylum albosparsum.—Aldrich, 1905, p. 222 [Spogostylum].

Anthrax albosparsus.—Painter and Painter, 1962, p. 69.

Argyramoeba angustipennis.—Williston, 1901, p. 275 [not

Macquart, 1840; misidentification].

Spongostylum angustipennis.—Aldrich, 1905, p. 222 [Spogostylum] [not Macquart, 1840; misidentification].

MALE.—Body black, tibiae red; integument mostly

gray and brown pruinose, eye margins and lower half of

occiput white pruinose, upper half of occiput blue

green pruinose. Front with black setae and lanceolate

black scales, a few white scales laterally below; face

with black setae extending almost to antennae, a few

white setae below; setae on occiput black, scales black,

a few white ones below; fringe of hairs on posterior
NUMBER 43 73

margin black externally and white internally above,

white below. First antennal segment about twice as long

as apical width; second segment lenticular, about as

wide as base of third segment which is flattened laterally and narrowed abruptly to styliform part; styliform

part about equal in length to base and style (Figure

111).

Mesonotum and scutellum with linear black scales,

some white scales on posterior margin of scutellum.

Sternopleuron, lower half of mesopleuron, and anterior

half of pteropleuron with black setae and hairlike white

scales. Prosternum and propleuron with white pile;

dorsal half of mesopleuron and anterior half of mesonotum with white pile and black setae; postalar tuft of

pile black. Fore coxa with white and black bristles, and

white scales; middle and hind coxae with black setae

and a few white scales, a few white setae present. Hypopleuron and metapleuron bare.

Wing (Plate 3h) brown anteriorly and basally, hyaline posteriorly; cells C and Sc filled with pigment,

cell Ri pigmented out to apex, a crescent-shaped hyaline area preapically and elongate hyaline areas at base

and submedially; cell R2+3 pigmented along upper margin as far as base of cell R4; cell R pigmented except for

a preapical hyaline spot; cell M entirely pigmented; extreme bases of cells 1M2 and Cux pigmented; apical

margin of pigment in cells 1A and 2A extending from

base of cell Cui to posterior margin just beyond base of

cell 2A. Base of cell R4 angled; r-m crossvein at basal

two-fifths of cell 1 M2, vein R2+3 arising slightly basad;

contact of cells 1M2 and Cux about 2.5 times longer

than width of base of cell Cui; posterobasal part of

wing greatly reduced; cell 1A broadly open; cell 2A

narrowed, much narrower than cell 1A postmedially,

posterior margin incurved; alula vestigial, posterior

margin straight. Stigmatic area lightly pigmented.

Calypter lightly pigmented, fringe of hairs white and

yellow. Halter brown, knob brown above except at

extreme apex, yellow below.

Scales on femora mostly black, a few yellow scales

posteriorly on fore and middle pairs; fore and middle

tibiae with black scales anteriorly and yellow and white

scales posteriorly; posterior tibia with black scales.

Middle femur with one or two macrochaetae anteroventrally; anteroventral row of macrochaetae on hind

femur incomplete basally. Anterodorsal surface of fore

tibia with a few weak macrochaetae; hind tibia with

single row of macrochaetae anterodorsally.

Sides of first abdominal tergum with white pile;

lateral margins of terga two through four with black

setae, hairs, and linear scales. Posterior margin of first

tergum with lanceolate white scales; discs of terga two

through four and middle of five with linear black scales.

Lateral thirds of fifth tergum, and sixth and seventh

with dense, overlapping, posteriorly produced, lanceolate-truncate, white scales. Sterna with fine, sparse,

white and black setae anteriorly and black setae posteriorly, with a few linear black scales.

MALE GENITALIA (Figure 67).—Gonocoxites short

and broad ventrally; apices with sharp dorsal lobes and

slightly obtuse ventral angles; setae evenly distributed

over anterior parts, extending almost to base ventrally.

Basal segment of gonostylus rounded in lateral view,

with a membranous lobe medially extending beyond

base of apical segment and covered with fine setae; base

of distal segment of gonostylus rounded transversely,

flattened apically, tapering outward to a long narrow

styliform part bent sharply upward and inward at apex.

Dorsal part of apex of epiphallus formed as a transverse shieldlike structure, convex exteriorly with acute

lateral angles; ventrolateral parts bluntly rounded

apically, extending far beyond dorsal part, thus forming a deep medial emargination below. Dorsal bands

joined mesally, very broad before junction, with a line

of setae extending almost to apex. Base of aedeagus

narrow, evenly rounded dorsally, undulated ventrally,

narrowing to junction with ventral bands before junction of dorsal bands.

FEMALE.—Setae on sternopleuron and lower half

of mesopleuron all white. Yellow scales more extensive

on posterior surfaces of fore and middle femora. Wing

(Plate 3i) more extensively pigmented; cell Ri without hyaline areas basally; cell R2+3 pigmented out to

base of cell R4 except for a concave hyaline area before

base of cell R4, cell R5 with a large spot extending from

cell R2+3 between bases of cells R5 and R4; bases of cells

1M2 and Cux broadly pigmented; cell 1A pigmented

except at extreme apex; contact of cells 1M2 and Cui

about twice as long as width of base of cell Cui. Posterior margin of first abdominal tergum with dense,

long, lanceolate white scales; posterior margins of second and third terga with small spots of white scales

medially; white scales on posterior segments restricted

to lateral margins of fifth tergum. First two sterna with

mixed black and white setae, posterior sterna with

black setae and black scales.

FEMALE GENITALIA (Figure 87).—Tenth tergum

with about 18 spines on each side. Ventral arm of

nine tergum undulating and expanding mesally into

a broad, dorsally convex plate at apex. Dorsomedial
74

angle of sclerite on each side of gonopore rounded, not

produced; lateral arm tapering to a sharp, upturned

apex; ventral arm tapering and slightly curving mesad

toward apex. Each spermathecal duct about twice as

long as bulb; basal section very short; medial section

about 2.5 times longer than apical section which is

straight; bulb elongate, tapering from just before apex

to base and sharply rounded apically.

DISTRIBUTION.—Anthrax a. albosparsus occurs in

the forested areas of southern Mexico (Map 20). The

few specimens available indicate that it may be allopatric to a. argyropygus on the east. The one specimen

of this species collected by the author was found in a

steep, rocky canyon 18 miles north of Chilpancingo,

Guerrero. The vegetation was an open, subtropical

evergreen forest with numerous cacti and other xerophytic plants.

N EW MATERIAL EXAMINED.—Guatemala: ? , Quirigua, August (Schaus and Barnes).

Mexico: Guerrero, <$, Acapulco, VI-15-1935

(A. E. Pritchard); ? , 18 mi. N Chilpancingo, VIII-7 -

1962 (N. Marston-8). Morelos, $ , 11 mi. E Cuernavaca, Lobo Canyon, VIII-15-1962, 3900 feet (R. H.

and E. M. Painter); d1, 12 mi. E Cuernavaca, Lobo

Canyon, VIII-14-1962, 3600 feet (R. H. and E. M.

Painter). Oaxaca, cf, Tehuantepec (Sumichrast).

TYPE.—According to Painter and Painter (1962)

the type of Argyramoeba albosparsa Bigot, a female,

is in the Bigot collection in the British Museum. It

carries the label "Argyramoeba albosparsa $ , n. sp.

In6dit. Quinz. 1888 J. Bigot. Am. du Nord Colorado."

The redescription by Painter and Painter along with

the photograph of the wing of the type indicates that

this specimen is a typical member of the species

argyropygus closely allied to the subspecies from

eastern United States and Mexico.

DISCUSSION.—Because the subspecies of argyropygus

have not been collected in Colorado, it seems unlikely

that this specimen came from there. Also, the remainder of Bigot's species described from specimens collected in 1888 were from the Pacific coast. It is possible

that the "Colorado" refers to the Colorado River, but

this seems unlikely since the females of the Arizona

subspecies have most of cells 1A and 2A hyaline whereas the type has these cells completely pigmented.

Another possibility is that the type came from southern

Mexico. If this is the case, the type-locality may be the

small town of Tierra Colorado just north of Acapulco.

This seems plausible since the person who collected

Bigot's specimens apparently was traveling by ship and

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

probably stopped at Acapulco either on his way to

California or in returning. As the description of the

type fits very well a specimen collected by the author

18 miles north of Chilpancingo, Guerrero, the name

albosparsus is being used for this southern Mexico

subspecies.

A female collected near Cuernavaca, Morelos, differs from the typical form in having the wing more

heavily pigmented with the margin indistinct. Cell Rx

is entirely pigmented as far as the apex of vein Sc, and

the pigment extends across cell R2+3 into the base of

cell R*; cells R2+3 and R5 are pigmented as far as the

base of cell Mi, and the pigment extends across the

apex of cell 1M2 into the base of cell Mi and the

anteromedial angle of cell 2M2; cells 1M2 and Cux

are pigmented beyond the base of cell 2M2; cell 1A is

entirely pigmented and cell 2A is subhyaline. The posterobasal part of the wing is less reduced than in the

typical form, with cell 2A as wide as cell 1A postmedially. Whether or not this specimen belongs to

a. albosparsus is a matter of conjecture. It is being

placed here since it seems to be only an extreme manifestation of the normal variation noted in the species

argyropygus.

A female from Quirigua, Guatemala, is tentatively

placed with this species since it lacks the gold scales

that seem to characterize Mexican specimens of

a. argyropygus. It differs from the female described in

having the wing more extensively pigmented, cells 1A

and 2A being entirely filled; cell 2A is wider, almost as

wide as cell 1A. The scales on the first abdominal

tergum and the scales and setae on the sternopleuron

and mesopleuron are black.

The limited material available of a. albosparsus indicates that the females of this species differ from females

of a. argyropygus from Mexico in lacking gold scales

posteriorly on abdominal terga two and three and in

having the posterior margin of the first tergum lined

with dense white scales. Males of a. albosparsus differ

from those of a. argyropygus in lacking gold scales on

the thorax and abdomen and in having the ventrolateral parts of the apex of the epiphallus projecting

forward, thus creating a deep medial emargination in

dorsal view.

Anthrax innubilipennis, new species

MALE.—Body black, mostly brown and gray pruinose, margins of eyes and lower part of occiput silver

pruinose. Front with black setae and a few linear black
NUMBER 4 3 75

scales. Face with mixed black and white setae extending almost to antennae. Occiput with black setae and

scales on upper half, with white setae and scales on

lower half; fringe of hairs on posterior margin black

externally and white internally on upper half, white on

lower half. First antennal segment about twice as long

as apical width; second segment globular; base of third

segment flattened laterally, narrowing abruptly to styliform part, about as wide as second segment; styliform

part of third segment about as long as base, about

1.5 times longer than style (Figure 109).

Disc of mesonotum mostly with linear black scales;

lateral and anterior margins and triangular area on

posterior margin with linear gold scales. Disc of scutellum with linear black scales; anterior and posterior

margins with linear gold scales. Sternopleuron with

fine yellow setae and linear yellow scales; mesopleuron

and anterior half of pteropleuron with yellow and gold

setae and hairs, some coarse black and gold setae above,

and some white pile anteriorly on mesopleuron. Pile on

prosternum, propleuron, and anterior margin of mesonotum white, some gold hairs on propleuron. Postalar

tuft of pile mixed gold and black. Metapleuron with

linear scales posteroventral to spiracle; hypopleuron

with some hairs and scales on anteroventral corner.

Fore coxa with gold setae and lanceolate white scales;

middle and hind coxae with gold and black setae and

linear gold scales.

Wing (Plate 5c) mostly hyaline, cells C and Sc,

extreme bases of cells R, M, 1A, and 2 A, small spot

below base of vein R5 and base of wing grayish brown.

Sectoral crossvein present; vein R* evenly curved at

medial angle; cell R2+3 narrow, less than half as wide

medially as apically; r-m crossvein located at basal

third of cell 1M2, vein R2+3 arising slightly basad of r-m

crossvein; contact of cells 1M2 and Cu! more than twice

as long as width of base of cell Cui; posterior part of

wing narrowed, cell 2A narrower than cell 1A, but not

incurved; alula greatly reduced, posterior margin

straight. Opening of anal cell half the width of r-m

crossvein. Halter brown, knob brown above except

at extreme tip, yellow below except on margins.

Fore and middle femora with cuneate gold and

yellow scales posteriorly, mixed gold and black scales

anteriorly; hind femur with black scales along ventral

margin and apically, with yellow scales elsewhere. Fore

and middle tibiae with black scales anteriorly and gold

and yellow scales posteriorly; hind tibia with black

scales and a few yellow ones basally. Middle femur

with three or four weak macrochaetae anteroventrally

toward base and one or two strong macrochaetae postmedially on anterior side; hind femur with a row of

weak macrochaetae anteroventrally; fore femur with a

postmedial bristle anteriorly. Fore tibia with complete

row of macrochaetae anterodorsally; hind tibia with

single row of macrochaetae anterodorsally.

Lateral margins of first abdominal tergum with

white pile, few gold hairs present posteriorly; lateral

margins of terga two through four with sparse black

setae and pile, and long, linear, erect, black and gold

scales; terga five through seven with gold setae laterally. Discs of terga two through four and medial part

of five with linear black scales; posterior margins of

one through four with linear and lanceolate gold scales.

Remainder of terga with overlapping, ovate and ovatetruncate, shining white scales. Venter with lanceolate

white scales and fine, sparse white, yellow, and gold

hairs, gold predominating posteriorly.

MALE GENITALIA (Figure 73).—Gonocoxites broad,

tapering upward from ventral margins to a sharp apical

angle in lateral view, apices with acute lateral angles

in ventral view, setae extending about halfway to base,

densest medially. Basal segment of gonostylus greatly

enlarged and flattened, appearing as a broad, apically

rounded, flat plate in ventral view, lateral margin

angled upward and medial line raised as a sharp ridge

in lateral view. Distal segment of gonostylus formed

dorsally to apex of gonocoxites, quadrangular with an

outfolded dorsoapical margin and a long truncate basal

lobe in lateral view, with a short, truncate lobe apicolaterally and a broad, lanceolate, acute lobe basolaterally in ventral view. Apex of epiphallus with a rounded

medial part dorsally from which long, upcurved, sharp

horns extend laterally; ventrolateral part broad and

rounded apically in lateral view, with medial longitudinal fold. Dorsal bands united mesally, with a

hump just after the junction, without setae. Base of

aedeagus bulbous, bent sharply downward apically;

junction with basal bands not visible in lateral view.

VARIATION.—The bristles on the coxae may be predominantly black. The scales on the femora and tibiae

may be more extensively black than in the type.

DISTRIBUTION.—This species is known only from

the three specimens listed below, although it probably

has a wider range throughout the mountains of southern Mexico. The holotype and one paratype were collected "hovering on sunny side of cement culvert."

HOLOTYPE.—cf j 25 mi. S Iguala, Guerrero, DC-14—

1963, 2400 feet (R. H. and E. M. Painter) (RHP).
76

PARATYPES.—Mexico: Guerrero, d1, 25 mi. S Iguala,

LX-14-1963, 2400 feet (R. H. and E. M. Painter)

(NLM). Morelos, d\ 3 mi. N Alpuyeca, V-9-1959,

3400 feet (H. E. Evans) (RHP).

DISCUSSION.—If the female of innubilipennis differs

from the male in the same way that the female of

delicatulus Walker differs from the male of that species,

it probably lacks the sectoral crossvein and has the

wing more extensively pigmented. The presence of

a postmedial bristle on the anterior side of the middle

femur should serve to separate it from other related

species from Mexico.

Anthrax delicatulus Walker

Anthrax delicatulus Walker, 1849, p. 266 [delicatuld\.—Osten

Sacken, 1858, p. 39 {delicatula).—Aldrich, 1905, p. 230

[delicatula].—Painter and Painter, 1962, p. 73.

FEMALE.—Body black, tibiae reddish brown; integument brown to blue green pruinose, margins of eyes,

face and lower half of occiput white pruinose. Front

with black setae. Face with mixed black and white

setae. Occiput with black setae and linear scales on

upper third, with white setae and linear scales on

lower two-thirds; fringe of hairs on posterior margin

black exteriorly and white interiorly on upper third,

white below. First antennal segment about 1.5 times

longer than apical width; second segment globular;

base of third segment flattened laterally, about as wide

as second segment in lateral view, tapering abruptly

to styliform part which is slightly shorter than base

and about 1.5 times longer than style (Figure 108).

Disc of mesonotum with linear black scales; lateral

margin in front of base of wing and posterior margin

with long, linear white scales. Scutellum with linear

black scales. Mesopleuron, sternopleuron, and anterior

half of pteropleuron with fine yellow setae and long,

linear, white scales, some coarse yellow setae above on

mesopleuron and pteropleuron, and some white pile

anteriorly on mesopleuron. Prosternum, propleuron,

and anterior margin of mesonotum with white pile;

postalar tuft of pile mixed black and white. Metapleuron with large patch of linear white scales posteroventral to spiracle; hypopleuron with small patch of

white scales and hairs anteroventrally. Coxae with

mixed black and white setae, and linear white scales.

Wing (Plate 5b) brown anteriorly and basally, hyaline posteriorly; pigment filling cells C, Sc, and Rla

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

extending from tip of cell Ri across tip of cell R2+3

into tip of cell R4, filling cell R2+3 beyond and into base

of cell R4, cell R5 filled to base of cell R4 and extending faintly along base of cell Mi; cell 1M2 pigmented

basally and along anterior margin three-fourths of way

to apex; basal fifth of cell Cui and extreme base of

cell 2M2 pigmented, pigment extending basally from

cell Cui to posterior margin near base of cell 2A,

leaving apical fifth of cell 1A and posteroapical twothirds of cell 2A hyaline. Sectoral crossvein incomplete,

base of cell R4 angled and with long spur, medial angle

slightly recurved and sharply bent forward. R-m crossvein at basal third of cell 1M2, vein R ^ arising opposite. Contact of cells Cui and 1M2 three times as long

as width of base of cell Cui. Posterobasal margin of

wing reduced, but margin not incurved, cell 2A as

wide as cell 1A preapically; alula greatly reduced,

posterior margin straight. Stigma tic area lightly pigmented. Calypter pigmented, fringe of hairs black.

Halter dark brown, knob black, extreme tip lighter.

Fore and middle femora with cuneate white scales

posteriorly and black scales anteriorly; hind femur

with white scales on basal half posteriorly and at base

anteriorly, remainder with black scales. Fore and middle tibiae with yellow and white scales posteriorly and

black scales anteriorly; hind tibia with black scales.

Middle femur with a large postmedial bristle on anterior side, without bristles anteroventrally; hind femur

with row of bristles anteroventrally incomplete basally.

Fore tibia with complete row of anterodorsal macrochaetae; hind tibia with single row of anterodorsal

bristles.

First abdominal tergum and anterior part of second

with white pile laterally, posterior part of second tergum, and third and fourth with black pile, setae and

lanceolate scales laterally, some white setae posteriorly

on fourth. Hind margin of first tergum with long,

linear white scales. Discs of following terga with linear

black scales except for patches of lanceolate, truncate,

white scales laterally on five and six. Anterior sterna

with fine, white and black setae and long, linear and

lanceolate, white scales, some medial black scales;

posterior sterna with black setae and linear black scales.

MALE.—Similar to female. Wing (Plate 5a) less

extensively pigmented, pigment not completely filling

cell R a at apex, filling cells R2+3 and R5 only halfway

to base of cell R4, extending only slightly into bases of

cells 1M2, Cul5 and 1A, and leaving Cell M subhyaline
NUMBER 43 77

medially and hyaline apically. Sectoral crossvein complete, vein R4 strongly sinuate medially; contact of

cells 1M2 and Cui about 1.66 times longer than width

of base of cell Cux; posterior part of wing less narrowed

than in female, cell 2A wider than cell 1A postmedially.

Middle femur with several macrochaetae anteroventrally. Last three abdominal terga with overlapping

ovate scales with bifid apices. Posterior sterna with

lanceolate white scales.

MALE GENITALIA (Figure 74).—Gonocoxites very

broad in ventral view, with a preapical fold, tapering

concavely to acute, upcurved apices in lateral view;

apices broad, sharply rounded apically in ventral view,

mesal margin rounded to narrow mesal sulcus; setae

very sparse and coarse basad of preapical fold, finer and

more dense apically. Basal sclerite of gonostylus very

small, not visible in lateral or ventral view, with long

setae apically; distal segment with a long, sharp,

slightly outcurved, mesal lobe and a shorter, broader

lateral lobe, fine setae at base and on lateral lobe. Apex

of epiphallus flattened dorsoventrally, apex broadly

rounded, a short, sharp lobe curving upward and backward from each side; a flat longitudinal plate extending ventrally under lateral lobes. Dorsal bands united

medially, with numerous setae. Base of aedeagus narrow, tapering irregularly to junction with ventral

bands below junction of dorsal bands; ventral bands

with a pouchlike protuberance halfway to apex.

DISTRIBUTION.—Only two specimens of this species

are available, a female in excellent condition from

Clarendon, Portland Ridge (N. side), Jamaica, West

Indies, VII-19-1958 (T. H. Farr) and a male missing

the head and anterior part of the thorax from San

Lorenzo, Dominican Republic, VI-27-29-1915.

TYPE.—According to Painter and Painter (1962)

the type female of A. delicatulus is in the British

Museum. It is well preserved except that the head is

missing. The type-locality is Jamaica.

There is some doubt about the association of this

male and female. Because the two specimens are

similar in all respects except the wing pattern and venation and because a similar sexual dimorphism occurs

in the closely related A. angustipennis Macquart, the

male and female are regarded here as the same species.

The male of A. delicatulus may be separated from

the male of A. innubilipennis by the heavier infuscation of the wings. Both males and females may be distinguished from other closely related species by a postmedial bristle on the anterior side of the middle femur.

Anthrax macquarti d'Andretta and Carrera

Anthrax leucopygus Macquart, 1855, p. 76 [leucopyga].—

KertSsz, 1909, p. 43 [leucopyga] [not Macquart, 1840;

preoccupied].

Anthrax macquarti d'Andretta and Carrera, 1952, p. 296.

MALE.—Body black; integument mostly grayish and

brownish pruinose, eye margins and lower part of occiput silver pruinose. Front with black setae and lanceolate black or mixed black and white scales; face

with black, yellow or white setae extending almost to

antennae. Occiput with black, white or mixed setae

and linear scales, fringe of hairs on posterior margin

white, black, or black above and white below. First

antennal segment about as long as apical width; second segment globular; base of third segment flattened

laterally, slightly wider than second segment in lateral

view, narrowing abruptly to styliform part which is

about as long as base and about 1.6 longer than style

(Figure 113).

Discs of mesonotum and scutellum with fine black

setae and linear scales; gold scales sometimes in front

of base of wing, along posterior margin on mesonotum,

and anteriorly and laterally on scutellum; white scales

often present laterally on mesonotum and posteriorly

on scutellum. Sternopleuron, lower half of mesopleuron, and anterior half of pteropleuron with fine

black or white setae and linear scales. Upper half of

mesopleuron, prosternum, propleuron, and anterior

margin of mesonotum with white, or mixed black and

white pile. Postalar tuft of pile black, gold or mixed.

Hypopleuron and metapleuron bare. Coxae with

coarse black setae and black or white, linear and lanceolate scales.

Wing (Plate 5k) dark brown basally, hyaline apically, pigment entirely filling cells C and Sc, extending

out to a line running from a point on vein Rx basal

to tip of Sc obliquely across cells R! and R2+3, basally

almost to r-m crossvein, posteriorly to extreme base of

cell 2M2, and thence to posterior margin leaving apices

of cells 1A and 2A hyaline for a distance equal to twice

width of r-m crossvein, a small pigmented area extending from cell R2+3 into cell R5 postbasally. Base of

cell R4 angled; r-m crossvein located at basal third of

cell 1M2; contact of cells 1M2 and Cui slightly less or

slightly more than width of base of cell Cux, apex of

cell 1A punctiform or narrowly open. Stigmatic areas

very lightly pigmented. Alula narrowed, posterior

margin straight or slightly convex or concave. Calypter
78

pigmented, fringe of hairs black, white or brown. Halter brown, knob broadly yellow apically.

Femora and tibiae with cuneate black scales, yellow

and white scales often present posteriorly and basally.

Middle femur with one to three macrochaetae anteroventrally; hind femur with anteroventral row of

macrochaetae incomplete basally; anterodorsal surface

of fore tibia with few macrochaetae apically; hind

tibia with a single row of macrochaetae anterodorsally.

Lateral margins of first abdominal tergum with

white pile, often some black hairs, pile sometimes entirely black; lateral margins of terga two through four

with black pile and setae, sometimes white hairs anteriorly on two. Posterior margin of first tergum and

discs of terga two through four with fine black setae and

linear scales, sometimes long white scales laterally on

one; terga five through seven with long, overlapping,

lanceolate-truncate, silver scales, linear black scales

sometimes present medially on five. Sterna with fine

black or black and gold scales, some lanceolate white

scales on five and six and sometimes laterally on one

and two.

MALE GENITALIA (Figure 70).—Gonocoxites rather

broad basally, tapering apically; apices rounded in

ventral view; dorsal margins with setae; ventral parts

with setae almost to base. Basal segment of gonostylus

flattened dorsally in lateral view, rounded apically

with a few fine setae in ventral view. Distal segment of

gonostylus with a bulbous base (triangular in ventral

view) tapering to a styliform apex extending upward

and outward; apex sharply curved dorsolaterally; ventral part of base with fine setae. Dorsal part of apex

of epiphallus raised above ventrolateral parts, cuneate

apically, spatulate basally in dorsal view, convex and

sharply pointed basally and apically in lateral view;

ventrolateral parts extending downward and backward

from dorsal part, apices evenly rounded, not reaching

apex of dorsal part. Dorsal bands without setae, imperfectly united apically. Base of aedeagus broad, but

not bulbous, tapering gradually to junction with ventral bands beyond junction of dorsal bands.

FEMALE.—Similar to male. Silver scales on posterior

terga restricted to lateral margins of terga five and six.

FEMALE GENITALIA (Figure 89).—Tenth tergum

with about 22 spines on each side. Ventral arm gradually tapering to acute apex. Dorsomedial angle of

sclerite on each side of gonopore sharply produced

dorsomedially; lateral arm narrow, sharply pointed

distally and curled under below; ventral arm narrow,

parallel sided and nearly straight. Each spermathecal

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

duct about 1.5 times longer than bulb; first sections

very short; second section about two-thirds as long as

third section which is nearly as long as bulb, slightly

curved and expanded apically; bulb elongate oval,

about twice as long as maximum width, slightly curved

and darker than third section of duct.

DISTRIBUTION AND ECOLOGY.—Anthrax macquarti

occurs in northeastern South America as far south as

the state of Sao Paulo in Brazil and into central Paraguay (Map 21). A specimen from the Rio Negro in

Amazonas indicates that it probably occurs throughout

the Amazon Basin.

Two specimens of this species were collected by the

author in Rio de Janeiro, Guanabara, Brazil: one at

Serro, Minas Gerais, and one at Petrolina, Pernambuco. The specimens from Rio de Janeiro and Serro

were collected in open areas in tropical forest. One of

the specimens was observed ovipositing in a steep bank.

The specimen from Petrolina was collected in an open

sandy area near the Rio Sao Francisco where the vegetation was typical of the semidesert areas of northeastern Brazil. These observations indicate that the distribution of macquarti is not limited to certain vegetation

zones as are many species of Bombyliidae.

TYPE.—Painter and Painter (unpublished notes)

found that the three specimens associated with the

name Anthrax leucopygus Macquart in the Bigot collection in the British Museum represented three species. A female agreeing with Macquart's description

and figure was considered the type and so marked. It is

headless and has several legs missing but is otherwise

in good condition. A photograph of the wing and redescription indicate that the type represents the species

described herein.

Macquart (1840) described another species which

he named Anthrax leucopygus from Timor, making

leucopygus Macquart, 1855, a junior homonym.

D'Andretta and Carrera (1952) proposed the name

macquarti for this species, although the specimen which

they considered to be leucopygus was actually a specimen of A. midas Fabricius. According to Mayr, Linsley, and Usinger (1953, Appendix G), it was ruled

at the Fourteenth International Congress of Zoology

in Copenhagen "that a substitute name ('nom. nov.')

which is in part also based on particular specimens

applies in all circumstances to that which the previously published name is applicable." Thus, the name

macquarti is to be applied as a new name for leucopygus Macquart, 1855, rather than being a synonym of

midas Fabricius.
NUMBER 43

DISCUSSION.—Anthrax macquarti exhibits a broad

range of variation. Specimens from the semidesert and

desert areas of northeast Brazil have much more extensive white pile on the thoracic pleura and the sides of

the abdomen, and the fringe of pile on the posterior

margin of the occiput becomes entirely or predominantly white. There are also much more extensive

patches of yellow or white scales on the thorax, and

the postalar tuft of pile is usually predominantly gold

79

or yellow, as are the setae on the face. Specimens from

forested areas have the white pile on the body more

or less restricted to the anterior margin of the thorax

and the sides of the first abdominal tergum, usually

with black, yellow or white scales on the thorax, and

have the postalar tuft of pile and the setae on the face

black. In the extreme, a male from Serro, Minas Gerais,

has only a few white hairs anteriorly on the mesonotum

and the usual covering of silver scales on the apical

MAP 21.—Distribution of Anthrax macquarti.

369-673 O—70 6
80

abdominal terga. The wing patterns are much the same

throughout the range although those from drier areas

tend to be less extensive. The genitalia of males from

the two areas appear almost identical.

If long series are acquired from various populations

throughout the range, it may be possible to differentiate

subspecies. With the limited material now available

and lack of definite gaps in the range of variation, however, it seems best to regard all of the populations as

a single taxon.

Anthrax macquarti may be separated readily from

repertus, clinopictus, and gideon by the straight or

slightly concave margin of the alula and by the postbasal spot extending from cell R2+3 into cell R5 and

isolating a small basal hyaline area. The relationship

between macquarti and angustipennis is discussed under the latter species.

Anthrax austrinus, new species

FEMALE.—Body generally black, femora reddish

brown basally and yellow apically, tibiae yellow; integument mostly brownish pruinose; lower part of occiput and margins of eyes silver pruinose. Front with

black setae and lanceolate black scales, some white

scales laterally below. Face with black setae extending

almost to antennae, some white hairs below. Upper

third of occiput with short black setae; lower twothirds with black and white setae, the white predominant posteriorly, and white scales; fringe of pile on

posterior margin black on upper third, becoming white

on lower two-thirds. First antennal segment about as

long as apical width; second segment globoid, slightly

flattened apically, about as wide as base of third segment in lateral view; basal part of third segment

flattened laterally, narrowing abruptly to styliform part

which is slightly longer than base and about twice as

long as style (Figure 114).

Disc of mesonotum and scutellum with fine black

setae and linear scales, mixed black and orange brown

on anterior and lateral parts of mesonotum, and posterior margin of scutellum, black elsewhere; lateral margins of mesonotum with coarse setae and linear, semierect, mixed white and orange-brown scales. Sternopleuron and anterior half of pteropleuron with fine

black setae and linear gold scales, some coarse black

setae dorsally on pteropleuron; mesopleuron with white

pile anterodorsally and black setae and white scales

posteroventrally. Pile on prosternum, propleuron, and

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

anterior margin of mesonotum white, some black setae

on latter; postalar tuft of pile black. Anterior coxa with

black and white bristles and white scales; middle and

hind coxae with black bristles and gold scales. Hypopleuron bare; metapleuron with several white scales

behind spiracle.

Wing (Plate 5/) dark brown basally, hyaline apically, pigment extending out to a line running from

after tip of Sc irregularly to vein M1+2 before base of

cell Mi, basally for distance equal to r-m crossvein,

back to vein R4+5 and basally leaving hyaline spot

basally in cell R5, posteriorly from a point distal to rm crossvein across extreme base of cell 2M2 to vein

2A before its apex, anteriorly again, leaving a hyaline

area subapically in cell 1A, and thence to posterior

margin two-thirds of way to apex of cell 2A. Vein

R4 sharply angled basally; r-m crossvein located

two-fifths of distance from base to apex of cell

1M2; base of vein R2+3 located basad to r-m crossvein

slightly less than width of that crossvein; contact of

cells IMo and Cux slightly greater than width of base

of cell Cux; cell 1A punctiform apically. Alula narrowed, the posterior margin straight. Halter light

brown, knob yellow tipped dorsally and mostly yellow

ventrally.

Fore and middle femora with mixed yellow and

white cuneate scales posteroventrally and black scales

anterodorsally; hind femur with yellow and white

scales basally and black scales apically; fore and middle

tibiae with yellow scales posteriorly and mixed gold

and black scales anteriorly; hind tibia with mixed

gold and black scales. Middle femur with one to three

strong macrochaetae anteroventrally; hind femur with

five to seven strong macrochaetae anteroventrally; fore

tibia with macrochaetae distally on anterodorsal side;

hind tibia with single row of macrochaetae anterodorsally.

First abdominal tergum with white pile laterally;

terga two through four with black pile laterally, some

white hairs and scales posteriorly on four; terga five

and six with long, narrow, silvery-white scales and

black setae laterally; seventh tergum with only black

setae laterally. Posterior margin of first tergum and

discs of remaining terga with fine black setae and linear

black scales; some linear, orange-brown scales on posterior margins and some yellow and white scales posterolaterally. Abdominal sterna with fine white and gold

setae and linear black scales; numerous white, yellow

and orange-brown scales posteriorly and laterally on all

segments.
NUMBER 43 81

MALE.—Similar to female. Lateral fourths of fifth

abdominal tergum, most of sixth tergum and all of

seventh with posteriorly produced, long, lanceolatetruncate, overlapping, white scales.

MALE GENITALIA (Figure 71).—Gonocoxites narrow, dorsal margins curving concavely to sharply

rounded apices in lateral view; apices narrowly

rounded in ventral view with blunt ridges medially,

medial sulcus narrow; setae sparse, short, extending

basally halfway from medial fold to base. Basal segment of gonostylus sinuate dorsally in lateral view,

blunt apically in ventral view, with fine setae apically.

Distal segment of gonostylus round basally, tapering

apically, styliform part projecting upward and outward, apex curved dorsolaterally; base with fine setae

ventrally and laterally. Apex of epiphallus with a raised

dorsal plate which is right angled basally and slightly

emarginate apically in dorsal view; ventrolateral part

curved downward and inward, rounded below, straight

apically in lateral view, extending slightly past middle

of dorsal plate. Dorsal bands united mesally, with

three small setae. Base of aedeagus tapering conically

to junction with ventral bands below junction of dorsal

bands.

VARIATION.—The only other specimen available of

this species is similar in all respects to the holotype.

DISTRIBUTION.—Anthrax austrinus is known from

only two localities, Nova Teutonia, Santa Catarina,

and "Granja Carola," Rio Grande do Sul, Brazil. It

probably occurs in forested areas from Parana southward.

HOLOTYPE.— $ , Nova Teutonia, Santa Catarina,

Brazil, 27°11'S, 52°23'E, XII-1-1938 (Fritz Plaumann) (BM).

PARATYPES.—Brazil: Rio Grande do Sul, d,

Granja Carola (Dr. M. Leitao) (NLM). Santa Catarina, ? , Nova Teutonia, 27°11'S, 52°23'E, XII-19-

1959 (Fritz Plaumann) (UCAL).

DISCUSSION.—The male paratype is not designated

as the allotype, since the head is missing and it has

only one leg and one wing. The female paratype lacks

one wing and two legs but is otherwise in good condition.

The distinctive wing pattern of this species will

serve to separate it from all other species in the cephus

group from North and South America. The structure

of the wings and the male genitalia indicate its close

relationship to macquarti.

Anthrax repertus Walker

Anthrax repertus Walker, 1852, p. 188.—Kertesz, 1909, p. 52.

Anthrax inexactus Walker, 1857, p. 140.—Kert&z, 1909, p.

42.

Anthrax analis.—Macquart, 1840, p. 345 [not Say, 1823].

MALE (TYPICAL) .—Body mostly black, femora and

tibiae reddish brown or black; integument brownish

pruinose. Front with black setae and lanceolate black

scales, sometimes a few lateral yellow scales. Face with

black setae extending to antennae, a few black or white

scales below. Occiput with sparse black setae and

lanceolate or linear black scales, sometimes a few yellow scales; fringe of pile on posterior margin black.

First antennal segment about as long as apical width;

second segment globular; base of third segment slightly

compressed laterally, about equal to second segment

in lateral view, narrowing abruptly to styliform part

which is about 1.5 times longer than base and about 3

times longer than style (Figure 119).

Mesonotum and scutellum with fine black setae and

linear black scales, longer and suberect laterally on

mesonotum; a few white or yellow lateral scales on

mesonotum and scutellum. Mesopleuron, sternopleuron, and anterior part of pteropleuron with black setae,

pile, and scales. Pile on prosternum and propleuron

black, on anterior margin of mesonotum mixed black

and white; postalar tuft of pile black or mixed black

and white. Hypopleuron and metapleuron bare. Coxae

with black setae and black scales.

Wing (Plate 4;) light or dark brown basally, hyaline

apically, pigment typically extending out as far as base

of cell R4 in cell Ri, two-thirds of way from base of

R2+3 to base of R4 in cells R2+3 and R5, one-third of way

from r-m crossvein to base of Mx in cell 1M2, extreme

base of cell 2M2, and thence running to posterior margin in cell 2A, leaving cells 1A and 2A hyaline apically

for a distance up to two times length of r-m crossvein.

Stigmatic area lightly pigmented. Base of cell R4 evenly

rounded. R-m crossvein located at basal third or twofifths of cell 1M2; contact of cells 1M2 and Cui slightly

longer to slightly shorter than base of cell Cui. Cell

1A closed and sometimes petiolate. Alula distinctly

rounded posteriorly. Calypter lightly pigmented, fringe

of hairs white. Halter dark brown, tip of knob yellow.

Scales on legs cuneate and black; sometimes a few

yellow scales on femora and more often on tibiae.

Middle femur without macrochaetae anteroventrally

or with only one or two; hind femur with three to five

macrochaetae anteroventrally. Fore tibia with a com-
82

plete row of small macrochaetae anterodorsally; hind

tibia with single row of macrochaetae anterodorsally.

Pile laterally on first abdominal tergum black, white

or mixed; on terga two, three, and four black; some

white or yellow setae sometimes present on four; terga

five, six, and seven with white or black setae and some

white scales laterally. Discs of terga one through four

with hairlike black scales and fine setae; some cuneate

lateral white scales on four, some linear yellow scales

sometimes posteriorly on anterior terga. Discs of terga

five and six with yellow or white setae and cuneate

or lanceolate scales, yellow or white posterolaterally,

black anteromedially; scales on seventh tergum entirely

white or yellow, or black scales intermixed medially.

Abdominal sterna with fine black setae and hairlike

black scales, sometimes a few yellow posterior scales.

MALE GENITALIA (Figure 65).—Gonocoxites broad,

dorsal margins tapering convexly to acute apices in

lateral view; apices rounded apically in ventral view,

mesal margins rounded, mesal sulcus narrow; setae on

gonocoxites sparse, evenly distributed basally beyond

medial fold. Basal segment of gonostylus sharply angled

basally, rounded apically, concave dorsally before apex;

distal segment rounded basally, somewhat laterally

flattened, with a styliform apical part arising from the

dorsal half and slightly curving upward and outward

distally; sparse setae on ventrolateral surface. Apex of

epiphallus with slightly convex, raised dorsal plate

formed at about a 30° angle, the plate right angled

basally and slightly rounded apically in dorsal view,

curved downward laterally; ventrolateral part broadly

rounded ventrally, extending apically as far as middle

of dorsal plate. Dorsal bands united mesally. Aedeagus

narrow basajly, gradually tapering to junction with

ventral bands below junction of dorsal bands.

FEMALE.—Similar to male. Femora sometimes yellowish. Pigmentation of wing more extensive, reaching apex of vein Rx in cell R1} base of cell R* in cell

R2+3 and base of cell Mi in cell R4+5. Light scales on

abdomen restricted to lateral margins of fourth and

sometimes fifth and sixth terga.

FEMALE GENITALIA (Figure 90).—Tenth tergum

with only one spine on each side. Ventral arm of ninth

tergum narrow, apex not enlarged. Dorsomedial angle

of sclerite on each side of gonopore expanded dorsally

and sharply rounded; lateral arm broad, tapering to

upturned apex, ventral margin curled under; ventral

arm narrow, parallel sided, recurving mesally below.

Each spermathecal duct about 2.5 times longer than

bulb; first section very short, middle section about 1.5

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

times longer than straight third section; bulb globoid,

symmetrical, slightly contracted just before middle.

DISTRIBUTION AND ECOLOGY.—Anthrax repertus occurs throughout northeastern South America from Rio

Grande do Sul, Brazil, and central Paraguay to the

Caribbean Sea, north into Central America as far as

Honduras and south along the Pacific coast into Ecuador and Peru (Map 22). Specimens have been collected by the author in Rio de Janeiro, Guanabara;

Guaiba, Rio Grande do Sul; and Remanso, Bahia,

Brazil. The first locality was a high hill covered with

grass in the dominantly tropical forest of eastern Brazil; the second locality was a dry hillside covered with

grass and forbs in the predominantly grassland area of

southern Rio Grande do Sul; the third area was an

open sandy spot near the Rio Sao Francisco in the

"cerrado" of central Brazil.

TYPES.—According to Painter and Painter (unpublished notes) the type female of Anthrax reperta

Walker is in the British Museum. It carries the green

type label with "A. reperta Walk." on the back, another "68-4," and another "S. America pres. W. W.

Saunders, B. M. 1868-4." The head has been glued on,

but otherwise the type is well preserved. The photograph of the wing and redescription of the type show

it to be a typical specimen of the taxon described here.

The type of Anthrax inexactus Walker is also in the

British Museum. According to notes made by Painter

and Painter, the body is greasy and covered with

fungus and dust, the wings are mounted on a celluloid

square below the specimen, the head is glued on sideways, and the antennae and four and a half legs are

missing. It carries the green type label, another label

"Amaz," another "68-4," and a folded label with

"inexacta Wlk." in Walker's handwriting. A photograph of the wing and redescription indicate that this

specimen falls within the range of variation of repertus.

DISCUSSION.—Like macquarti, repertus exhibits

extensive variation throughout its range. The most

marked difference is in the color of the scales and

pile on the body. In one extreme, a male from Guaiba,

R. G. S., Brazil, has only a few white hairs on the

anterior margin of the mesonotum, a few white scales

on the posterior abdominal terga, and the normal white

fringe on the calypteres. In the other extreme, a specimen from Remanso, Bahia, Brazil, has all of the black

hairs and scales on the body replaced by yellow ones.

A specimen from Limoeiro, Ceara, is intermediate between the typical form and the specimen from

Remanso. Specimens from the coastal area of Peru
NUMBER 4 3 83

also have numerous yellow scales and hairs on the

body. In this case, two specimens from Ecuador are

intermediate between the typical and light-colored

forms.

The extent of the wing pattern is also highly variable. Specimens from areas of high humidity and rainfall usually have a pattern similar to that described

above. In some cases, however, the pattern may be

much more extensive, reaching the apex of cell R1}

extending into the base of cell R4 and extending past

the base of cell Mx in cell R5. Apparently, a specimen

with a pattern like this served as type for Walker's

inexactus which was described from the "Valley of the

Amazon." The only specimen studied from the lower

Rio Amazonas has wings with only slightly more

extensive pigment than that described for the typical

form. At the other extreme, specimens from desert

areas have less extensively pigmented wings. In this

MAP 22.—Distribution of Anthrax repertus.
84

case, the pigment may extend apically only one-third

of the way from the r-m crossvein to the base of cell

R4 in cells R2+3 and R5, and may extend in cell Rx only

halfway from the r-m crossvein to its tip. In these

specimens the pigment is also less extensive posteriorly,

leaving as much as the apical third of cells 1A and 2A

hyaline.

As in macquarti, this variation seems to be correlated

with climate, those species from drier areas having less

extensive wing pigment and lighter scales and hairs.

Although some of the variants may represent distinct

species or subspecies, further collecting will be required

to determine their true relationship. Since describing

the variants from only one or a few specimens now

would complicate the work of future students, the

entire range is included here under repertus.

Two additional distinctive specimens closely related

to repertus have been studied by the author. One, a

female from Lagoa Santa, Minas Gerais, Brazil, in

the collection of the author, has the margin of pigment

on the wing extending from below the tip of vein Sc

across the base of cell R5 to the base of cell Cux and

thence basally, leaving the apical third of cell 1A and

all but a narrow area along the anterobasal margin of

cell 2A hyaline. This specimen also has the pile and

scales on the thoracic pleura entirely white and has

some linear gold scales on the mesonotum, scutellum,

and posterior margin of the first abdominal tergum.

The other, also a female, is from Rio de Janeiro and

is in the collection of the Instituto Oswaldo Cruz. It

has the wing distinctly broader than in typical specimens of repertus and has the pigment entirely filling

cells 1A and 2A and extending out to the apical fourth

of cell Cui, whereas anteriorly it reaches only as far as

the tip of vein Sc. The margins of the front are nearly

parallel. It is much smaller than typical specimens of

repertus, the wing length being only 5.7 mm.

Anthrax repertus may be readily separated from

analis and gideon by the hyaline apices of cells 1A and

2A. It differs from macquarti and austrinus in having

the posterior margin of the alula distinctly rounded.

The relationship of repertus and clinopictus is discussed under the latter species.

Anthrax clinopictus, new species

MALE.—Integument mostly black, brownish pruinose, femora and tibiae yellow. Front with black setae

and lanceolate gold scales. FaCe with black setae extending almost to antennae, a few linear yellow scales

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

below. Setae on occiput black around vertex, yellow

elsewhere, scales gold; fringe of hairs on posterior margin yellow below, black exteriorly and yellow interiorly

above. First antennal segment about as long as apical

width; second segment globular, about as wide as base

of third segment in lateral view; base of third segment

slightly compressed laterally, narrowing abruptly to

styliform part which is about 1.5 times as long as base

and about twice as long as style (Figure 118).

Disc of mesonotum with fine black setae and mixed

black and gold, linear scales; lateral margins with

linear, semierect gold scales. Disc of scutellum with

fine black setae and mixed black and white, linear

scales; posterior and lateral margins with linear gold

scales. Mesopleuron, sternopleuron, and anterior half

of pteropleuron with fine, black and gold setae, and

linear gold scales; some coarse black and gold setae and

yellow hairs above on mesopleuron and pteropleuron.

Pile on prosternum and propleuron mixed yellow and

brown, yellow predominating on prosternum; pile on

anterior margin of mesonotum mixed yellow and

white; postalar tuft of pile yellowish white. Coxae with

black setae and linear gold scales. Hypopleuron and

metapleuron without scales or setae.

Wing (Plate 4A) dark brown basally, hyaline apically, the pigment filling all of cell Ri, cell R2+3 past

base of cell R4, cell R5 to base of cell Ml 5 basal threefifths of cell 1M2, extreme base of cell 2M2, and cell

Cu! out to base of cell 2M2, leaving apices of cells 1A

and 2A hyaline for a distance twice length of r-m crossvein. Stigmatic area very lightly pigmented. Base of

vein R4 evenly rounded; r-m crossvein placed at basal

third of cell 1M2; base of vein R2+3 slightly basad of r-m

crossvein; contact of cells 1M2 and Cux about equal to

base of cell Cui; cell 1A punctiform apically. Alula

rounded posteriorly. Calypter lightly pigmented, fringe

of hairs white.

Scales on femora and tibiae cuneate and lanceolate,

yellow ventrally, mixed yellow and black dorsally. Middle femur without macrochaetae anteroventrally; hind

femur with two to four macrochaetae anteroventrally;

fore tibia with a row of small macrochaetae anterodorsally; hind tibia with a single row of macrochaetae

anterodorsally.

First abdominal tergum with white pile laterally;

terga two through four with black hairs and setae

laterally, some yellow hairs present posteriorly on four;

terga five through seven with black and yellow setae

laterally. Posterior margin of terga one through three

with linear gold scales posteriorly, some long, white
NUMBER 43 85

and yellow, lanceolate scales laterally; remainder of

discs of terga two and three, and anterior half of fourth

tergum with hairlike black scales; terga five through

seven and posterior margin of fourth tergum with

ovate-truncate and lanceolate, white or yellowish-white

scales, the darker scales anterior on the segments, some

linear gold scales intermixed. Venter with sparse yellow setae and linear and lanceolate gold scales, a few

black scales anteriorly on the segments.

MALE GENITALIA (Figure 62).—Gonocoxites broad,

dorsal margins broadly rounded to sharp apices in

lateral view, rounded mesally to narrow mesal sulcus;

apices sharply rounded in ventral view; setae sparsely

distributed basally beyond medial fold. Basal segment

of gonostylus convex dorsally in lateral view; distal

segment bulbous basally, with styliform apex extending outward and upward from dorsal side, apex curved

dorsolaterally, sparse setae lateroventrally. Apex of

epiphallus with a raised dorsal plate convex dorsally

in lateral view, angulate basally and slightly emarginate apically in dorsal view, ventrolateral part curled

upward below, extending past middle of dorsal plate.

Dorsal bands united mesally. Base of aedeagus narrow,

tapering to junction with ventral bands below junction

of dorsal bands.

FEMALE.—Similar to male. Yellow and white scales

restricted to lateral parts of posterior abdominal terga,

centers of discs with anterior linear black scales and

posterior linear gold scales.

FEMALE GENITALIA (Figure 91).—Tenth tergum

with eight spines on each side. Ventral arm of ninth

tergum undulate and not enlarged apically. Dorsomedial corner of sclerite on each side of gonopore

produced dorsally as a narrow, apically rounded lobe;

lateral arm short, turned upward to extreme apex

which is bent posteriorly. Each spermathecal duct

about four times longer than bulb; first section very

short; second section about two-thirds as long as apical

section which is about twice as long as bulb and expanded distally; bulb asymmetrical, elliptical with

apex bent to one side, and not well defined from duct.

VARIATION.—The scales on the front may be yellow

rather than gold and there may be a few dorsal black

scales. The setae on the occiput may be entirely black.

The scales on the mesonotum vary from light to dark

gold. The hairs on the prosternum may be entirely

yellow or predominantly light brown. The middle

femur may have two or three macrochaetae anteroventrally.

DISTRIBUTION.—Specimens of clinopictus have been

collected from several localities in Sao Paulo, and from

western Parana and northern Argentina. It probably

also occurs in southern Minas Gerais and possibly in

southwestern Rio de Janeiro. A typical specimen of

repertus was collected along with clinopictus at the

type-locality.

HOLOTYPE.—c?j Onda Verde, Faz. Sao Joao, Sao

Paulo, Brazil, 1-1946 (F. Lane) (SASP).

ALLOTYPE.— $ , same data as holotype (SASP).

PARATYPES.—Argentina: cf, $ , "Tapikiol6," XII25-1-26 (Lind.; D. Chaco Exped.) (Staat).

Brazil: Parana, d, Iguassu, XII-1941 (Com. E.

N. V.) (IOC). Sao Paulo, 2c?,? , Onda Verde, Faz.

Sao Joao, 1-1946 (F. Lane) (SASP, NLM); d,

Perube, XII-1946 (M. Carrera) (SASP); ? , Praia

Grande, Faz. Rondonea, 11-1945 (M. Carrera)

(SASP); d, Ubatuba, VI-1955 (F. Lane) (SASP).

DISCUSSION.—The diagnostic characters of clinopictus are in all cases ones which occur in one or more

of the variants of repertus. Only in clinopictus, however, are they found in combination. Since this particular combination of characters is found in both males

and females at several localities with similar habitats,

they likely represent a genetic population distinct from

repertus. Since a typical specimen of repertus was collected along with specimens of clinopictus at the typelocality and no intermediates have been seen from

within the range of clinopictus, it is being described as

a distinct species.

The male genitalia of clinopictus differ from those of

repertus in that the ventral margin of the ventrolateral

part of the apex of the epiphallus is curled outward

and upward. The female genitalia differ in having nine

bristles on each side of the tenth tergum, rather than

only one, and in having the bulb of each spermatheca

asymmetrical.

Anthrax gideon Fabricius

Anthrax gideon Fabricius, 1805, p. 124.—Wiedemann, 1828,

p. 311.—Macquart, 1840, p. 342.—Osten Sacken, 1858,

p. 40.—Curran, 1934, p. 363.—Wolcott, 1951, p. 450.

Argyramoeba gideon.—Schiner, 1868, p. 122 [Argyromoeba].—Osten Sacken, 1878, p. 90.—Kertesz, 1909, p. 64.

Spongostylum gideon.—Aldrich, 1905, p. 233 [Spogostylum].—Johnson, 1913, p. 56 [Spogostylum].

Anthrax acroleucus Wiedemann, 1828, p. 312 [acroleucd].—

Curran, 1934, p. 363 [acroleuca].

Argyramoeba acroleuca.—Osten Sacken, 1886, p. 101.—Kert6sz, 1909, p. 59.
86

Spongostylum acroleucum.—Williston, 1901, p. 275 [Spogostylum acroleuca].—Aldrich, 1905, p. 222 [Spogostylum acroleuca].—Johnson, 1913, p. 56 [Spogostylum acroleuca].

Argyramoeba sp.—OstenSacken, 1886, p. 101.

MALE.—Integument mostly black, legs and pleura

often brown; front, mesonotum, scutellum, and

abdominal terga velvet black or brown pruinose, remainder of body brown or gray pruinose. Setae and

scales on front black; face with black setae extending

to antennae, a few white hairs ventrally. Setae and

scales on occiput black; fringe of pile on posterior margin black. First antennal segment about as long as

apical width; second segment globular; base of third

segment globular (often collapsed), slightly smaller

than second segment, narrowing abruptly to styliform

part which is about one-half longer than base and

about one-third longer than style (Figure 117).

Discs of mesonotum and scutellum with short black

setae and threadlike black scales; lateral and anterior

margins of mesonotum with longer black hairs, erect

linear scales, and black setae. Sternopleuron, mesopleuron, and anterior half of pteropleuron with fine

black setae and threadlike black scales, some dorsal

long black hairs and coarse black setae on mesopleuron

and pteropleuron. Prosternum and propleuron with

black pile; postalar tuft of pile black. A patch of black

hairs often behind spiracle. Coxae with black hairs,

setae, and threadlike scales.

Wing (Plate 4t) pigmented velvet black or brown

basally, hyaline apically; apical margin of pigment distinct, running from tip of vein Rj across base of cell

R4 to vein Mx slightly beyond base of cell Mx, basally

on vein R5 to slightly beyond r-m crossvein and thence

curving posteriorly through extreme base of cell 2M2 to

tip of vein 1A; usually a narrow hyaline area posteriorly

in cell R5. Basal angles of veins R* and R2+3 without

spurs, or having spurs shorter than basal segments of

veins. R-M crossvein located at basal third or two-fifths

of vein M1+2; contact of cells 1M2 and Cux about equal

to base of cell Cui. Posterobasal part of wing not narrowed, cell 2A slightly wider than cell 1A, alula

rounded posteriorly. Apex of wing often milky colored

as far back as tip of vein R^ Stigmatic area heavily

pigmented. Calypter pigmented, fringe of hairs usually

white in specimens from northern South America and

Central America, usually black or brown in specimens

from southeast Brazil. Halter brown, extreme tip of

knob yellow.

Scales on legs black. Middle femur without macrochaetae anteroventrally or with only one or two short

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ones; hind femur with three to five weak macrochaetae

anteroventrally; hind tibia with single row of macrochaetae anterodorsally.

Sides of abdominal terga with dense black pile;

sterna and discs of terga with sparse black setae and

black, threadlike scales.

MALE GENITALIA (Figure 63).—Gonocoxites broad,

dorsal margins curving concavely to ventral margins to

form acute apices in lateral view; apices broadly

rounded in ventral view, with sharp ridges mesally.

Basal segment of gonostylus sharply pointed basally,

convex toward apex in lateral view; distal segment

round in cross section basally, tapering apically; sharp

longitudinal ridge dorsally in lateral view; apex sharply

bent upward and outward, or straight in some specimens from eastern Brazil. Setae on gonocoxites extending almost to base, evenly distributed, absent at extreme apices; setae over interior and ventral surfaces of

basal part of distal segment of gonostylus. Apex of

epiphallus with a basally acute, apically emarginate

dorsal plate formed at about a 30° angle; ventrolateral

part broad, not extending to apex of dorsal plate; dorsal bands very narrowly separated medially, without

setae. Base of aedeagus broad, but not bulbous, tapering only slightly to junction with ventral bands below

junction of dorsal bands.

FEMALE.—Similar to male. Fringe of hairs on posterior margin of occiput sometimes white below. Pile

on prosternum, propleuron, anterior half of mesopleuron, and anterior margin of mesonotum occasionally partly or wholly white. Apices of wings never

milky. Sides of first and anterior half of second abdominal terga often with white pile. Sides of fourth

tergum occasionally with a patch of long, overlapping,

acute silver scales, and a few white seta.

FEMALE GENITALIA (Figure 94).—Tenth tergum

with about 18 spines on each side. Ventral arm of

ninth tergum narrow, parallel sided and nearly straight,

apex not enlarged. Dorsomedial angle of sclerite on

each side of gonopore produced mesally as a short

narrow, truncate lobe; lateral arm broad basally, tapering to an acute apex, curled under below; ventral arm

narrow and curved sharply inward below. Each spermathecal duct about four times as long as bulb; first

section very short; middle section slightly shorter than

apical section which is about twice as long as bulb and

broadly expanded almost to width of bulb apically;

bulb asymmetrical, slightly longer than maximum

width, constricted at junction with duct
NUMBER 43 87

DISTRIBUTION AND ECOLOGY.—Anthrax gideon occurs from southern Mexico into the Guianas and Bolivia in northwestern South America, and in the Serra

do Mar in southeast Brazil (Map 23). Two specimens

MAP 23.—Distribution of Anthrax gideon.

were collected by the author in Rio de Janeiro, Brazil,

along the edge of an open sand and gravel area in the

tropical forest covering the mountainous southwestern

part of the city. No specimens of this species have

been seen from the Amazon Basin, nor from the extensively collected Barro Colorado Island in the Canal

Zone, Panama. This may indicate that gideon inhabits

areas of greater relief, which would explain its discontinuous distribution.

TYPES.—According to notes made by Painter and

Painter, there are two females of Anthrax gideon Fabricius in the Universitets Zoologiske Museum, Copenhagen. One carries the red type label only, while the

other carries the red type label and another label reading "A. gideon ex Am. Mer. Schmidt." A homotype

compared with the type is typical of female specimens

from Central America and northwestern South

America.

Painter and Painter (personal notes) found a series

of syntypes of Anthrax acroleuca consisting of five

males and one female in the Vienna Museum. All are

apparently from Brazil. No lectotype was chosen since

none of the specimens was perfect nor complete. A

homotype compared with the type series is typical of

specimens from southeast Brazil.

The type female of Argyramoeba propinqua Schiner

is in the Vienna Museum. According to notes made by

Painter and Painter, it carries the labels "Venezuela"

and "propinqua Alte Sammlung." The specimen is

partly glued together and greasy, with the antennae

plastered down or missing, the last pair of legs missing,

and one wing broken. The photograph and redescription of the type indicate that it is a typical specimen

of the form propinquus discriminated below.

DISCUSSION.—Anthrax gideon is a highly variable

species. In general, however, those from southeast Brazil show much less variation than those from the remainder of the range. Specimens from the coastal area

of Brazil usually have the pile and scales on the body

entirely black in both sexes, although the pile laterally

on the first abdominal tergum is often white in females.

The pruinosity on the dorsal parts of the body is velvet

black and the wing pigment is velvet black. The wing

pattern in this area is quite stable, like that shown in

Plate 4i.

Specimens from northwest South America, Central

America, and southern Mexico, on the other hand,

show a wide range of variation. The wing pattern

varies from the condition of the form propinquus to a

condition approaching that of analis. The pigment is

typically velvet black, but may be brown in specimens

from Colombia, Venezuela, and British Guiana. The

pruinosity on the dorsal surface of the body also is

typically velvet black, but may be brown on those

specimens with lighter wings. Many males from the

northwest part of the distribution have the fringe of

pile on the calypteres white, a condition found in only

one Brazilian specimen. The females often have extensive white or yellowish white pile on the anterior

part of the thorax and the lateral parts of the first abdominal tergum, and patches of white scales laterally

on the fifth tergum.

Throughout the range of gideon, some of the males

have the apex of the wing milky white rather than

hyaline. No females have been examined that show

this condition.

Wiedemann (1828) described the Brazilian form

with entirely black hairs and scales as acroleucus. Since

the extent of variation in these characters broadly overlaps in the two areas, they are regarded here as the

same species. It seems probable that the population in

southeast Brazil was derived relatively recently from an

ancestral population in the area of Bolivia with a low

incidence of specimens with white pile and scales.
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Anthrax gideon i. propinquus (Schiner)

Argyramoeba propinqua Schiner, 1868, p. 123.—Kertesz,

1909, p. 66.

Two males and two females from Fusagasuga, Cundin

Amarca, Colombia, have a wing pattern like that of

Argyramoeba propinqua Schiner. The pigment extends in cell R5 more than twice the length of the r-m

crossvein past the base of cell Mi and extends slightly

into the base of cell Ma. These specimens agree in all

other respects with the redescription of the type made

by Painter and Painter. Although no type-locality was

given in the original description, the type is labeled

"Venezuela." As in the forms of Anthrax analis, this

form does not diverge in other respects from the normal range of variation in this area. For this reason, and

since a male with normal wing pattern has been studied

from Caquesa, Cundin Amarca, propinquus is regarded

here as simply a form of gideon. A long series of specimens will be required to determine conclusively the

true relationship of the taxa.

The relationship of gideon and analis is discussed

under the latter species. Anthrax gideon differs from

funebris in that the pigment on the wing does not fill

more than half of cell 1M2, and differs from the other

members of the analis complex in having cells 1A and

2A entirely pigmented, or with only the extreme apex

of cell 1A subhyaline.

Anthrax analis Say

Anthrax analis Say, 1823, p. 45.—Osten Sacken, 1858, p.

39.—Johnson, 1925, p. 108 [anale].—Curran, 1927, p. 85.—

Painter, 1930, p. 794.—Painter, 1933, p. 5 [anale].—

Maughan, 1935, p. 32.—Brimley, 1938, p. 341.—Strickland, 1938, p.195.

Argyramoeba analis.—Osten Sacken, 1877, p. 242.—Osten

Sacken, 1878, p. 89.—Osten Sacken, 1886, p. 101.—Coquillett, 1894, p. 95.—Johnson, 1895, p. 325.—Rau, 1926, p.

231 [Argyromoeba anale].

Spongostylum anale.—Aldrich, 1905, p. 222 [Spogostylum].—

Johnson, 1913, p. 55 [Spogostylum].—Shelford, 1913,

p. 213 [Spogostylum].—Malloch, 1915, p. 328 [Spogostylum]—Cole and Lovett, 1921, p. 243 [Spogostylum].—

Cole, Malloch, and McAtee, 1924, p. 185 [Spogostylum].

Anthrax georgicus Macquart, 1834, p. 406 [georgica].—Macquart, 1840, p. 346 [georgica].—Osten Sacken, 1858, p.

40.— Painter and Painter, 1962, p. 75.

Argyramoeba gideon.—Coquillett, 1894, p. 95 [not Fabricius,

1805].

Spongostylum gideon. Aldrich, 1905, p. 223 [Spogostylum]

[not Fabricius, 1805].

Spongostylum occidentalis.—Johnson, 1913, p. 56 [Spogostylum].

MALE.—Body black, tibiae sometimes red or yellow,

femora sometimes red; integument light and dark

brown pruinose. Front with black setae and scales;

face with black setae extending almost to antennae.

Occiput with sparse black setae and a few black scales;

fringe of hairs on posterior margin black. First antennal segment about as long as apical width; second segment biscuit shaped, flattened apically; base of third

segment flattened laterally, narrowing abruptly to

styliform part, about as wide as second segment; styliform part about one-half longer than base, two to

three times longer than style (Figure 116).

Discs and lateral margins of mesonotum and scutellum with linear black scales. Sternopleuron, lower

half of mesopleuron, and anterior part of pteropleuron

with black setae and linear black scales; upper half of

mesopleuron with black pile and fine and coarse black

setae. Prosternum, propleuron, and anterior margin

of mesonotum with black pile, some white pile often

present on latter; postalar tuft of pile black, or rarely

mixed black and white. Coxae with black bristles and

scales.

Wing typically dark brown basally, hyaline apically

(Plate 3a); pigment extending out to a line from tip

of vein Rx across cell R^ slightly basad along vein R2+3

and perpendicularly across cells R2+3 and R5, or diagonally across cells R2+3 and R5 about to bifurcation of

vein M1+2, along vein Mi+2 halfway to r-m crossvein,

and thence posteriorly to tip of cell 2A; extreme tip of

cell 1A sometimes hyaline. Base of cell R4 evenly

rounded; r-m crossvein at basal two-fifths of cell 1M2,

vein R2+3 arising opposite. Contact of cells 1M2 and

Cui slightly shorter or longer than width of base of

cell Cui; cell 1A closed at margin or short petiolate;

cell 2A not narrowed, wider than cell 1A postmedially;

alula not reduced, posterior margin rounded. Stigmatic

area pigmented. Calypter pigmented, fringe of hairs

white or brown. Stem of halter brown, knob brown

basally, yellow apically.

Legs with black scales. Middle and hind femora

with one or two macrochaetae anterodorsally, or with

only a few toward the apex; hind tibia with a single

or double row of macrochaetae anterodorsally.

Sides of first abdominal tergum with black, white,

or mixed pile; terga two through four, and sometimes

five, six, and seven with black pile, setae and scales

laterally. Posterior margin of first tergum with linear

black scales; discs of terga two through four, and middle

of five with linear black scales; remainder of terga with

dense or sparse, linear or lanceolate, truncate, pos-
NUMBER 43 89

teriorly produced, white scales sometimes restricted to

lateral margins and occasionally absent. Venter with

black setae and linear scales.

MALE OENITALIA (Figure 64).—Gonocoxites very

broad basally, dorsal margins tapering concavely to

acute apices in lateral view; apices bluntly rounded in

ventral view, mesal sulcus very narrow; setae evenly

distributed as far as midway between medial fold and

base. Basal segment of gonostylus elongate basally,

slightly convex dorsally in lateral view; distal segment

round basally in cross section, tapering to acute apex

which is slightly bent upward and outward; ventral

margin curved sharply upward before styliform part;

setae present laterally and ventrally on basal part. Apex

of epiphallus with a raised dorsal plate formed at a

45° angle, plate right angled basally, truncate apically

in dorsal view; ventrolateral part curved inward below,

extending out as far as middle of dorsal plate. Dorsal

bands united mesally, without setae. Aedeagus narrow,

tapering to junction with ventral bands before junction

of dorsal bands.

FEMALE.—Similar to male. White scales on posterior

abdominal terga restricted to lateral margin of fifth

tergum or absent.

FEMALE GENITALIA (Figure 93).—Tenth tergum

without spines. Ventral arm enlarged and bilobate apically. Dorsomedial angle of sclerite on each side of

gonopore produced dorsointeriorly as a short, sharp,

acuminate lobe; lateral arm tapering and curved upward apically; ventral arm broad basally, tapering to

sharp apex which is slightly curved mesad. Each

spermathecal duct about as long as bulb; first section

very short; second section enlarged apically, about onefourth longer than third section which is about half

as long as bulb; bulb ovoid, narrowing basally to duct

and somewhat constricted premedially.

DISTRIBUTION AND ECOLOGY.—Anthrax analis occurs throughout North America from northern Canada

to Central America (Map 24). It is found in areas of

sand dunes, along beaches, and in sandy or gravelly

areas along rivers. It seems to be replaced in the forested

areas of Mexico and Central America by gideon, although their distributions broadly overlap.

TYPES.—The type of A. analis Say apparently has

been lost along with the remainder of Say's collection.

Say's description, however, leaves little doubt as to

the identity of the species. The type-locality is Georgia.

Painter and Painter (1962) found the type of

Anthrax georgicus Macquart, a female, in the Ville

de Lille, Museu d' Histoire naturelle. The head has

been glued on, the antennae are nearly gone, and it

has been in liquid at some time. The photograph of

the wing is like that of a typical specimen of A. analis.

The type-locality is Georgia.

The type female of Anthrax cedens Walker is in

the British Museum. According to Painter and Painter

(1962), it "bears the green type label with 'A cedens

Walk.' on back, a second label '68-^J-,' a small yellow

label with no writing, and 'U.S.A. pres. W. W. Saunders 1868-4.' " A photograph of the wing of the type

shows it to be a rather heavily pigmented female like

those discriminated below.

Painter and Painter (1962) stated that there were

three female specimens in the Bigot collection in the

British Museum under the name Hemipenthes latelimbatus. One specimen carried the pin label "Hemipenthes latelimbatus m, f, n. sp. Inedit. Avril 1889 J.

Bigot Jun (?) Carolin." The best preserved specimen

was selected as a lectotype. The photograph of the

wing and description given by Painter and Painter

clearly indicate that this species is synonymous with

A. analis, f. cedens Walker.

The type male of Spongostylum grossbecki Johnson

is deposited in the collection of the American Museum

of Natural History and has been studied by the author.

It lacks the head but is otherwise in excellent condition and is typical of males of the form grossbecki

discussed later. Johnson (1913) erroneously referred

the female of this form to A. cephus Fabricius. The

type-locality is Lakeland, Florida.

The cotypes of Spongostylum occidentalis Johnson,

from Denver, Colorado, and Seattle, Washington, are

in the Museum of Comparative Zoology and have been

seen by the author. Johnson (1913) named this species

in a key to the species related to analis in North and

South America but did not give a detailed description.

He implied that one of the specimens was a male, although both are females. The specimen from Denver

was in good condition except that one of the forelegs

and the last segment of the antenna were missing. The

specimen from Seattle was missing the head, one foreleg, and one hind leg, and was slightly rubbed. Both

specimens fall within the range of variation of analis

in the western United States.

BIOLOGY.—Shelford (1913) gave an excellent account of the life history and habits of analis which he

reared from the burrows of "Cicindela scutellaria Say,

var. lecontei Hald." (Coleoptera: Cicindellidae). Malloch (1915) described the larva and pupa, utilizing

specimens collected by Shelford. Anthrax analis has
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MAP 24.—Distribution of Anthrax analis (horizontal lines) and A. funebris (vertical lines).
NUMBER 43 91

been collected many times by the author in sandy

areas where the females were seen ovipositing in holes

in the sand. This species may parasitize bees and wasps

nesting in such areas, in addition to tiger beetles.

DISCUSSION.—The extent of the pigment on the

wings is variable in specimens from Florida and the

coastal plain of the southeastern United States. It

seems probable that a population was isolated in Florida in the past which evolved more extensively pigmented wings similar to those of the form grossbecki

separated below. When this population came in contact

with the mainland population again, hybridization took

place so that now a gradient from normal to heavily

pigmented forms occurs in that area. This fits the

evolutionary scheme given by Hubbell (1954). The

forms do not differ in other characters and they are all

found sympatrically at the present time. Specimens

that have the area of pigment slightly expanded also

have been collected as far north as New York and as

far west as Alabama. The two principal forms are

distinguished below.

Anthrax analis f. grossbecki (Johnson)

Anthrax cephus.—Macquart, 1840, p. 337.—Osten Sacken,

1858, p. 39 [part] [not Fabricius, 1805].

Argyramoeba cephus.—Osten Sacken, 1878, p. 90 [part].—

Coquillett, 1894, p. 95.—Johnson, 1895, p. 325 [part]

[not Fabricius, 1805].

Spongostylum cephus.—Johnson, 1913, p. 56 [Spogostylum]

[not Fabricius, 1805].

Spongostylum grossbecki Johnson, 1913, p. 56 [Spogostylum].

In this form, the wing of the female is completely black

(Plate 3c) and the wing of the male has only a narrow

hyaline area along the posteroapical margin (Plate 36).

It is restricted to Florida. When Johnson (1913) described grossbecki from the male, he erroneously referred the female to cephus Fabricius.

Anthrax analis f. cedens (Walker)

Anthrax cedens Walker, 1852, p. 190.—Osten Sacken, 1858,

p. 39.—Osten Sacken, 1877, p. 238.—Kertesz, 1909, p.

31.—Painter and Painter, 1962, p. 72.

Spongostylum cedens.—Johnson, 1913, p. 56 [Spogostylum].

Hemipenthes latelimbatus Bigot, 1892, p. 351.

Argyramoeba latelimbata.—Johnson, 1895, p. 325 [latilimbata].

Anthrax latelimbatus.—Aldrich, 1905, p. 232 [latelimbata].—

Painter and Painter, 1962, p. 75.

Spongostylum latelimbatum.—Johnson, 1913, p. 55 [Spogostylum latelimbata].

In the female of this form, the pigment of the wing

extends well beyond the base of cell R* and extends

from cell Rs into or across cell M1; leaving the tip

of cell 1M2 hyaline or subhyaline (Plate 3e). In the

male, the pigment is less extensive, but extends broadly

into the base of cell R4 (Plate 3d). This form occurs

from Florida north along the Atlantic coast to New

York and west along the Gulf coast to Alabama.

Anthrax analis is very closely related to gideon. Most

specimens of gideon have cell 1M2 less than half pigmented and have the pigment in cell R8 extending as

far as the base of cell Mi, but some specimens have

been noted with a wing pattern similar to that of

analis in that the pigment fills more than half of cell

1M2. In this case, the velvet black pruinosity on the

dorsum of the thorax and abdomen of gideon may

serve to separate the species.

Specimens with wings like those of analis and with

other characters like those of gideon have been seen

from several localities in Mexico, Central America,

and northwestern South America. These may be only

a form of gideon or they may be intermediates between gideon and analis. The first possibility seems

most likely since the variant form has been collected

as far south as Peru, far beyond the range of analis.

It is possible that as in Florida the genetic potentiality

for the intermediates originated from crosses between

reunited populations. Extensive collecting and ecological observation may be necessary to determine the true

relationship of the populations of gideon and analis.

Anthrax analis has been confused with argentatus

from northwestern United States, but may be separated

by the absence of patches of silver scales posterolaterally

on abdominal terga two and three. The black-winged

females of the form grossbecki have been confused

with cephus and koebelei. They differ from the former

in lacking hairs or setae on the anterior corner of the

hypopleuron, and from the latter in having the styliform part of the third antennal segment at least 1.5

times as long as the base and in having the integument

dull and heavily pruinose.

Anthrax funebris Macquart

Anthrax funebris Macquart, 1840, p. 344.—Osten Sacken,

1858, p. 40.—Aldrich, 1905, p. 231.—Kertesz, 1909, p.

38.—Painter and Painter, 1962, p. 75.

FEMALE.—Integument generally black, pleura and

legs sometimes reddish brown; integument grayish

to brownish pruinose. Front with black setae and
92

lanceolate black scales. Face with black setae extending almost to antennae, a few white scales below. Occiput with short black setae and linear black scales;

fringe of hairs on posterior margin black. First antenna!

segment about as long as apical width; second segment

globular, about as wide as base of third segment in

lateral view; base of third segment slightly compressed

laterally, narrowing abruptly to styliform part which is

one-half longer than base and about twice as long

as style (Figure 115).

Discs of mesonotum and scutellum with fine black

setae and linear scales, some longer, semierect scales

and coarse setae laterally on mesonotum. Sternopleuron, mesopleuron, and anterior half of pteropleuron

with fine black setae and linear scales, some black hairs

and coarse setae dorsally on mesopleuron and pteropleuron. Pile on prosternum, propleuron, and anterior

margin of mesonotum black, white hairs sometimes on

propleuron and anterior margin of mesonotum; postalar tuft of pile black. Coxae with black setae and linear

black scales. Hypopleuron and metapleuron bare.

Wing (Plate 4k) pigmented with dark brown,

hyaline areas subapically in cell R2+3, apically in cells

R4, Cul5 and 1M2; a narrow hyaline area sometimes

along posterior border connecting hyaline areas in

cells R4 and Cui, and sometimes connected to hyaline

area in cell 1M2 across m crossvein. Base of cell R4

edged by subhyaline. Base of vein R4 evenly rounded;

r-m crossvein located at basal two-fifths of cell 1M2;

contact of cells 1M2 and Cux equal to or slightly greater

than base of cell Cux; cell 1A punctiform apically.

Alula well developed, posterior margin rounded. Stigmatic area lightly pigmented, sometimes only partially.

Calypter lightly pigmented; fringe of pile white. Halter

dark brown, tip of knob yellow.

Femora and tibiae with lanceolate and cuneate

black scales. Middle femur with one or two macrochaetae anteroventrally; hind femur with three or four

macrochaetae anteroventrally; fore tibia with incomplete row of macrochaetae anterodorsally; hind tibia

with single row of macrochaetae anterodorsally.

First abdominal tergum with white, pile laterally;

terga two through four with black pile and setae laterally; terga five through seven with black setae laterally.

Posterior margin of first tergum and discs of posterior

terga with fine, black setae and linear, black scales; a

few white scales sometimes laterally on four and five.

Abdominal sterna with fine, black setae and linear,

black scales.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FEMALE GENITALIA (Figure 92).—Tenth tergum

with one or no spine on each side. Ventral arm of

ninth tergum straight, slightly wider medially than at

acute apex. Dorsomedial corner of sclerite on each side

of gonopore produced dorsally as a short, sharply

angled lobe; lateral arm narrow and curved upward

apically; ventral arm narrow, twisted and slightly recurved mesally below. Each spermathecal duct about

twice as long as bulb; first section very short; second

section about as long as third, which is abruptly broadened apically to junction with bulb; bulb symmetrical,

globoid, somewhat longer than maximum width and

darker than duct.

DISTRIBUTION.—Specimens of A. funebris have been

studied from Haiti, Dominican Republic, and South

Caicos Island in the Bahamas. It may occur also

throughout the Greater Antilles and Bahamas, and

possibly in the Lesser Antilles (Map 24).

N EW MATERIAL EXAMINED.—Bahama Islands: Turks

and Caicos Islands, ? , South Caicos Island, II—11-

1953 (E. B. Hayden and G. B. Rabb).

Dominican Republic: ? , "St. Dom., 51.1."

Haiti: ? , Bois Caradeux, VIII-10-1934 (E. M.

Ducasse); ? , Pivert, IV-1-1922, about 250 feet,

"F. 4657" (C. H. Curran, Ace. 31144); $ , St. Marc,

III-30-IV-2-1922, "F. 4655."

TYPE.—Painter and Painter (1962) stated that the

type of Anthrax funebris Macquart was in the Paris

Museum. Only part of the thorax and most of

one wing remained. It carried the museum labels

"A. funebris Macqt. St. Domingue, 1607," and the pin

labels "No. 907, Anthrax funebris" and "A. in St.

Domingue." The distinctive wing pattern left little

doubt as to the identity of the species.

DISCUSSION.—Anthrax funebris is most closely

related to analis and closely resembles specimens of

the form cedens. In funebris the pigment on the wing

extends from the tip of cell Rx across the apex of cell

R2+3 leaving a subapical hyaline area in cell R2+3. In

analis, if the pigment extends from the tip of cell R4

into cell R2+3, the latter cell is entirely pigmented.

All of the specimens of funebris examined are female.

If funebris has the same kind of sexual dimorphism

shown by the forms cedens and grossbecki of analis, the

males will have less extensively pigmented wings.

Anthrax koebelei, new species

MALE.—Body generally black or brown, legs other

than coxae reddish brown; integument shining, sparsely
NUMBER 4 3

covered with gray and brown pruinosity, eye margins

and lower half of occiput silver pruinose. Setae and

scales on front black; setae on face mixed black and

white extending to antennae. Dorsal part of occiput

with black setae, ventral part with white setae and a

few white scales; fringe of hairs on posterior margin

black above, mixed black and white below. First antennal segment slightly longer than apical width, subcylindrical and angled toward meson; second segment

large, globoid, slightly smaller than length of first segment and distinctly wider than diameter of base of

third segment in mesal view; base of third segment

flattened laterally, narrowing abruptly to styliform

part about equal in length to base and style.

Mesonotum and scutellum with black setae and

scales; a few white scales in front of wing. Pile and

setae on mesopleuron, sternopleuron, and anterior part

of pteropleuron white. Hypopleuron and metapleuron

bare. Pile on prosternum, propleuron, and anterior

margin of mesonotum white. Postalar tuft of pile white.

Coxae with black and white setae and white scales.

Wing entirely translucent brown. Base of cell R4

angular, with a short spur; base of cell R2+3 slightly

basad of r-m crossvein; contact of cells 1M2 and Cui

about equal to base of cell Cui. Apex of cell 1A narrowly open. Cell 2A about as broad as cell 1A; alula

well developed, posterior margin rounded. Stigmatic

area in cell M reduced in size, not reaching apex of cell,

not pigmented. Calypter pigmented light brown, fringe

of hairs black. Base of halter dark red, knob dark red

basally, yellowish at extreme apex above, mostly red

below.

Scales on legs black, hind femur with two macrochaetae anteroventrally toward apex; middle femur

without strong macrochaetae. Fore tibia with anterodorsal, posterodorsal, and posteroventral rows of macrochaetae; hind tibia with a single row of macrochaetae anterodorsally.

Pile and setae on sides of first abdominal tergum

yellowish white; pile and setae on lateral margins of

terga two through seven mostly white, with a few

black setae. Discs of terga two through five with

black and white setae and linear black scales, some

white scales laterally on five; terga six and seven with

long, narrowly lanceolate, silvery-white scales. Venter

with sparse white setae.

MALE GENITALIA (Figure 55).—Gonocoxites broad

basally, dorsal margins curving gradually to ventral

margins apically in lateral view; apices bluntly rounded

in ventral view, and narrowly separated medially;

93

setae sparse and evenly distributed beyond medial

fold. Basal segment of gonostylus poorly developed,

with an obtuse angle dorsally in lateral view, bluntly

rounded apically and scarcely visible in ventral view.

Distal segment of gonostylus bulbous basally with

styliform apical part; apex of styliform part curved

upward and outward to a sharp point; basal part with

numerous fine setae. Apex of epiphallus with a broad

dorsal shield, slightly emarginate apically and with

three blunt teeth basally in dorsal view, angled downward at about 30° with apical part slightly depressed

in lateral view; ventrolateral part formed behind and

below dorsal shield, with nearly straight ventral margin about twice as long as slightly curved anterior

margin in lateral view. Dorsal bands not uniting

mesally, without setae; ventral bands uniting medially

over aedeagus and below juxtaposition of dorsal bands.

Aedeagus narrow basally, tapering gradually to junction with ventral bands; apex not visible in either

dorsal or lateral view.

FEMALE.—Similar to male. Pile, setae, and scales on

body entirely black.

VARIATION.—The only other specimen of this species

available, a male, is smilar in every respect to the holotype except that the fore femur has some white scales

posteriorly and the posterior cells of the wing are subhyaline medially.

DISTRIBUTION.—The three specimens available of

this species are simply labeled "Arizona A. Koebele,"

and "2542."

HOLOTYPE.—cf, Arizona (A. Koebele) (2542)

(CAS).

ALLOTYPE.— $ , same data as holotype (CAS) .

PARATYPE.—c?, same data as holotype (NLM).

BIOLOGY.—All three specimens of koebelei were

reared, but there are no data with the specimens.

DISCUSSION.—Anthrax koebelei appears to have no

close relatives in North and South America, but it

may be confused with other species with entirely black

wings. It may be distinguished readily from cephus and

aterrimus by the absence of setae or hairs at the anterior

corner of the hypopleuron. It differs from analis f.

grossbecki in having the basal part of the third antennal

segment about as long as the style rather than about

half as long.

Anthrax inaquosum, new species

FEMALE.—Body black, tibiae yellow; integument

mostly gray and brown pruinose; margins of eyes, face,
94

and lower part of occiput silver pruinose. Front with

black setae; face with white setae extending almost to

antennae. Occiput with black setae on dorsal third,

remainder with white setae, fringe of hairs on posterior

margin black exteriorly and white interiorly above,

white below. First antennal segment much shorter than

apical width; second segment lenticular; base of third

segment flattened laterally, distinctly broader than

second segment in mesal view, narrowing abruptly to

stylif orm part which is short, about as long as base and

slightly shorter than style (Figure 100).

Discs of mesonotum and scutellum with linear yellow scales, margins with white scales. Sternopleuron,

mesopleuron, and anterior half of pteropleuron with

white setae and hairs and lanceolate white scales.

Prosternum, propleuron, and anterior margin of mesonotum with white pile. Postalar tuft of pile white.

Hypopleuron and metapleuron bare. Coxae with white

bristles and lanceolate white scales.

Wing (Plate 3k) mostly hyaline, cells C and Sc

subhyaline, faint light-yellow spots at bases of cells

R2+3, Rs, 1M2, Cui, Ri, and in cell R below base of Rx;

base of wing lightly pigmented. Base of cell R* angled,

without spur. R-m crossvein at basal two-sevenths of

cell 1M2, vein R2+3 arising slightly basad. Contact of

cells 1M2 and Cux 3.5 times longer than width of base

of cell Cui. Anal cell open for distance equal to onehalf width of r-m crossvein. Posterobasal part of wing

not narrowed, cell 2A slightly wider than cell 1A postmedially; alula distinctly rounded posteriorly. Calypter

not pigmented, fringe of hairs white. Halter yellow

basally; knob light brown and yellow.

Femora with cuneate white scales, a few black scales

apically on posterior pair. Fore and middle tibiae with

light scales; hind tibia with white scales posteriorly

and mixed black and white scales anteriorly. Middle

femur without macrochaetae anteroventrally, hind

femur with three macrochaetae anteroventrally toward

apex; fore tibia with one bristle anterodorsally toward

apex; hind tibia with incomplete row of macrochaetae

anterodorsally.

First abdominal tergum with white pile laterally;

lateral margins of posterior terga with scattered white

hairs, black and white setae, and appressed white scales.

Posterior margin of first tergum with lanceolate white

scales; discs of posterior terga with linear black scales;

yellow scales anteriorly on two and on seven; mixed

yellow and white, lanceolate scales along posterior marSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

gins. Venter with lanceolate white scales and white

setae.

DISTRIBUTION.—Only one specimen of inaquosum is

known. It was collected by the author along the Rio

Mossoro near the railroad southwest of Mossoro, Rio

Grande do Norte, Brazil. The area was sandy and apparently derived subterranean moisture from the river.

The species probably occurs throughout the caatinga

of northeastern Brazil.

HOLOTYPE.— $ , Mossoro, Rio Grande do Norte,

Brazil, XII-5-1960 (N. Marston—2) (USNM).

DISCUSSION.—Anthrax inaquosum apparently has

no close relatives in the genus Anthrax in North and

South America. It superficially resembles Anthrax

pauper (Loew) from North America but may be distinguished readily by setae on the upper half of the face.

Anthrax trimaculatus Group

The trimaculatus group is a heterogeneous assemblage

of species confined to South America, except for one

specimen of trimaculatus Macquart collected at Barro

Colorado Island, Canal Zone, Panama. Most species

may be readily separated from those of the cephus

group by absence of setae on the upper half of the face

just below the antennae. The one species that has such

setae, plurinotus (Bigot), has distinct spots at the bases

of cells R4 and Mi (Plate 6a), while those spots are

absent in species of the cephus group. The Neotropical

species of the oedipus group differ by having short

spurs at the medial angles of vein R4 and the m crossvein of the wing, and by having numerous spots along

the veins other than at the bases of cells. The species

of the tigrinus group may be readily distinguished by

the distinctive crossvein between the m crossvein and

vein Cui on the wing.

No good characters separate the trimaculatus and

the albofasciatus groups, but each species of the former

has some distinctive characters not found in the latter.

Rather than risk confusion by a long statement of

variable characters, the group key simply separates the

two groups on a geographical basis. They appear to

have been derived from a common early Pleistocenic

ancestor and to have evolved differences in a variety

of characters.

No comprehensive studies of the trimaculatus group

have been reported previously, although Edwards

(1930) discussed the Chilean species and Stuardo
NUMBER 4 3 95

Ortiz (1946) listed them in his catalog of the Chilean

Diptera. Nine species are separated in this group; three,

baliopteros, latibasis, and caatingensis, are described

as new.

Although collections of most major museums in

North and South America have been borrowed, only

96 specimens of the trimaculatus group have been obtained, of which 79 are trimaculatus. Thus, all notes

regarding distributions and geographic variation are

tentative. With further collecting, new species probably

will be discovered in areas such as Bolivia, Argentina,

and Chile.

Group Description

Body generally black, femora, tibiae, proximal tarsal

segments and margins of abdominal sterna and posterior terga often yellow or red. Pruinosity mostly gray

or brown, sometimes blue green; eye margins and

front with silver pruinosity. Head globoid; eyes

separated by 1.0 to 3.0 times width of ocellar tubercle

at vertex; antennal sockets separated mesally by about

0.5 their width and about 0.5 to 1.0 of their width from

eye margins. Face and lower part of front produced

slightly above eye margins; oral margin obtusely

angular. Front covered with fine setae and lanceolate

scales; lower half and lateral margins of face with

setae, setae sometimes present on upper half extending almost to antennae. Occiput with short, semirecumbent setae and sparse, small, lanceolate scales;

fringe of pile on posterior margin white to black, usually with lighter tips, some white hairs often present

behind vertex. First antennal segment cylindrical or

enlarged mesad apically, varying in length from 0.5

to 1.5 times apical width. Second segment globular or

greatly flattened apically. Third segment rounded basally, more or less flattened laterally, tapering abruptly

or gradually to styliform part, which varies from 0.5

to 1.5 the length of the base; style 0.2 to 1.5 times the

length of the styliform part.

Disc of mesonotum with fine black setae and linear,

curly scales; scales mixed yellow and white, or forming

complex pattern of spots and stripes of white, gold

and black scales; lateral and posterior margins with

curly white scales. Scutellum with sparse, black setae

and linear or lanceolate scales that are mixed white

and yellow, or more commonly, black in submedial

spots, white laterally or anteriorly and at apex, and

369-673 O—70 7

yellow or gold elsewhere. Macrochaetae black or,

rarely, more or less red. Sternopleuron, mesopleuron,

and anterior part of pteropleuron with white setae and

scales, some black setae and yellow or white pile often

present dorsally on mesopleuron and pteropleuron.

Prosternum, propleuron, and anterior margin of

mesonotum with white or yellow pile, some black setae

on latter. Postalar tuft of pile white or yellow. Hypopleuron and posterior part of pteropleuron bare. Metapleuron bare or with white scales in patches dorsally

and ventrally or directly behind spiracle. Coxae with

black, gold, white or mixed, setae and white scales.

Wing (Plate 6a-i) highly variable, largely hyaline

or with discrete or coalesced spots at bases of some or

all of major cells; rarely with spots at apices of veins

M2, Cui, and Cu2; postmedial spot often present in

cell Ri. Sectoral crossvein present or absent; when

absent, spur at base of cell R4 0.3 to 2.0 times width

of base of cell; spur at base of cell R2+3 absent or as

much as 2.0 times longer than width of base of cell. Cell

2A well developed with rounded margin, or greatly

narrowed with straight margin. Calypter unpigmented,

fringe of hairs white. Stem of halter light brown, knob

light or dark brown basally, yellow or white apically.

Scales on femora mostly white, some black, or rarely

yellow, scales present anterodorsally toward apices,

especially on fore and middle pairs. Scales on tibiae

black anteriorly and white posteriorly, or sometimes

entirely white. Fore femur with incomplete row of

macrochaetae anterodorsally; middle femur with

anteroventral row (sometimes incomplete); hind

femur with anteroventral and sometimes more or less

complete posteroventral rows of macrochaetae; scattered bristles sometimes present dorsally toward apex.

Tibiae with rows of macrochaetae at all four angles,

except anteroventral angle of fore pair.

Discs of abdominal terga usually with scattered

black setae and abundant linear or lanceolate scales,

rarely with abundant, long black setae and curly, hairlike scales. Black scales present in most species in mesally interrupted bands medially on two and anteriorly

on posterior terga, sometimes absent on five, six, and

seven; white scales along posterior margins and sometimes on meson; remainder of scales gold or yellow;

black scales in continuous bands in squalidus. Pile

on lateral margins of first tergum white or yellow;

scales on posterior margin white, sometimes dense and

curly. Lateral margins of terga two, three, and rarely
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

four with black setae and with black, hairlike to

obovate, short or long scales anteriorly and white or

yellow scales posteriorly, black scales rarely absent;

posterior terga with black setae and white or yellow

scales. Venter with sparse, yellow, white, or black setae

and linear to obovate white scales.

Gonocoxite of male genitalia (figures 128-137)

broad basally, tapering or curving to sharp to broadly

rounded apex; apex rounded or flattened, usually with

coarse or fine, dense or sparse setae; medial sulcus

broad or narrow, with sharp or gradually curved margins. Basal segment of gonostylus simple, platelike, with

or without apical setae. Distal segment of gonostylus

highly variable, often triangular basally and tapering

upward and outward to sharp apex, with various teeth,

spines or ridges, sometimes dactylate, ovate and flattened, or with dactylate lateral lobe. Dorsal part of

apex of epiphallus highly variable, often with dorsocentral flat plate or spine, deeply emarginate to sharply

rounded apically in dorsal view; ventrolateral part

broad or narrow, acute or truncate distally; dorsomesal

Dorsal band usually with few setae in minimaculatus,

with preapical, apically projecting lobe in squalidus.

Base of aedeagus bulbous and narrowing abruptly or

narrow and gradually tapering to junction with ventral

bands before or after junction of dorsal bands. Lateral

apodemes of aedeagus open dorsoapically or apically;

basal apodeme large, semicircular or circular.

Ninth tergum of female genitalia (figures 122-126)

simple, apices of ventral arms acute or slightly expanded with short, curved apical lobe. Tenth tergum with

six to ten spines on each side. Sclerite on each side of

gonopore hatchet shaped or club shaped; lateral arm

broad or narrow, acute or truncate distally; dorsomesal

angle sharp, obtuse, rounded or produced as broadly

rounded lobe; ventral arm narrow, tapering or parallel

sided, and curving slightly mesad. First section of each

spermathecal tube short, smooth; second section slightly shorter or slightly longer than bulb, granularly pubescent; third section short, tapering to bulb; bulb

oblanceolate or obovate, of equal size.

Taxonomic Characters

The species of the trimaculatus group have undergone a great amount of differentiation so that there are

discrete differences between most taxa. The wings of

the trimaculatus group (Plate 6a-i) are highly variable

between species, ranging from nearly hyaline in squalidus to heavily marked with pigment on the anterobasal

portion and at the bases of the posteroapical cells in

baliopteros and plurinotus. All species except plurinotus, squalidus, and mystaceus have a distinctive quadrate spot in cell Ri below the tip of vein Sc. Significant

variation in the wing pattern has been noted only in

trimaculatus and bellulus, but in both it is simply an

expansion or contraction of the area of pigment, the

extent of which may depend on environmental factors

as much as on inherited differences. Specimens of the

same species of Anthrax generally have been noted to

be darker from wet areas than those from dry areas.

Two species in the trimaculatus group, plurinotus

and bellulus, have the posterobasal area of the wing

greatly reduced, similar to the species related to argyropygus in the cephus group. The base of cell 2A is almost punctiform and the cell is distinctly narrower than

cell 1A; the alula is reduced to a narrow strip of membrane with a straight posterior margin. Again, no species of the albofasciatus group show that character. A

complete sectoral crossvein is present in bellulus and

latibasis. It often is a somewhat plastic character in

other groups, so specimens may lack the crossvein in one

or both wings. The sectoral crossvein that Macquart

(1840) noted on the wing of mystaceus and that he

used as a diagnostic character for the genus Spongostylum is clearly spurious, judging from the figure of

the wing. One specimen of mystaceus examined showed

the crossvein, and one specimen had a short spur extending anteriorly toward vein R2+3 (Plate 6t) in the

same position as the crossvein in Macquart's figure.

The vesture of the body is similar in all species of the

group except squalidus, which has distinctive long black

setae on the abdominal dorsum and black and white

pile on the sides of the abdomen. Patterns of scales and

hairs on the remaining species are essentially similar,

although the extent of the colors varies somewhat.

However, mystaceus has a distinctive patch of white

scales on the metapleuron directly behind the spiracle,

and latibasis, bellulus, and minimaculatus have dorsal

and ventral patches of white scales on the metapleuron.

The alternating tufts of erect, white or yellow and black

scales on the sides of abdominal terga two, three, and

four are peculiar to trimaculatus, latibasis, minimaculatus, and bellulus.

The male genitalia vary widely interspecifically in

the trimaculatus group and are excellent taxonomic

characters, as illustrated in figures 128-137. Forms of
NUMBER 4 3 97

the apex of the gonocoxite, distal segment of the gonostyluSj and the apex of the epiphallus, in particular, are

distinctive in each species. Significant variation is

found in the structure of the genitalia of trimaculatus,

the only species represented by enough specimens to

allow a study. The limits, however, are not so wide as

to lead to confusion with other species. The female

genitalia also vary between species, although less strikingly than in males. Here, the shape of the sclerite on

each side of the gonopore is most distinctive. The

various structures of the male and female genitalia are

named and discussed under the cephus group. The form

of the male and female genitalia is not emphasized in

keys and diagnoses because there are excellent external

characters for separating the species. The externally

visible parts of the male genitalia, however, provide

an excellent supplementary character to check determinations.

Key to the Neotropical Species of the Anthrax trimaculatus Group

1. Wing with a distinct spot postmedially in cell Ri (Plate 6b-g) 2

Wing hyaline postmedially in cell Ri (Plate 6a,h-i) 8

2(1). Bases of cells Mi and 2Ms of wing pigmented (Plate Sb,e-g) 3

Bases of cells Mi and 2Ma of wing hyaline (Plate 6c—d) 6

3(2). Vein Cu» of wing without spot apically; spots at bases of cells Ri and Mi not expanded

along veins (Plate 6e-g). Metapleuron with at least some scales above and behind

spiracle. Western South America 4

Vein Cuj and often vein Cui of wing with spots apically; pigment at bases of cells R« and

Mi expanded along veins (Plate 6b). Metapleuron bare. Southern Brazil and Uruguay.

baliopteros, new species

4(3). Metapleuron with patch of white scales ventrally behind spiracle. Wing usually with

sectoral crossvein (Plate 6f-g) 5

Metapleuron bare ventrally. Basal angle of vein R« of wing with long spur, but not joined

to vein R2+s to form sectoral crossvein (Plate 6e). Western Argentina

minimaculatus Oldroyd

5(4). Cell 2A of wing greatly narrowed on basal half, distinctly narrower than cell 1A; alula

greatly narrowed, posterior margin straight (Plate 6g). Chile bellulus Philippi

Cell 2A of wing not narrowed on basal half, wider than cell 1A; alula broad, posterior

margin convex (Plate 6/). Peru latibasis, new species

6(2). Basal half of cell 2A of wing broader than cell 1A; alula well developed with convex

posterior margin (Plate 6c-d). Eastern South America to Panama 7

Cell 2A of wing greatly narrowed on basal half, narrower than cell 1A; alula greatly

reduced, posterior margin straight (Plate 6g). Chile bellulus Philippi

7(6). Abdominal terga two, three, and four with tufts of erect black scales laterally; discs with

medially interrupted bands of black scales. Southern Brazil and northern Argentina

to Panama (Map 25) trimaculatus Macquart

Abdomen without black scales, with white setae and few white scales laterally. Northeast

Brazil caatingensis, new species

8(1). Bases of cells in outer part of wing pigmented (Plate 6a,i). Abdomen without long, black

and white pile on lateral margins 9

Wing nearly hyaline (Plate 6/t). Abdomen with long, black and white pile on sides. Chile.

squalidus Philippi

9(8). Metapleuron bare. Setae on face extending nearly to antennae. Chile and western

Argentina plurinotus (Bigot)

Metapleuron with dense patch of scales behind spiracle. Setae on face restricted to lower

half. Chile mystaceus (Macquart)
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Anthrax trimaculatus Macquart

Anthrax trimaculatus Macquart, 1848, p. 194 [trimaculata].—

Kert6sz 1909, p. 56.—Marston, 1964, p. 104.

Argyramoeba imitans Schiner, 1868, p. 122.—Van der Wulp,

1882, p. 86.—KertSsz, 1909, p. 64.

Anthrax imitans.—Painter, 1933, p. 5 [initans],

MALE.—Body generally black, tibiae, margins of

apical abdominal segments, and sometimes femora and

margins of proximal abdominal segments red orange.

First antennal segment inverted conical, about 1.5

times apical width; second segment buttonlike, about

one-half wider than long; bulbous basal part of third

segment slightly narrower than second segment, attenuating to styliform part, which is about as long as base

and twice as long as style. Front with black setae and

scattered, linear, yellow scales; face with black and

white setae along oral and lateral margins, bare on

upper half below antennae. Occiput with black setae

and linear gold scales on upper third, with white setae

and scales below; fringe of pile on posterior margin

black or yellow on upper third, white below and behind

vertex.

Disc of mesonotum with fine black setae and linear

scales, white in posteriorly bifurcate medial stripe and

transversely in front of bases of wings, light yellow to

gold elsewhere, submedial spots of black scales present

posteriorly on specimens with gold scales; lateral and

posterior margins with long, linear, semirecumbent

white or white and yellow scales and black setae and

macrochaetae. Scutellum with black setae and linear

scales varying from yellow on disc and white on margins to black on disc and gold on margins. Sternopleuron, mesopleuron, and anterior part of pteropleuron with fine white setae and linear white scales,

some black and gold macrochaetae present dorsally on

last two. Propleuron, prosternum, and anterior margin

of mesonotum with white or yellowish-white pile, some

black setae on latter. Postalar tuft of pile white or

yellowish white. Metapleuron bare or with few linear

white scales just above coxa. Coxae with black, gold

and white bristles and linear and lanceolate white

scales.

Wing (Plate 6c) hyaline with light or dark brown

spots at bases of cells R2+3, Rs, 1M2, Cui, postmedially

in cell Ri, and below base of cell Rx in cell R; spots

varying from small and distinct to large and confluent;

base of cell 2M2 with very small spot in darker specimens. Bases of cells R2+3 and R4 with basal spurs about

as long as basal sections of veins. R-m crossvein at basal

two-fifths of cell 1M2, vein R2+3 arising opposite; contact of cells 1M2 and Cui about 1.5 times longer than

width of base of cell Cux. Cell 2A 1.0 to 1.5 times as

wide as cell 1A; alula well developed with posterior

margin distinctly convex. Calypter unpigmented, fringe

of hair white. Stem of halter light to dark brown, knob

dark brown basally, yellow apically.

Anterior sides of femora with black and dark brown

scales apically and yellow to gold scales basally, posterior sides with white scales. Fore and middle tibiae

with black scales anteroventrally and white scales elsewhere; hind tibiae with mixed black, white, and yellow scales. Middle femur with anteroventral row of

macrochaetae; hind femur with anteroventral and

posteroventral rows.

First abdominal tergum with white or yellowishwhite pile laterally, some black hairs on darker specimens; posterior margin with long, curly, lanceolate

white scales. Lateral margins of terga with long, erect,

black pile, lanceolate scales and setae medially on two,

and anteriorly on three, four, and sometimes five; remainder of lateral margins with black and white setae,

and lanceolate and ovate, white, yellow, and sometimes

gold scales. Discs of terga two through seven with fine

black setae and linear scales, black in medially interrupted transverse bands or spots of varying extent,

white along meson, and medially and laterally along

posterior margins of all but three and four, yellowish

white to reddish brown over remainder. Venter with

white or yellowish-white setae and sparse white scales.

MALE GENITALIA (Figure 132).—Gonocoxites broad

basally, bilobed apically; apical lobes membranous

laterally, separated mesally by deep sulcus; dense, fine

hair on lobes and mesally before bases of lobes. Basal

segment of gonostylus simple, slightly convex dorsally,

with fine hairs apically. Distal segment of gonostylus

projecting apicolaterally, with flat basal section from

which the apical section curves dorsally and laterally;

apex of dorsal section acute or flat and rounded, dorsal

margin flat, with ridges, or with preapical tooth; preapical ventral tooth on one specimen. Dorsal part of

apex of epiphallus flat, acute basally in lateral view,

sharply angled basally in dorsal view; apex broadly

rounded in dorsal view, right angled dorsally and

broadly rounded ventrally in lateral view. Ventrolateral

part of apex of epiphallus extending downward and

backward from dorsal part, with sharp ridge curving

upward and apically from point below proximal end of

dorsal part. Dorsal band unmodified. Base of aedeagus

tubular, tapering gradually to junction with ventral

bands. Epandrium simple with irregular margins.
NUMBER 4 3 99

Cerci narrow, with few setae, with sharp ventral lobe.

FEMALE.—Similar to male.

FEMALE GENITALIA (Figure 125).—Tenth tergum

with ten spines on each side. Ventral arm of ninth tergum broad, S-shaped distally, with small, dactylate

ventral projection. Dorsal part of sclerite on each side

of gonopore produced dorsolaterally as a broad, flat

plate with blunt dorsomedial angle, sharp dorsal angle,

and broadly rounded ventrolateral angle; ventral arm

narrow and curved mesad at tip. Each duct of spermathecae slightly longer than bulb; first section short,

clear; second section about as long as bulb, covered

with granular pubescence; third section very short;

bulb elongate oval, tapering basally to junction with

duct.

DISTRIBUTION AND ECOLOGY.—Anthrax trimaculatus

has been collected east of the Andes Mountains from

Uruguay and northern Argentina to Panama (Map

25). It occurs in a wide variety of habitats, from the

caatinga of northeastern Brazil to the grasslands of

southern Brazil and the tropical rain forest of Panama.

Specimens were collected by the author throughout

much of eastern Brazil, usually along paths or near soil

or clay banks where bees were nesting.

BIOLOGY.—One reared specimen of this species has

been studied. It was reared from the nest of a species

of Diadasia (Hymenoptera: Apoidae) by Pe. Jesus S.

Moure. The pupa was described previously (Marston,

1964).

TYPES.—The type male of Anthrax trimaculatus

Macquart is in the Bigot collection in the British

Museum. According to notes by R. H. and E. M.

Painter, the right wing and third antennal segments

are missing, the head is glued on, and some fungus is

present. It carries the label "Anthrax trimaculata nov.

sp. cf Brasil Macq. D. Ex. nom. +. " A photograph

and redescription of the type made by the Painters

agrees well with specimens from northeastern Brazil.

The type-locality given by Macquart is "Brasil."

The type male of Argyramoeba imitans Schiner is in

the Vienna Museum. R. H. and E. M. Painter found it

"in good condition except that it is greasy." It is labeled

"Lindig 1864 Venezuela" and "imitans Schin. Alte

Sammlung." A specimen from Corazal, Canal Zone,

Panama, agrees with the redescription made by the

Painters. They also labeled a specimen from Paraiba,

Brazil, as a homotype.

DISCUSSION.—Edwards (1930) recorded imitans

from Chile, but he was undoubtedly referring to a lightcolored specimen of bellulus Philippi.

Anthrax trimaculatus is a highly variable species.

Specimens from forested areas in southern Brazil have

the spots on the wings largely or entirely coalesced;

whereas, at the other extreme, specimens from the

desert area of northeastern Brazil may have small and

distinct spots. The male genitalia also vary considerably especially the distal segment of the gonostylus,

which may have the dorsoapical projection sharply

tapered or flattened and broadly rounded distally, and

with or without dorsal ridges or a dorsal spine. A complete range of variation has been noted in a long series

of specimens from Itu, Sao Paulo, Brazil. As variation

is wide within habitats and intergradations occur between habitats, all variations noted are regarded here

as the same species.

One specimen from Tucuman, Argentina, has a

short preapical tooth ventrally on the dorsoapical part

of the distal segment of the gonostylus, and has the

dorsal part of the apex of the epiphallus distinctly

shorter than in other males of trimaculatus examined.

However, because the specimen's external characters

fall within the range of variation of specimens from

eastern Brazil, and since there is considerable variation in the structure of the male genitalia in that area,

the specimen from Tucuman is regarded simply as a

variant.

Anthrax trimaculatus differs from latibasis and most

specimens of bellulus in lacking spots at the bases of

cell R4 and Mi and in lacking a sectoral crossvein.

Specimens of bellulus with light-colored wings may be

distinguished readily by the reduced anal margin of

the wing. Anthrax trimaculatus differs from other

species with a postmedial spot in cell Ri by having long,

erect black scales and pile on the sides of abdominal

terga two, three, and four.

Anthrax latibasis, new species

MALE.—Body mostly black, brownish and grayish

pruinose; legs except metatarsi and dorsoapical parts

of femora, margins of thoracic pleura, lateral and

posterior margins of abdominal terga, posterior margins of abdominal sterna, and genitalia orange. Front

with black setae and lanceolate white scales. Face with

black setae laterally and along epistomal margin, upper

half bare below antennae; some yellowish-white hairs

below. Occiput with fine, short, black setae and lanceolate scales, yellow along eye margins on upper third,

white elsewhere; fringe of pile on posterior margin

white below and behind vertex, black above. First an-
100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MAP 25.—Distribution of Anthrax trimaculatus.
NUMBER 4 3 101

tennal segment inverted conical, about as long as apical

width; second segment buttonlike, narrower than apex

of first segment and slightly wider than base of third;

third segment bulbous basally, tapering abruptly to

styliform part, which is about as long as base and about

twice as long as style; setae on basal segments black.

Disc of mesonotum with fine black setae and linear

black, gold, and white scales forming complex pattern

of longitudinal bands and spots. Lateral and posterior

margins with coarse black setae and semierect, curly,

linear and lanceolate white scales. Anterior margins

and apical spot on scutellum with linear white scales;

disc with patches of linear black scales on each side;

remainder of disc and posterior margin with gold

scales. Mesopleuron, sternopleuron, and anterior part

of pteropleuron with fine, yellowish-white setae and

erect, oblanceolate, curly white scales; some coarse gold

setae and bristles above on mesopleuron and pteropleuron. Prosternum, propleuron, and anterior margin

of mesonotum with white pile, some black setae present on latter. Postalar tuft of pile white. Metapleuron

behind spiracle with small tuft of lanceolate white

scales above and larger patch of linear white scales

below. Coxae with black setae and linear and lanceolate

white scales.

Wing (Pate 6/) largely hyaline, cells C and Sc and

extreme base subhyaline; brown spots at bases of cells

R4, Mi, 2M2, R2+3, and R5, in cell Ri above base of sectoral crossvein, running from base of cell Rt

 to base

of cell Cui, and from base of cell R to base of cell 2A.

Base of cell Rt connected to vein R2+3 by an evenly

curved crossvein without spurs. R-m crossvein located

halfway from base to apex of cell 1M2; vein R2+3 arising

opposite, with basal spur slightly longer than r-m crossvein. Contact of cells 1M2 and Cui about 1.5 times

longer than base of cell Cux. Cell 2A not reduced,

slightly wider than cell 1A; alula broad, posterior margin distinctly convex. Calypter subhyaline, fringe of

hairs black. Stem of halter light brown, with lanceolate

white scales at base; knob dark brown basally, light

yellow at apex.

Femora and tibiae with ovate-truncate, white scales

except for some yellow and black scales anterodorsally

at apices of femora. Middle femur with complete row

of macrochaetae anteroventrally; posterior femur with

complete anteroventral and posteroventral rows of

macrochaetae and few scattered dorsally at apex.

Lateral margins of first abdominal tergum with white

pile; posterior margin with long, curly, lanceolate white

scales. Lateral margins of terga two, three, and four

with long, erect, lanceolate-truncate, dark brown and

white scales anteriorly, and white scales along posterior

margins; lateral margins of terga five, six, and seven

with shorter, ovate-truncate white scales posteriorly.

Discs of terga two and three with linear scales, white

anteriorly on first and in medial posterior spots on

both, black in transverse, medially interrupted bands

on posterior margins, mixed yellow and gold elsewhere.

Fourth tergum with submedial patches of linear black

scales separated by mixed yellow and white scales, and

mixed yellow and white, lanceolate-truncate scales

laterally. Fifth, sixth, and seventh terga with lanceolatetruncate, yellow scales anteriorly and lanceolate-truncate white scales posteriorly, small patch of linear black

scales present medially on five. Entire dorsum with

scattered black setae. Venter with fine yellow setae and

scattered, short, lanceolate- and ovate-truncate white

scales.

MALE GENITALIA (Figure 128).—Gonocoxites broad

basally, tapering in both lateral and ventral view to

sharp apices; medial sulcus narrow and deep, widened

toward base; lateral margin unsclerotized medially;

fine setae scattered from base to apex. Basal segment

of gonostylus platelike and rounded dorsally in lateral

view, not extending past base of distal segment, with

few fine setae apically. Distal segment of gonostylus

with rectangular, proximally emarginate basal part

curving mesad dorsally, from which a dactylate, apically broadened and truncate lobe extends distally. Dorsal part of apex of epiphallus with short, narrow, dorsomesal part and apically rounded, ventrobasally acute,

larger lobes extending downward and forward. Ventrolateral part of apex of epiphallus very narrow apically,

but recurving ventrally to form broad, rectangular

preapical plate. Dorsal bands simple, narrow after

junction above aedeagus. Basal part of aedeagus narrow, tapering gradually to tip with slight enlargement

below junction of dorsal bands. Epandrium simple,

with coarse bristles. Cerci oblong, tapering from dorsal

margin near base to rounded apex; dorsal two-thirds

velvet black, bottom third orange; with dense, minute

setae giving velvety appearance.

FEMALE.—Similar to male. Wing pigment slightly

more extensive, cell M largely pigmented. Fore and

middle tibiae with black scales anteriorly. Dorsum of

abdomen with black and white scales more extensive

and gold scales less extensive.

FEMALE GENITALIA (Figure 124).—Tenth tergum

with six spines on each side. Ventral arm of ninth tergum parallel sided and tapering slightly to acute apex.
102

Dorsomedial angle of sclerite on each side of gonopore

extending mesally as broad, apically rounded lobe

about equal in length to lateral arm, which is broad

and shallowly emarginate apically; ventral arm narrow, parallel sided and slightly curved mesad. Each

duct of spermathecae about twice as long as bulb; first

section short, unpigmented; second section about 1.5

times longer than bulb, pigmented and covered with

granular pubescence; third section short; bulb ovoid,

with acutely rounded tip.

VARIATION.—The remaining specimens of this species seen by the author do not vary beyond limits of the

holotype and allotype.

DISTRIBUTION.—The few specimens of latibasis

available indicate that it occurs in the desert area along

the coast of Peru and perhaps into northern Chile. The

female specimens from Matucana were collected along

a railroad track in an area with only a few cacti and

tufts of grass among rocks.

HOLOTYPE.— $ , Matucana, Lima, Peru, V-21-1920

(Cornell Univ. Exped. Lot 569) (AMNH).

ALLOTYPE.— $, Matucana, Lima, Peru, X-31 -

1964, 8000 feet (R. H. Painter) (RHP).

PARATYPES.—Peru: Arequipa, cf, $ , Arequipa,

1922 (Dt. E. P. Reed) BM 1931-360 (BM). Lima,

9 , Chosica, 111-23-1964 (Peter Hocking) (caught in

short grass by path) (RHP); 2 $ , Matucana, X-31 -

1964, 8000 feet (R. H. Painter) (RHP, NLM).

"Canta," VI-26-1955, 2800 m (Weyrauch) (WKW).

DISCUSSION.—Both latibasis and bellulus have the

spur at the base of cell R, of the wing connected to

vein R2+3 to form a stable sectoral crossvein. In this

case, the crossvein is evenly curved and has no short

spurs extending from it. In some other species some

specimens may have a spurious crossvein, in which case

it is angled and bears short spurs; there may also be

differences between the two wings and other evidence

of abnormal venation such as short spurs and crossveins. Judging from the illustration given by Macquart,

the type of Spongostylum mystaceum has such an abnormal crossvein and is probably a species that normally has only two submarginal cells.

Anthrax latibasis is most closely related to bellulus

Philippi. The two species are quite similar in external

appearance, but may be readily distinguished by the

reduced anal margin of the wing of the latter. The

male genitalia also are quite different as shown in the

illustrations. Anthrax latibasis and bellulus differ from

other species with a postmedial spot in cell Rx by the

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

presence of a stable sectoral crossvein and patches of

white scales on the metapleuron behind the spiracle.

Anthrax bellulus Philippi

Anthrax bellulus Philippi, 1865, p. 675 [bellula].—Kert6sz,

1909, p. 30.—Oldroyd, 1938, p. 84.—Stuardo Ortiz, 1946,

p. 96 [bellula].

Spongostylum inappendiculatum Bigot, 1892, p. 351 [Spogostylum].—Kert6sz, 1909, p. 58.

Anthrax inappendiculatus.—Edwards, 1930, p. 172.—Stuardo

Ortiz, 1946, p. 93.

Anthrax imitans.—Edwards, 1930, p. 172 [not Schincr, 1868,

misidentification].

MALE.—Body mostly black, legs orange or with

dorsoapical parts of femora dark red; apices of sterna

broadly orange, of terga narrowly orange. Front with

black setae and scattered, lanceolate white scales.

Lower half and lateral margins of face with mixed black

and yellow setae; upper half of face bare below antennae. Occiput with fine, scattered setae, black on upper

two-thirds, white below; scales linear, gold along eye

margins on upper third, white elsewhere; fringe of pile

on posterior margin black behind gold scales, white elsewhere. First antennal segment about as long as apical

width; base of third segment slightly narrower than

second segment, bulbous, constricting abruptly to a

styliform part which is about as long as base and about

twice as long as style.

Disc of mesonotum with fine black setae; linear scales

white in medial stripe which divides a round patch of

black scales on posterior half, black and orange laterally

on anterior half; lateral and posterior margins with

woolly, semierect, linear white scales, and black setae

and macrochaetae. Scutellum with fine black setae

and linear scales, gold along margins, black on disc,

few white scales anterolaterally and posteromedially.

Sternopleuron, mesopleuron, and anterior part of

pteropleuron with white pile and linear scales, gold

and black macrochaetae dorsally on last two. Prosternum, propleuron, and anterior margin of mesonotum

with yellow and white pile, some black setae on latter;

postalar tuft of pile yellow. Metapleuron with patches

of linear scales dorsally and ventrally behind spiracle.

Coxae with black and yellow setae and linear and

lanceolate white scales.

Wing (Plate 6g) hyaline with light brown spots at

bases of cells R2+3 and RB, R4, Ml5 2M2, in cell Ri

above base of sectoral crossvein, running from base of

cell Rx to base of cell Cui, and at bases of cells R,

M, and 1A; spots occasionally reduced or absent at

bases of cells R,, Ml5 and 2M2; base of wing and cells
NUMBER 4 3 103

C and Sc light brown. Sectoral crossvein usually

present, without angles and spurs. Basal angle of vein

R2+3 with long spur. Contact of cells 1M2 and Cua

about three times longer than width of base of cell

Cux. R-m crossvein about halfway to apex of cell 1M2;

vein R2+3 arising basad about length of r-m crossvein.

Cell 2A greatly narrowed basally; alula reduced,

posterior margin straight. Calypter subhyaline, fringe

of hairs brown above, white below. Stem of halter

brown, darker above, with few white scales at base;

knob dark brown, tip yellow.

Scales on femora mostly white, some black ones

anterodorsally on apical halves, especially on fore and

middle pairs. Scales on tibiae white. Middle femur with

row of macrochaetae anteroventrally; hind femur with

anteroventral row of macrochaetae and some scattered

anterodorsally toward apex.

First abdominal tergum widi white pile laterally and

linear, curly white scales along posterior margin.

Lateral margins of terga two through four with scattered black setae and long, erect, lanceolate-truncate

scales, mostly black anteriorly on segments, yellowishwhite posteriorly; lateral margins of terga five, six,

and seven with ovate-truncate, yellowish-white and

white scales along posterior margins. Discs of terga two

through seven with scattered black setae and linear

scales, black in submedial patches posteriorly on two

and anteriorly on three and four, white anteriorly on

two, longitudinally along meson, and posterolaterally,

especially on posterior terga, scales mostly gold elsewhere. Venter with light-yellow setae and lanceolate

white scales.

MALE GENITALIA (Figure 133).—Gonocoxites broad

in ventral view, narrow apically in lateral view; apices

gradually rounded to rounded mesal angles in ventral

view; mesal sulcus narrow; lateral margins unsclerotized apically; setae scattered over apical fourfifths. Basal segment of gonostylus simple, slightly convex dorsally in lateral view, with few fine setae apically.

Distal segment of gonostylus projecting distally, a flat

plate curving upward and outward with acute distal

and lateral angles apically. Dorsal part of apex of

epiphallus with raised transversely rounded mesal

ridge, basal angle rounded in dorsal view, acute in

lateral view; apex sharply rounded in lateral view,

extending beyond tip of aedeagus. Ventrolateral part

of apex of epiphallus extending downward and backward from dorsal part, unmodified. Dorsal bands

simple, narrow after junction. Base of aedeagus tubular, narrow, tapering gradually to apex. Epandrium

simple; each cercus ovoid with blunt ventroapical angle

in lateral view, with fine, dense setae giving velvety

appearance.

FEMALE.—Similar to male.

FEMALE GENITALIA (Figure 123).—Tenth tergum

with about ten spines on each side. Ventral arm of ninth

tergum narrow, acute apically, posterior margin curled

toward meson medially. Dorsomedial angle of sclerite

on each side of gonopore large, projecting dorsornedially as a broadly rounded lobe, broader than

lateral arm which is slightly curved upward apically;

ventral arm slightly curved laterad above, with sharp

ventral point. Each duct of spermathecae about twice

as long as bulb; first sections short, unsclerotized; second section about 1.5 times as long as bulb, sclerotized

and covered with granular pubescence; third section

short, clear; bulb ovoid with short stem.

DISTRIBUTION.—Only three specimens of this species

have been seen by the author. One is labeled "Chile.

A. Faz," the second "Perales. 1-26," and the third

"Perales. 1-25" and "Valparaiso, Chile. A. Faz." No

specific type-locality is given, although it probably is

the province of Santiago, Chile, since most of Philippi's

types came from there. The specimen referred to below

as inappendiculatus was collected at Majada Blanca,

Manquehua, Coquimbo, Chile. This species probably

occurs throughout central Chile and possibly in

western Argentina.

TYPES.—The type of Anthrax bellula Philippi,

which was presumably in the Santiago Muesum, has

not been seen by the author. The specimens at hand

agree perfectly with the description, however, and, as

Philippi stated, bellulus cannot be confused with any

other Chilean species. Philippi makes no mention of

the presence of a sectoral crossvein on the wing, but

from his description it may be inferred. Of the specimens examined, two have the crossvein present and

the other has it present on one wing but not on the

other.

According to notes by R. H. and E. M. Painter, the

type female of Spongostylum inappendiculatum Bigot

is in the Bigot collection of the British Museum. It is

headless, as Bigot stated, but is otherwise well preserved. It carries the label "Spogostylum inappendiculatum ? Inedit. n. sp. Quinze Aout 1888. J. Bigot,

Chili."

A specimen in the collection of the author from

Coquimbo, Chile, agrees well with the redescription

and photograph of the type furnished by Dr. and Mrs.

Painter. The two specimens apparently differ from
104

typical specimens of bellulus only in having the pigmentation on the wings reduced with the spots at the

bases of cells R4, Mx, and 2M2 absent or represented

by faint clouds. Further collecting may show that

bellulus and inappendiculatus are separate taxa, but

with the limited number of specimens available, it

seems best to regard them as one species.

DISCUSSION.—Specimens of bellulus without spots

at the bases of cells R4, Mi, and 2M2 may be separated

from similar species in northern and eastern South

America by the reduced anal margin of the wing,

dorsal and ventral patches of white scales on the metapleuron behind the spiracle, and by the presence of a

sectoral crossvein on the wing. Characters separating

typical specimens from latibasis and other related

species are given under latibasis.

Anthrax minimaculatus Oldroyd

Anthrax minimaculatus Oldroyd, 1937, p. 83.

MALE.—Basal antennal segments, legs, margins of

thoracic pleura, and margins of abdominal segments

reddish or orange. Front with black setae and linear

gold scales. Face with mixed yellow and black setae

and yellow scales. Occiput with fine yellow setae and

yellow, gold and white scales; fringe of pile on posterior

margin yellow with light tips. First antennal segment

about 1.5 times longer than wide, cylindrical; second

segment globular, slightly narrower than apex of first

segment, base of third segment narrower than second

segment, elongate globular, about three-fourths as long

as styliform part, which is about 1.5 times longer than

style.

Disc of mesonotum with fine black setae and mixed

gold, yellow, and white, linear scales, few black scales

medially on posterior half; lateral margins with longer,

suberect, linear, gold, yellow and white scales, and

coarse black setae and macrochaetae. Scutellum with

gold, yellow and white, linear scales on margin

and along mesal line, with linear black scales in submedial patches. Pile, setae, and scales on sternopleuron,

mesopleuron, and anterior part of pteropleuron white,

setae on last two yellow. Pile on prosternum, propleuron, and anterior margin of mesonotum yellowish

white, some black hairs on latter; postalar tuft of pile

white. Metapleuron with few lanceolate, white scales

above and behind spiracle, bare below. Coxae with

yellow setae and black scales.

Wing (Plate 6e) largely hyaline, cells C and Sc subhyaline. Light-brown spots present at bases of cells

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

R2+3, R5, R4, Mi, and 2M2, postmedially in cell Ra,

extending from base of cell Rj to base of cell Cuj, and

at bases of cells R, M, and 2A. Basal angle of vein R«

with long spur, but not connected to vein R2+3 to form

sectoral crossvein. R-m crossvein at basal two-fifths of

cell 1M2; contact of cells 1M2 and Cui about 1.5 times

longer than width of base of cell Cu,. Cell 2A slightly

broader than cell 1A, not narrowed basally; alula well

developed, posterior margin convex. Calypter subhyaline, fringe of hair white. Stem of halter yellow;

knob yellow basally, white apically.

Femora and tibiae with ovate white scales except

for few yellow scales anteriorly toward apices of femora. Middle femur with complete anteroventral row

of macrochaetae; hind femur with complete anteroventral and posteroventral rows and few scattered

dorsally at apex.

Lateral margins of first abdominal tergum with

yellowish-white pile, few brown hairs posteriorly; posterior margin with fine, curly, yellow and white scales.

Lateral margin of second tergum with white setae and

scales anteriorly, with black setae and long, lanceolate,

brown and white scales posteriorly; terga three and

four with black setae and lanceolate-truncate scales,

black anteriorly and white posteriorly; lateral margins

of remaining terga with fine white setae and sparse,

short white scales. Discs of terga with fine black setae

and linear and lanceolate scales, white and yellowish

white in medial transverse band on two, in mesal

longitudinal spots and posterolaterally on posterior segments, mixed gold, yellow and white elsewhere, few

black scales submedially at middle of two and submedially along anterior margin of three and four.

Venter with fine white setae and lanceolate scales.

MALE GENITALIA (Figure 129).—Gonocoxites broad

basally, tapering with slight preapical emargination to

broadly rounded apices in ventral view, narrow with

acute apices in lateral view, medial sulcus narrow and

shallow, deepening apically to separate distinct distal

lobes; setae short, becoming sparse basally. Basal segment of gonostylus platelike, slightly rounded dorsally,

extending to base of distal segment. Distal segment

broadest mesolaterally, triangular basally, rounding

dorsomedially to sharp apex; large, sharp, preapical

tooth present laterally. Dorsal part of apex of epiphallus flattened dorsoventrally, with sharp, recurved basal

spine; apex broadly rounded in dorsal view, mesal

part raised and curved to apex in lateral view. Ventrolateral part of apex of epiphallus extending downward

and backward from before apex, narrow, tapering, and
NUMBER 4 3 105

sharp distally. Dorsal bands simple, with row of fine

erect setae before and after junction. Epandrium simple, with coarse setae; cercus oblong dorsoventrally, entirely pale, with fine yellow setae.

FEMALE.—Similar to male, judging from Oldroyd's

description.

DISTRIBUTION.—Anthrax minimaculatus is known

only from the type series from La Rioja, Patquia,

Argentina.

TYPE.—The type female and 12 paratypes of Anthrax minimaculatus Oldroyd are in the British

Museum. A photograph by R. H. and E. M. Painter

shows the type female is in good condition. Although

Oldroyd stated that the 12 paratypes were all females,

the two paratypes loaned for study are males.

DISCUSSION.—Anthrax minimaculatus may be

readily distinguished from both bellulus and latibasis by

absence of a complete sectoral crossvein on the wing,

and by absence of scales on the metapleuron behind

and below the spiracle. In addition, it differs from

latibasis by absence of a distinct, medially interrupted

crossband of black scales on the second abdominal

tergum; bellulus differs by having the posterobasal

part of the wing reduced with the alula narrow and

straight margined. Anthrax minimaculatus differs from

other species in western South America by having a

postmedial spot in cell Ri of the wing.

Anthrax baliopteros, new species

MALE.—Integument mostly black, grayish to brownish pruinose; bases of femora, tibiae, proximal tarsal

segments, apices of abdominal segments, and genitalia

reddish orange. Front with fine black setae and lanceolate, semierect scales, yellowish white above and on

each side of antennae, gold elsewhere. Face with mixed

black and gold setae extending almost to antennae

medially; few lanceolate, erect white scales along oral

margin. Occiput with sparse black setae and linear

and lanceolate, recumbent scales, white below emargination of eye, gold above; fringe of pile on posterior

margin black behind gold scales, white behind vertex,

mixed brown and white behind white scales. Antennae

black, first two segments with black and white setae;

first segment short, about as wide as long; second segment biscuit shaped, about half as long as wide; third

segment with bulbous basal portion small, narrower

than second segment, about as long as styliform portion

which is twice as long as style.

Mesonotum with fine black setae and linear, recumbent gold and white scales; margins with semierect gold

and white scales. Scutellum with fine black setae and

linear scales, black in submedial spots, mixed gold

and white on anterior margin, gold elsewhere. Mesopleuron, sternopleuron, and anterior part of pteropleuron with white setae and scales, mesopleuron with

some gold macrochaetae above. Prosternum, propleuron, and anterior margin of mesonotum with white

pile, few black setae present on latter. Postalar tuft of

pile mixed gold and white. Metapleuron bare. Coxae

with yellowish-white setae and white scales.

Wing (Plate 66) hyaline with brown maculation;

dark spots present in cell Ri below tip of subcosta, at

bases of cells R2+3 and Rs, and extending from base of

cell Cux to base of cell Ri; lighter spots at base of cell

R4 and extending along vein R4 two thirds of way to

apex, at base of cell Mx and along vein M2 and m

crossvein, at base of cell 2M2, and apically in cell 1A;

faint subhyaline areas near apices of veins Mi and Cui.

Cells C and Sc, and base of wing lightly infuscated,

base of cell M darker. Veins R2+3 and R4 with basal

spurs about as long as basal sections. Contact of cells

1M2 and Cux twice as long as base of cell Cux. R—m

crossvein at basal two-fifths of cell 1M2; vein R2+3 arising opposite. Anal margin of wing well developed,

alula broad with rounded posterior margin.

Legs with ovate-truncate scales, yellow anterodorsally at apices of femora, white elsewhere. Middle

femur with anteroventral row of macrochaetae; posterior femur with complete anteroventral and

posteroventral rows of macrochaetae and scattered

macrochaetae anterodorsally toward apex.

Lateral margins of first abdominal tergum with

dense white pile, with few black hairs posteriorly;

posterior margin with few linear white scales on lateral

thirds. Lateral margins of second tergum mostly with

white pile, few semierect, lanceolate, yellowish-white

scales posteriorly; lateral margins of terga three and

four with short, lanceolate, erect black scales anteriorly

and semierect, lanceolate, yellowish-white and yellow

scales posteriorly; lateral margins of terga five, six,

and seven with few black and yellow setae and some

recumbent, lanceolate cuneate, white scales. Disc of

second tergum with mixed gold and white, linear

scales, predominantly gold posteriorly, two small submedial spots of linear black scales near posterior margin; terga three and four with linear gold scales on

lateral and posterior margins, with linear yellow scales

medially and linear black scales elsewhere; terga five
106

and six with small submedial spots of linear black

scales, with mixed yellow and white, lanceolate scales

medially and laterally, and linear gold scales between

black and light scales; seventh tergum with mixed yellow and white, lanceolate scales medially, a few black

scales submedially, linear gold scales sublaterally and

a few lanceolate cuneate, yellow scales laterally. Venter

with fine gold setae and small white scales.

MALE GENITALIA (Figure 131).—Gonocoxites broad

basally, tapering to acute apices in both dorsal and

lateral view; medial sulcus shallow, narrow, setae scattered from near apices to base. Basal segment of gonostylus simple, slightly rounded dorsally in lateral view,

with small setaceous lobes apically in ventral view.

Distal segment of gonostylus projecting dorsolaterally,

dactylate with anterior margin narrowing postmedially

to transversely broadened, sharply rounded apex. Dorsal part of apex of epiphallus with large, preapical,

basally directed sharp spine in back of a deep, broad

mesal emargination. Ventrolateral part of apex of epiphallus extending downward and backward from dorsal part, not notably modified. Dorsal bands simple.

Base of aedeagus narrowly bulbous, tapering gradually

to junction with ventral bands. Epandrium about

one-third narrower than high, with coarse setae apically. Cerci not heavily sclerotized, with fine setae.

FEMALE.—Similar to male. Scales on front white

below, yellowish white above. Postalar tuft of pile

entirely white. Maculation of wings somewhat darker;

fusion of cells 1M2 and Cui only 1.5 times as long as

width of base of cell Cu!. Femora entirely yellow except

for dorsoapical parts of posterior pair; macrochaetae

on posteroventral side of posterior femur restricted

to apex. Gold scales more extensive and yellow and

white scales less extensive on abdomen.

VARIATION.—The only other specimen of this species

available, a male, is similar to the holotype except

that its wings have slightly less extensive pigment.

DISTRIBUTION.—The three specimens available indicate that baliopteros probably occurs throughout the

drier tropical and subtropical areas of southern Brazil

and Uruguay, and possibly into northeast Argentina

and southern Paraguay.

HOLOTYPE.—cf, Bage, Rio Grande do Sul, Brazil,

1 March 1961 (N. Marston-1) USNM).

ALLOTYPE.— ? , same data as holotype (USNM).

PARATYPE.— cf, Araraquara, Sao Paulo, Brazil, I17-18-1941 (M. Carrera) (26146) (SASP).

DISCUSSION.—The holotype and allotype were collected along an overhanging bank of a stream near a

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

patch of woods in the "campos" or grassland area of

southern Rio Grande do Sul.

Anthrax baliopteros may be readily separated from

other species with a postmedial spot in cell Ri by the

peculiar wing pattern in which the pigment runs along

vein R4, M2 and the m crossvein from the bases of

cells R4 and Mi. The small apical spot in cell 2A also

is distinctive.

Anthrax caatingensis, new species

FEMALE.—Face, lower part of front, first antennal

segments, legs except apical tarsal segments, margins

of thoracic pleura, posterior part of scutellum, posterior abdominal segments, and posterior margins of anterior abdominal segments reddish orange; remainder

of body black. Front with black setae above, with

mixed black and white setae and linear white scales

below. Face with yellowish-white setae extending threefourths of way to antennae and few linear white scales.

Occiput with yellowish-white setae and lanceolate

scales; fringe of pile on posterior margin white below,

yellowish white above. First antennal segment about

as long as apical width; second segment buttonlike;

base of third segment bulbous, tapering to styliform

part, which is about as long as base and about twice

as long as style.

Disc of mesonotum with yellowish-white and brown

scales, and fine setae; margins with yellowish, curly

scales, gold setae and few black macrochaetae. Scutellum with lanceolate brown scales medially and yellowish-white scales along posterior margins. Sternopleuron, mesopleuron, and anterior part of pteropleuron with white setae and scales, some gold bristles

dorsally on last two. Prosternum, propleuron, and

anterior margin of mesonotum with white and yellowish-white pile. Postalar tuft of pile white. Metapleuron

with patch of linear white scales above coxa. Coxae

with white bristles and lanceolate scales.

Wing (Plate 6d) with light-brown spots postmedially in cell Rl5 at bases of cells R2+3 and R5, extending from base of cell Cua to base of cell Rl5 and at bases

of cells R and M; anterobasal part of wing between

spots yellowish brown to subhyaline. Basal angle of vein

R2+3 with spur about as long as basal section; basal

angle of vein R4 with short spur about one-fourth as

long as basal section. R-m crossvein at basal two-fifths

of cell 1M2; vein R2+3 arising slightly basad. Contact

of cells 1M2 and Cui about twice as long as width of

base of cell Cui. Cell 2A about 1.5 times as wide as cell
NUMBER 4 3 107

1A; alula well developed, posterior margin convex.

Calypter subhyaline, fringe of hairs white. Stem of

halter brown, knob brown basally, yellow apically.

Femora and tibiae with white and yellowish-white,

lanceolate scales. Middle femur with incomplete anteroventral row of macrochaetae. Hind femur with

anteroventral and anterodorsal rows of macrochaetae

incomplete basally.

Lateral margins of first abdominal tergum with yellowish-white pile; posterior margin with curly, lanceolate, yellowish-white scales. Lateral margins of terga

two through seven with white setae and some semirecumbent, lanceolate white scales, few black setae posteriorly. Discs of terga two through seven with sparse

black setae, and linear and lanceolate scales, yellow

and gold anteromedially, white posterolaterally, especially on posterior terga. Venter with fine white setae

and ovate white scales.

FEMALE GENITALIA (Figure 122).—Tenth tergum

with seven spines on each side. Ventral arm of ninth

tergum narrow, apex not enlarged. Dorsomedial angle

of sclerite on each side of gonopore not produced;

lateral arm narrow, dorsal margin undulate, ventral

margin curving up to acute apex; ventral arm narrow,

curving mesad below. Each duct of spermathecae

slightly longer than bulb, first section short and clear;

second section about as long as bulb and covered with

granular pubescence; third section very short before

expanding to stem of bulb; bulb ovoid with stem about

half as long as distal portion.

VARIATION.—The only other specimen available, a

female, is similar in most respects to the holotype. The

scales on the upper half of the occiput may be yellow.

DISTRIBUTION AND ECOLOGY.—The two specimens

known of caatingensis were collected near a railway

along the river just southwest of Mossoro. The area

was typical caatinga; the soil was sandy and apparently

plants derived subterranean moisture from the river.

This species was taken along with A. inaquosum

Marston. Anthrax caatingensis probably occurs

throughout the desert area of northeastern Brazil.

HOLOTYPE.— $ , Mossoro, Rio Grande do Norte,

Brazil, XII^196 0 (N. Marston-1) (USNM).

PARATYPE.— 9 , Mossoro, Rio Grande do Norte,

Brazil, XII-5-1960 (N. Marston-2) (NLM).

The holotype is in good condition except that the

abdomen is slightly rubbed, the mesonotum is slightly

greasy, and one middle leg is missing. The paratype

lacks one of the third antennal segments and the mesonotum and scutellum are greasy. Both specimens may

be slightly teneral; for that reason the extent of the red

body color has not been emphasized.

DISCUSSION.—Anthrax caatingensis differs from

baliopteros by lacking spots of pigment at the bases of

cells R4, Mi, and 2M2. It may be readily distinguished

from other species with a postmedial spot in cell Rx by

the absence of erect black scales on the sides of abdominal segments two, three, and four.

Anthrax mystaceus (Macquart), new combination

Spongostylum mystaceum Macquart, 1840, p. 53 [Spogostylum].—Kert6sz, 1909, p. 58.—Stuardo Ortiz, 1946, p. 92

[Spogostylum].

Anthrax duodecimpunctatus Philippi, 1865, p. 670 [duodecimpunctata].—Ken€sz, 1909, p. 35.—Edwards, 1930,

p. 172 [decempunctatus].—Oldroyd, 1938, p. 84 [duodecimopunctatus].

MALE.—Body mostly black, purplish or bluish-gray

pollinose; legs, sutures on pleura, posterior margins of

abdominal sterna and basal antennal segment more or

less orange. Front with black setae on upper threefourths, with gold setae just above antennae; setae on

lower two-thirds and lateral margins of face gold, a

few black setae sometimes present medially, white setae

sometimes intermixed along oral margin; scales on

front white below, gold above. Occiput with scattered

yellow setae and scales; fringe of pile on posterior margin yellow. First antennal segment about as long as

apical width; second segment buttonlike, about as

broad as base of third segment; base of third segment

narrow, funnel shaped, tapering to styliform part,

which is one to three times as long as style.

Disc of mesonotum with fine black setae and linear,

hairlike scales, white in submedial longitudinal bands

and extending inward in front of wings, gold elsewhere; lateral margins with longer, fine white and

yellowish-white, semirecumbent scales and gold

bristles; posterior margin with curly, fine white scales.

Scutellum with black setae and bristles and fine scales;

white in lateral and posterior spots, gold elsewhere.

Mesopleuron, sternopleuron, and anterior half of pteropleuron with white hairs and linear or lanceolate

scales, some gold bristles on upper half of mesopleuron.

Prosternum, propleuron, and anterior margin of mesonotum with yellowish-white pile, some black setae on

latter. Metapleuron with dense tuft of white scales

behind spiracle. Postalar tuft of pile white. Coxae with

gold bristles and white scales.

Wing (Plate 6») mostly hyaline; extreme base and

spots at bases of cells R2+3, R4, R5, Mi, 2M2, and Culs
108

and in cell R below base of vein R: light brown. R-m

crossvein near basal two-fifths of cell 1M2, vein R2+3

arising slightly basad; bases of veins R2+3 and R5 with

short spurs more or less as long as r-m crossvein. Contact of cells 1M2 and Cux 1.5 to 2.0 times as long as

base of cell Cux. Cell 2A about as wide as cell 1A; alula

well developed, posterior margin rounded. Calypter

unpigmented, fringe of hairs white. Stem of halter

yellow; knob brown basally, yellow apically.

Femora and tibiae with obovate and oblanceolate,

truncate white scales, yellow scales sometimes present

anterodorsally on femora and anteriorly on tibiae. Fore

femur with scattered bristles anteroventrally toward

base; middle femur with anteroventral row of bristles

and one or several postmedially on anterior side; hind

femur with complete anteroventral and posteroventral

rows of bristles and some dorsally toward apex.

Lateral margins of first abdominal tergum with

yellowish-white pile; lateral margins of posterior terga

with black and yellow setae and yellow and white scales

as continuations of dorsal bands, second tergum with

some gold hairs anteriorly. First and second terga

fringed with curly, fine white scales; terga three

through seven with ovate and lanceolate white scales

posterolaterally, becoming more extensive posteriorly;

remainder of terga covered with linear gold scales

except for mesally interrupted bands of linear black

scales medially on two and anteriorly on three, four,

and five. Sterna with scattered yellow setae and ovate

white scales.

MALE GENITALIA (Figure 134).—Gonocoxites narrow and parallel sided apically in lateral view, broad in

ventral view, and curving inward apically before rounding to glabrous apices; medial sulcus narrow and deep;

scattered fine setae present toward base. Basal segment

of gonostylus not produced dorsally or apically, narrow

in lateral view. Distal segment of gonostylus higher

than broad with apex curled outward and ending in a

sharp distal tooth and blunt medial tooth. Dorsal

part of apex of epiphallus nearly transparent, tapering

toward apex in dorsal view, extending beyond tip of

aedeagus and bluntly rounded apically in lateral view;

an angular projection extending backward above pigmented area formed from dorsal bands. Ventrolateral

part of apex of epiphallus curving downward and

backward to junction with ventral bands. Dorsal

bands simple, broadly united before joining epiphallus.

Epandrium simple, cerci unsclerotized and much

broader than long.

FEMALE.—Similar to male.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FEMALE GENITALIA (Figure 126).—Tenth tergum

with six spines on each side. Ventral arm of ninth

tergum undulating, sharply rounded apically. Sclerite

on each side of gonopore hatchet shaped; ventral arm

parallel sided and slightly curved mesad below; lateral

arm parallel sided, slightly turned upward apically to

sharp dorsal angle. Each duct of spermathecae about

1.5 times as long as bulb; first section of duct short,

second section slightly longer than bulb and covered

with granular pubescence; third section short before

expanding gradually to ovoid bulb.

DISTRIBUTION.—The few specimens available of

mystaceus indicate that it occurs in mountains of central Chile and northwestern Argentina. Specimens

have been seen from "Cuesta, Pajonales," Valle Piquenes, Aconcagua; "Las Mercedes"; "C. Chile"; and

"Chile."

TYPES.—The type specimen of Spogostylum mystaceum Macquart was not found by R. H. and E. M.

Painter in a survey of type specimens of North and

South America Bombyliidae in European museums.

Macquart's rather brief description, however, indicates that it is the species described here. Macquart

indicates that the wings are three times as long as the

abdomen, but this character varies with telescoping

of the abdominal segments. The legs were said to be

black; specimens examined by the author vary only

from red orange to about half black. Macquart's figure

of the wing shows no spots at the bases of cells Mi

and 2M2, although his description indicates spots at

the bases of all principal cells, which was true of all

specimens examined by the author. Macquart's figure

clearly indicates that the crossvein between veins R4

and R2+3 is spurious, a common situation in the genus

Anthrax. Only one of the specimens examined shows

the character, and one has a short spur extending

anteriorly toward vein R2+3 from the basal spur of vein

R4 (Plate 6i) in the same position as the crossvein

figured. Other characters of the species agree with

those described.

The type-locality given by Macquart is "du Bresil

ou du Chili."

Mr. Luis Pefia indicates that he has been unable to

locate Philippi's bombyliid types in the Santiago Museum and that they may have been destroyed in an

earthquake. Philippi's description of duodecimpunctatus indicates that he was referring to this species. The

type-locality is the province of Santiago, Chile.

DISCUSSION.—None of the characters given by Macquart in his description of the genus Spongostylum
NUMBER 43 109

have generic value as they are presently interpreted.

The presence of a sectoral crossvein, particularly, varies

between closely related species, and sometimes within

species. Specimens with a spurious crossvein on one

wing quite often lack it on the other, or have it in

another position.

Anthrax mystaceus appears to be most closely related

to squalidus, although they differ widely. It may be

readily distinguished from squalidus and all other

South American species of Anthrax by the dense patch

of white scales on the metapleuron directly behind the

spiracle.

Anthrax squalidus Philippi

Anthrax squalidus Philippi, 1865, p. 672 [squalida].

Argyramoeba squalida.—Kert^sz, 1909, p. 67.

Aphoebantus squalidus.—Edwards, 1930, p. 171.—Stuardo

Ortiz, 1946, p. 92.

Argyramoeba crinita Bigot, 1892, p. 349 [Argyromoeba].—

Kertdsz, 1909, p. 62.

Spongostylum crinitum.—Kert6sz, 1909, p. 92 [Spogosty-

/um].—Edwards, 1930, p. 172 Icrinita}.—Stuardo Ortiz,

1946, p. 92 [Spogostylum crinita].

MALE.—Body black, tibiae and genitalia reddish;

integument mostly bluish gray pruinose. Front with

black setae and scattered linear white scales below.

Upper half of face bare medially; eye margins and

lower half of face with black setae, few yellow setae

intermixed along oral margins. Occiput with scattered,

short, mixed black and yellow setae; fringe of pile on

posterior margin yellow. First antennal segment about

two-thirds as long as apical width; second segment

biscuit shaped, more flattened laterally; base of third

segment conical, somewhat flattened mesolaterally,

tapering to styliform part; style about one-sixth as long

as third segment.

Disc of mesonotum with fine black setae and scattered, hairlike yellow scales, a few black scales postmedially; margins with longer setae and longer curly,

hairlike white scales. Sternopleuron, mesopleuron, and

anterior part of pteropleuron with sparse, erect, hairlike, white scales, dense black setae and some white pile

present dorsally on last two. Propleuron, prosternurn,

and anterior margin of mesonotum with white pile,

black setae intermixed on mesonotum. Hypopleuron,

metapleuron, and posterior part of pteropleuron bare.

Coxae with black and yellow setae and linear white

scales.

Wing (Plate 6h) nearly hyaline, yellowish at base.

R-m crossvein located at basal third of cell 1M2; vein

R2+3 arising opposite, without basal spur. Vein R4 with

short basal spur about one-third as long as basal section. Contact of cells Cui and 1M2 slightly longer than

width of base of cell Cuj. Cell 1A narrowly open. Cell

2 A wider than cell 1 A, alula not reduced. Calypter unpigmented, fringe of hair white, stem of halter yellow,

knob yellow basally, paler apically.

Femora with ovate-truncate scales, black anterodorsally, white posteroventrally. Fore femur without

macrochaetae; middle femur with anteroventral row;

hind femur with anteroventral and incomplete posteroventral rows. Tibiae with black scales anteriorly, and

white scales posteriorly.

First abdominal tergum with dense yellowish-white

pile laterally. Lateral margins of terga two through

seven with dense black and white pile, and long black

setae. Discs of terga two through five with extremely

fine, recumbent black scales anteromedially, their area

decreasing posteriorly; remainder of discs of two

through five, and six and seven with mixed yellow and

white, fine, curly, recumbent and semirecumbent

scales; all terga with long, erect black setae especially

laterally and along posterior margins. Abdominal

sterna with fine yellow setae, and fine, curly white

scales.

MALE GENITALIA (Figure 130).—Gonocoxites broad,

rounding to acute apices in lateral view, broadly

rounded apically in ventral view; medial sulcus broad,

gonocoxites folded over sulcus and meeting mesally at

apex; setae scattered on gonocoxites from bases nearly

to apices. Basal segment of gonostylus sharply rounded

apically with a dorsomedial, truncate basal fold; a few

fine setae present apically. Distal segment of gonostylus

projecting dorsolaterally, cylindrical, tapering nearly

to apex, which is slightly enlarged distally, with a narrow, dactylate lobe extending laterally from near base

and curved upward apically. Dorsal part of apex of

epiphallus extending upward like a hood in lateral

view, dorsal margin bent downward to sharp apical

angles, which extend above low, truncate, lateral projections; a deep, angular, mesal emargination defined

by sharp angle present in dorsal view; a small nodelike projection present medially just above aedeagus.

Ventrolateral part of apex of epiphallus projecting as

narrow truncate lobes toward apex of aedeagus. Dorsal

bands forming broadly angular, tapering, apically

directed, lateral lobes before joining above aedeagus.

Dorsal margin of epandrium curving downward to

cerci; cerci sclerotized, triangular with sharp ventro-
110

apical angles, fitting over remainder of genitalia when

in normal position.

FEMALE.—No females of this species have been

studied. Notes by R. H. and E. M. Painter show the

female type of crinitus (Bigot) has faint smoky brown

pigment at base of cell R4, base of cell R5, and in the

center of cell R of the wing. Both cells R2+3 and R4

have recurrent stumps less than the length of the r-m

crossvein at their bases.

DISTRIBUTION.—Only two specimens of this species

have been seen by the author, both from 5 miles north

of Laguna Dam, Coquimbo, Chile (Ross and Michelbacher, collectors). The type-locality is the province of

Santiago, Chile.

TYPES.—The types of Anthrax duodecimpunctatus,

squalidus, and bellulus Philippi were not found in any

of the European museums R. H. and E. M. Painter

visited in their study of types of North and South

American Bombyliidae. Some of Philippi's types are

said to be in the Santiago museum, but Mr. Luis Pefia

was unable to find them there. He stated (personal

communication) that they may have been destroyed

during an earthquake.

The type of Argyramoeba crinita Bigot is in the

British Museum (Natural History). According to

Painter and Painter (unpublished notes), it is a female

carrying the labels "Argyramoeba crinita $ n. sp.

Inedit. Aout Quinz 1886. J. Bigot. Chili," and "Chili."

It has been badly covered with fungus but is otherwise

in good condition. A redescription of the type agrees

well with specimens of squalidus except that the wings

have slightly more pigment and cells R2+3 and R4 have

distinct spurs basally.

DISCUSSION.—Edwards (1930) erroneously referred this species to the genus Aphoebantus. He apparently mistook Philippi's statement that the bases

of cells R2+3 and R5 have recurrent spurs to mean that

vein R2+8 arises before the r-m crossvein. The two specimens seen by the author do not have the recurrent

spurs mentioned by Philippi, but this is a highly variable character. The long, erect black setae mentioned in

the description are distinctive. Also, Philippi specifically

mentioned setae at the tip of the third antennal segment, which are not found on the antennae of New

World species of Lomatiinae.

Anthrax plurinotus (Bigot)

Argyramoeba plurinota Bigot, 1892, p. 350 [Argyromoeba].—

Kert&z, 1909, p. 66.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Anthrax plurinotus.—Edwards, 1930, p. 172 [plurinotata].—

Stuardo Ortiz, 1946, p. 93.

MALE.—Body mostly black, pleural sutures, margins

of abdominal segments, tibiae, and basiventral parts

of femora reddish orange. Front with black setae and

few white scales; face completely covered with black

setae, few white setae along oral margin. Occiput with

fine black setae and scattered, lanceolate white scales;

fringe of pile on posterior margin white below, mixed

black and white above, white behind vertex. First and

second antennal segments closely joined, first segment

shorter than apical width; second segment spherical

but flattened basidistally; base of third segment flattened, styliform part arising from outer half, about as

long as style.

Disc of mesonotum with black setae and black, white

and gold, linear scales; posterior margin and lateral

margins in front of wings with curly, lanceolate scales.

Scutellum with black setae and linear scales, white on

anterior half, gold on posterior half. Sternopleuron and

mesopleuron with yellow setae and linear and lanceolate white scales, some gold setae on upper half of mesopleuron. Pteropleuron bare; metapleuron bare. Prosternum, propleuron, and anterior margin of mesonotum with white pile, some black setae on latter; postalar tuft of pile yellow. Coxae with black and white

setae, and linear and lanceolate, white scales.

Wing (Plate 6a) pigmented light brown basally and

anteriorly out to tip of vein Sc in cell Rx, to bases of

cells R2+3, R5,1M2, Cul5 and into basal third of cell 1A;

slightly darker spots present at bases of cells Rx, R2+3

and R5, R4, Mi, 2M2, and Cu,, no isolated spot present

postmedially in cell Rj. R-m crossvein about halfway

from base to apex of cell 1M2, vein R2+3 arising about

length of r-m crossvein basad; contact of cells 1M2 and

Cut about three times longer than width of base of cell

Cuj. Vein R2+3 without basal spur, vein R4 with short

basal spur. Cell 2A narrower than cell 1A, alula reduced, posterior margin straight. Calypter subhyaline,

fringe of hairs yellowish white. Stem of halter yellowish brown, knob brown basally, yellow at tip.

Scales on femora mostly white, black scales anterodorsally toward apices, especially on fore and middle

pairs. Hind tibia with black scales dorsally and white

scales ventrally; fore and middle pairs with white

scales. Middle femur with anteroventral row of macrochaetae; hind femur with anteroventral and posteroventral rows of macrochaetae and some scattered dorsally toward apex.
NUMBER 4 3 111

Lateral margins of first abdominal tergum with white

pile, posterior margin with lanceolate and ovate, white

and gold scales. Lateral margins of terga two through

seven with scattered black setae and few appressed,

white scales. Discs of terga two through seven with

scattered black setae; second tergum with anterior band

of linear gold scales followed by band of linear black

scales interrupted medially by yellow scales, posterior

margin with lanceolate and ovate, yellow and white

scales; third and fourth terga with medially interrupted

bands of black scales anteriorly followed by gold scales,

then white scales along posterior margins; fifth, sixth,

and seventh terga with black scales anteriorly, and

white scales posterolaterally and medially. Venter with

scattered black and yellow setae and white lanceolate

scales.

MALE GENITALIA (Figure 135).—Gonocoxites broad

basally in ventral view, narrow apically with sharp

apices in lateral view; mesal margins curving concavely

to sharp lateral apices in ventral view; setae scattered

over apical parts. Basal segment of gonostylus short,

concave dorsally with rounded apex in lateral view.

Distal segment of gonostylus ovate, dorsoapical margin

curled slightly laterad. Dorsal part of apex of epiphallus forming bluntly rounded preapical, dorsal angle and

right-angled apex in lateral view, narrow with shallow

apical emargination in dorsal view. Ventrolateral part

of apex of epiphallus not well defined from simple,

broad dorsal bands, extending downward and backward from dorsal part. Base of aedeagus narrow, cylindrical, tapering to apex.

DISTRIBUTION AND TYPE.—The type male of Argyramoeba plurinota is in the Bigot collection of the

British Museum. According to notes made by R. H.

and E. M. Painter, it carries the labels "Argyromoeba

plurinotatus [sic] n. sp. Inedit. Quinze Aout 1888. J.

Bigot. Chile" and "Chile." It is in good condition except that the left antenna is broken; the abdomen is

mashed so the genitalia are twisted and concealed. The

specimen may be somewhat teneral.

Only one specimen of this species has been seen by

the author, a male labeled "Llao-Llao, Argentina,

8-1-962." It agrees perfectly with a redescription of

the type made by Painter and Painter, with minor

exceptions due to the slightly teneral condition of

the type. The species probably occurs throughout central Chile and western Argentina.

Anthrax plurinotus differs from those species lacking a postmedial spot in cell Ri on the wing by having

369-673 O—70 8

the setae extend onto the upper half of the face below the antennae and by having a completely bare

pteropleuron.

Anthrax tigrinus Group

The tigrinus group is a small assemblage of species of

distinctive appearance that are parasitoids of carpenter

bees so far as known. The species cannot be confused

with those of any other group in the genus. The distinctive crossvein between the m crossvein and vein

Cui on the wing (Plate 6/-n) will readily separate

them from their congeners. In addition, the large size

of the specimens and the unusual wing pattern with

pigment expanded along the veins to form bands are

peculiar to the group.

The study is based on approximately 450 specimens,

the majority available from museums in North and

South America. The localities of the specimens range

from extreme southeastern Canada, throughout eastern

and southwestern United States, north along the Pacific coast to southern Oregon, along both coasts of

Mexico, and south into Central America. Only one

South American specimen is available (from Venezuela), so that conclusions regarding the Neotropical

distribution of the group are purely conjectural.

Further collecting may show that there are additional

species from South America, or that the distribution

of simson simson includes the Amazon Basin and southeastern Brazil. The numerous species of Neotropical

carpenter bees make it probable that at least one

species of the group occurs in that area.

Four names have been proposed in the group. Until

now, however, all have been generally regarded as

synonyms and the entire group has been considered one

species. Here, five taxa have been discriminated. Anthrax tigrinus, delila, and simson are regarded as good

species while scriptus is a synonym of tigrinus. In

addition, one species, xylocopae, and one subspecies,

simson habrosus, are described as new.

Group Description

Body mostly reddish brown to orange; mesonotum,

upper half of occiput and sometimes third antennal

segment and anterior abdominal terga black. Pruinosity

mostly gray or brown, silver along eye margins. Head

globoid, slightly higher than long; eyes separated by

about 2.5 to 3.0 times width of ocellar tubercle; antennal sockets separated by about 2.0 to 2.5 times their
112

diameter, about 1.5 to 2.0 times their diameter from

eye margin. Antennal area and face produced distinctly above eye margins, face rounded to oral cavity.

Anterior tentorial pits forming deep cavities separating

face from genae. Front covered with numerous black

or reddish setae, denser on upper third, and sparse,

lanceolate, black to yellowish-white scales. Lower half

and lateral margins of face with dense, black or reddish

setae, and intermixed, gold to white, erect lanceolate

scales, setae occasionally extending along midline to

join those on front. Genae with sparse black or reddish

setae. Occiput with sparse black, gold, or mixed setae

and black or yellow scales; fringe of pile on posterior

margin yellow to black with lighter tips above, becoming lighter below. First antennal segment cylindrical or

slightly enlarged apically, about as long as or slightly

shorter than apical width. Second segment short, cuplike, longest dorsomedially, with apical margin forming

a sharp flange surrounding base of third segment. Base

of third segment broadened dorsoventrally, rounded;

styliform part arising near outside edge above middle,

narrow, about as long as dorsoventral width of base;

style very short, with few fine setae apically. Setae on

first and second segments black.

Mesonotum with fine setae scattered over disc and

with coarser setae and bristles along posterior margins.

Scales mostly linear and curly, dense, black in broad,

longitudinal, sublateral bands broken dorsoanteriorly

to base of wing, and in submedial spots of variable size

on posterior half, mixed reddish brown and white elsewhere. Setae on scutellum fine and black; scales lanceolate, white laterally and in posterior patch, sometimes

gold in anteromedial patch. Macrochaetae on mesonotum and scutellum varying from red to black. Pile on

upper margin of mesopleuron and humeral callus

white, bristles red or black. Pile and scales on remainder

of mesopleuron, sternopleuron, and anterior half of

pteropleuron white, red or black; setae red, yellow, or

black. Posterior part of pteropleuron and hypopleuron

bare. Metapleuron bare except for small patch of setae

and scales above hind coxa. Pile on prosternum, propleuron, and anterior margin of mesonotum black,

white, or red. Postalar tuft of pile white. Coxae with

black or red setae and black, white, or gold scales.

Wing (Plate 6j~n) pigmented with light to dark

brown spots and bands along veins and at bases of cells,

hyaline elsewhere. Cells C and Sc light brown to subhyaline except for hyaline areas after humeral crossvein. Base of wing into basal parts of cells R, M, 1A,

and 2A pigmented. More or less coalesced spots extendSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ing from tip of vein 2A across bases of cells Cuj, 1M2,

and 2ML. to bases of cells Ri, and RL.+:, and R5. Pigment

usually extending along vein Rir5 to base of cell R4 and

R5 to wing margin. More or less coalesced spots at bases

of cells M! and 3M_., apically on veins RL.+ ), M,, M2,

and Cui, preapically on vein R2+;t, and in cell Ri below

apex of vein Sc. Veins yellow to dark brown or red.

Base of cell R4 with long or short spur, or connected to

vein R2+3 to form complete sectoral crossvein; medial

angle with short spur. Base of cell RJ+;, with short spur,

arising at or near r-m crossvein, which is about at middle of cell 1M2. Spurious vein between m crossvein and

vein Cui (rarely incomplete). Contact of cells 1M2 and

Cui one-fifth to one-half as long as width of base of

cell Cu,. Cell R5 broadly open. Cell 1A narrowly open

or closed at wing margin. Cell 2A and alula broadened,

wider than cell 1A. Calypter unpigmented, fringe of

scalelike hairs brown nearest wing, white otherwise.

Stem of halter yellow to dark brown, knob light brown

to black, tip yellow.

Scales on legs brown or black. Fore femur with poorly defined anteroventral row of weak or strong macrochaetae. Middle and hind femora with anteroventral

rows of weak to strong macrochaetae (sometimes incomplete) . Few macrochaetae sometimes present anterodorsally toward apex. Fore tibia with anterodorsal,

posterodorsal, and posteroventral rows of macrochaetae. Middle and hind tibiae with more or less complete

rows of macrochaetae at all angles; anterodorsal row

on hind tibia double, with semierect scales intermixed.

Sides of first abdominal tergum with dense, long pile;

sides of posterior terga with sparse or dense, short pile

anteriorly and a few setae and scales posteriorly. Posterior margin of first tergum with lanceolate scales,

long, dense and white laterally, sparse and brown or

white medially; black or red setae present medially.

Discs of remaining terga with fine black or red setae

and small, oblanceolate or obovate scales. White scales

in large posterolateral spots on two, five, and six, and in

small spots posterolaterally on three and four, and submedially on posterior margins of two through seven

(absent on seven in female); white patches more or less

coalesced on posterior terga; white scales sometimes

present anteromedially on two and brown scales sometimes present along posterior margins. Venter with

dense black or red setae and dense, ovate, black or

black and white scales.

Gonocoxites of male genitalia (figure 127) divided

basally, basal lobes curving sharply mesad, then distally
NUMBER 4 3 113

again before joining about halfway to apices; lateral

margins curving gradually to sharp apices in lateral

view, lateral margins curving forward then mesad to

broadly rounded, small apices in ventral view; apices

separated by broad shallow sulcus; medial parts of

gonocoxites swollen before curving to flat apices, setae

moderately numerous. Basal segment of gonostylus a

small, flat plate, about as broad as long. Distal segment

formed dorsolaterally, about three times as long as

basal width in lateral view, bifurcated at apical third

into basally curved, acutely tipped proximal section

and broad, flat distal section with acute distolateral tip

curved upward in lateral view, acuminate apically to

bifurcate apex in dorsal view, with high, dorsally

curved carina with short, sharp, upcurved proximal

end. Dorsal bands long and narrow, uniting about twothirds of way to dorsal carina. Aedeagus highly modified, with vestigial basal part and long, bowed, flat,

narrow distal part curving dorsoproximally and in a

semicircle down and under epiphallus. Apodemes at

base of aedeagus small and upside down in relation to

the genitalia of other species of Anthrax observed. Ventral bands extending proximally and uniting with

aedeagus at the proximal arc. Epandrium deeply emarginate proximally in dorsal view, with long, apically

expanded ventral lobes, with numerous strong setae

on apical part. Cerci unsclerotized, with fine setae.

Eighth tergum of female genitalia (Figure 121)

with dense, fine hairs extending apically; medial, proximal process short, flattened laterally, broadly rounded

dorsally and straight ventrally in lateral view. Ventral

arm of ninth tergum narrow medially, bending proximally and slightly upward at slightly expanded apex;

an interior preapical flange extending ventrally over

edge of sclerite on each side of gonopore; ninth tergum

with numerous, fine long hairs apically. Tenth tergum

with 22 to 32 stiff spines apically on each side surrounding the cerci; ventral arm broadened parallel to

meson, bent inward apically. Sclerite on each side of

gonopore broadened parallel to meson; apex bent

sharply outward, with broadly rounded distal section

and sharp, uncurved proximal tooth; basal section

narrow, bent downward and laterally. Spermathecae

emptying into large, membranous sac extending inward; ducts uniting before emptying into sac, long

and narrow, flexible, convoluted; medial sections more

or less sclerotized; bulbs obovate or oblanceolate in

outline, one bulb distinctly larger than other two.

Evolution

Superficially, the species of the tigrinus group seem to

have no close af; mities with species of other groups of

Anthrax, but on closer examination they have many

characters in common with the pluto subgroup of the

oedipus group. The saucer-shaped second antennal

segment, wing venation, distribution of spots on the

wing and pattern of scales on the thorax and abdomen

are similar. Also, species of both oedipus and tigrinus

groups have similar habits; both parasitize hymenopterous insects nesting in wood or other similar habitats. The species of the tigrinus group simply have

become more specialized in that they are restricted to

carpenter bees so far as known. The two groups may

have evolved from a common ancestor early during

the Pleistocene.

Anthrax delila shows the least specialized characters

in the tigrinus group, which may indicate that the

group was derived from an ancestral stock occurring

in south-central California. If so, a possible line of

evolution in the group might be as follows (Figure 7).

During the Illinoian (third) glacial period the cool

wet climate allowed a connection between the forests of

eastern Arizona and southern California by way of

southern Nevada, and through southern Texas to

southeastern United States and eastern Mexico. The

ancestral form, adapted to forested areas, could have

followed that route and successfully colonized the

areas. During the following interglacial period, the dry

climate broke the forest connections, allowing four

populations to differentiate—delila remaining in California and retaining primitive characters in its stable

environment, xylocopae becoming adapted to dry

forests of eastern Arizona, tigrinus expanding throughout deciduous forests of eastern United States, and

simson becoming adapted to tropical forests of southern

and eastern Mexico. During the Wisconsin (fourth)

glaciation the forests were again connected, but no

great changes would have taken place since the available habitats were occupied. However, the population of simson adapted to dry climate in southern Mexico probably would have been isolated from that in the

wet forests of eastern Mexico and Central America by

the cold uplands at the Isthmus of Tehuantepec, allowing it to differentiate to form the present subspecies,

simson habrosus, which would have been able to migrate northward to fill the vacant ecological area in the

deserts of Sonora, western Arizona, southern California, and Baja California. Anthrax delila probably
114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

colonized southern Baja California during the last

glacial period. A population has since been isolated

there by the recent trend toward warmer and drier

climate in that area. The relatively slight differentiation that has taken place does not seem to warrant

naming a distinct subspecies.

The fact that the Florida population of tigrinus has

not diverged from those in the remainder of the eastern

United States tends to substantiate the evolutionary

scheme given above. Since populations of other species

of Anthrax isolated in Florida during the interglacial

periods show distinct differences from mainland populations, it seems probable that tigrinus did not arrive

in the southeastern United States until the third interglacial period and did not migrate into Florida until

it was reconnected during the fourth glacial period.

Populations of simson may not have crossed the

Isthmus of Panama until the fourth glacial period, since

the one specimen available from Venezuela is not

noticeably different from specimens of simson from

Central America. If that is the case, any populations

found in the Amazon or southeast Brazil probably will

not be noticeably different from the Central American

population.

Taxonomic Characters

The species of the tigrinus group all are quite similar

in general appearance so characters used to separate

them are rather inconspicuous. None of the characters

has been mentioned heretofore in the literature except

that Loew noted the comparatively small, rounded

spots in the outer part of the wing of delila.

The wing pattern is essentially similar in all species

of the tigrinus group, although the wing of delila has

the spots at the bases of cells Mx, 2M2, and 3M2 small,

rounded, and more or less discrete from each other and

the spots on the apices of veins M1} M2, and Cui. This

results in the separation of the spots at the bases of cells

2M2 and 3M2 along vein Cul5 which is the most important diagnostic character for delila. In other species,

the spots in the apicoposterior part of the wing are

larger and expand along the veins to form bands. The

wing pigment is dark brown in xylocopae and tigrinus,

and is similar, but less intense, in delila. In the subspecies of simson the pigment is more reddish brown,

especially in males. Females have darker and more

extensive pigment than males in all species.

The wing veins are dark red or dark brown in

tigrinus, delila, and xylocopae, whereas in the subspecies of simson they are yellow or light red and contrast

with the surrounding pigment. The light veins add to

the reddish-brown appearance of the wings of simson

simson and s. habrosus. The venation is essentially

similar in all of the species, but in most specimens of

s. habrosus the spur at the basal angle of cell R4 is

elongate and connected to vein R2+3 to form a sectoral

crossvein. This crossvein is incomplete in some specimens, sometimes on only one wing, but then the spur

is more than twice as long as the r-m crossvein, whereas

in the other subspecies it is usually less than twice as

long as the r-m crossvein. The crossvein between the m

crossvein and vein Cut, which is used as a diagnostic

character for the tigrinus group, has not been noted in

any other species of Bombyliidae. It is a stable character, incomplete in only one of the specimens studied.

In that case, short spurs on the m crossvein and vein

Cut indicated its position.

The vesture of the body in general offers no good

characters to separate the species. Scales and hairs of

different colors vary greatly in extent between individuals and between sexes. The color of the pile on the

lateral margins of the first abdominal tergum, however,

is a good diagnostic character in one case. In tigrinus,

the pile is black or reddish black on the lower half and

white on the upper half, whereas in other species dark

hairs occupy the anteroventral portion and extend onto

the dorsal surface anteriorly. In delila, a few specimens

from southern California and those from Baja California have the pile almost entirely white or yellowish

white with no dark hairs dorsally, but these specimens

may be readily separated from tigrinus.

Unlike other groups of Anthrax, the male genitalia

of the tigrinus group show no significant variation between species. Their highly modified structure is peculiar to the tigrinus group and they show no relationship to those of any other species in the genus in North

and South America. The female genitalia are less

highly modified, but they also show little variation between species. In specimens examined, the number of

spines on the tenth tergum varies from 22 to 32. Also,

the sclerites on each side of the gonopore and the bulbs

of the spermathecae vary somewhat in form, but in

other respects the genitalia are quite similar. A distinct

feature of the tigrinus group is that one bulb of the

spermathecae is larger than the other two.
NUMBER 4 3 115

Key to the North and South American Species of the

Anthrax tigrinus Group

1. Spot at base of cell 3Mj of wing (Plate 6k-n) broadly connected along vein Cui to spot

at base of cell 2Mi; spots in outer part of wing expanded as bands along veins 2

Spot at base of cell 3Mi of wing (Plate 6;) not connected along vein Cui to spot at base

of cell 2M» (narrow thread of pigment rarely present above vein); spots in outer part of

wing rounded, not expanded as broad bands along veins (except on R« and R»).

California and southern Baja California (Map 26) delila Loew

2(1). Spur at base of cell R« of wing (Plate 6k, m-n) not connected to vein R2+,, usually less than

2.0 times longer than r-m crossvein 3

Spur at base of cell R« of wing (Plate 6/) connected to vein R2<., to form complete sectoral

crossvein, or, if not connected, spur at least 2.0 times longer than r-m crossvein. Southern

California, Baja California, Arizona, and southeast along coast of Mexico to Chiapas

(Map 27) sitnson habrosus, new subspecies

3(2). Lateral margins of first abdominal tergum with black hairs on anterior part of upper

surface. Arizona to Texas, Gulf coast of Mexico, Central America, and eastern and

northern South America 4

Pile on lateral margins of first abdominal tergum white above, black or brown below,

sometimes with white hairs below, but without dark hairs anterodorsally. Extreme

southeast Canada and eastern United States as far west as central Texas (Map 27).

tigrinus (DeGeer)

4(3). Veins on wing (Plate 6m) dark red or brown, not contrasting with dark brown pigment

Central Texas to eastern Arizona and into north-central Mexico (Map 26)

xylocopae, new series

Veins on wing (Plate 6k) light reddish brown, contrasting with surrounding pigment.

Gulf coast of Mexico, Central America, and eastern and northern South America

(Map 27) sitnson sitnson Fabricius

Anthrax delila (Loew)

Argyramoeba delila.—Loew, 1869, p. 28.—Loew, 1872, p.

142.—Osten Sacken, 1877, p. 241.—Osten Sacken, 1878,

p. 90.—CoquiUett, 1894, p. 95.—Kertesz, 1909, p. 67.

Spongostylum delila.—Aldrich, 1905, p. 222 [Spogostylum\.—Nininger, 1916, p. 162.

Anthrax delila.—Painter and Painter, 1965, p. 432.

Argyramoeba simson.—Davidson, 1893, p. 153 [not Fabricius,

1805; misidentification].

Spongostylum simson.—Aldrich, 1905, p. 223 (part) [Spogostylum] [not Fabricius, 1805; misidentification].

Anthrax simson.—Priddy, 1939, p. 45.—Hurd, 1959, p. 54

[not Fabricius, 1805; misidentification].

MALE.—Setae on front and face mostly black or

reddish brown, gold setae sometimes present on face

and lower part of front; scales on upper part of front

usually black, on lower part mixed gold and yellowish

white, on face mixed white and yellowish white. Fringe

of pile on posterior margin of occiput gold or light

brown basally, with white tips, rarely entirely white.

Scales on mesopleuron, sternopleuron, and anterior

part of pteropleuron white or yellow, black scales sometimes on anterodorsal part of pteropleuron; setae usually black, sometimes partly reddish brown, or partly or

entirely yellow. Prosternal pile usually mixed brown

and white, sometimes entirely brown or entirely white.

Propleural pile usually white or yellowish white, sometimes partly or largely brown. Scales on coxae mixed

yellowish white, white and black or entirely white.

Wing pattern as in Plate 6;. Spots in outer part of

wing not elongated along veins except on veins R4 and

R5; spots at bases of cells Mx and 3M2 separated from

spots at apices of veins Mx, M2, and Cui; spots at base

and apex of cell 2M2 not connected along vein Cui,

or only very narrowly connected by thin thread of pigment. Vein dark brown, of same color as pigment or

darker. Base of vein R4 with short spur, but not connected to vein R2+3 to form sectoral crossvein.

Pile on lateral margins of first abdominal tergum

usually entirely white, sometimes partly brown or black

anteroventrally. Dorsum of abdomen with some brown

scales in addition to black and white scales. Setae on

venter black to yellowish, scales entirely white, or rarely

with few black scales on first sternum and anterior part

of second.

MALE GENITALIA.—See group description and Figure

127.

FEMALE.—Similar to male. Pile on prosternum and

propleuron entirely brown or black or with some white

hairs, rarely predominantly white. Setae on remainder

of pleura and coxae black. Scales on mesopleuron,
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sternopleuron, and anterior part of pteropleuron white

or yellowish white, black scales usually present on

upper half of sternopleuron. Wing pattern slightly more

extensive than in male, but spot at base of cell 3M2

not joined to those at bases of cells Mx and 2M2. Pile

on lateral margins of first abdominal tergum usually

black or dark brown on anteroventral half to threefourths, rarely almost entirely white. Setae and scales

on venter of abdomen entirely black or few white scales

present along posterior borders, especially posteriorly.

FEMALE GENITALIA (Figure 121).—Ninth tergum

with about 22 spines on each side. Middle sections of

ducts of spermathecae lightly sclerotized, only slightly

darker than other sections; smaller bulbs oblanceolate

in outline, about 2.75 times longer than maximum

diameter; larger bulb obovate, about 2.0 times maximum diameter.

DISTRIBUTION AND ECOLOGY.—Anthrax delila occurs

in forested areas from northern California south along

the Coast Range and the west side of the Sierra Nevada

into the mountainous area north and east of Los

Angeles, and at the southern tip of Baja California Sur,

Mexico; perhaps also in forested areas in extreme

southwestern California and on mountains in central

and northern Baja California (Map 26).

MAP 26.—Distribution of Anthrax delila (dots and vertical

lines) and A. xylocopae (squares and horizontal lines).

TYPE.—The type male of Argyiamoeba delila Loew

is in the Museum of Comparative Zoology, Harvard

University. Notes on the type made by R. H. Painter

indicate that it is the species described here. The type

specimen is lighter than most specimens of the species,

with numerous brown scales and few black hairs on

the dorsum of the abdomen. The type-locality is

"California."

BIOLOGY.—Anthrax delila has been reared from the

nests of Xylocopa tabaniformis orpifex Smith by Nininger (1916) and Davidson (1893). Hurd (1959) reviewed information provided by them. The life cycle

is typical of the species of Anthrax studied.

DISCUSSION.—Three males and one female collected

in southern Baja California differ from typical California specimens by the lighter pile and scales on the

thorax and abdomen. Males have the pile on the

prosternum, propleuron, and first abdominal tergum

entirely white or yellowish white, and the setae and

scales on the mesopleuron, sternopleuron, and anterior part of the pteropleuron yellow and white. The

female has only a few brown hairs on the propleuron

and first abdominal segment while the prosternum is

black haired. Setae on the remainder of the pleura are

black, while the scales and pile are white and yellowish

white. Since a male similar in all respects to those from

southern Baja California has been taken with several

darker males at Monrovia Canyon, Los Angeles

County, California, the two populations are regarded

as conspecific. Separation of the populations is probably a recent result of the progressively drier climate

since the last glacial period. Intermediate populations,

however, may still exist on the mountains in central

and northern Baja California.

Anthrax delila differs from other species in the

tigrinus group by the separation of the spots at the

bases of cells 2M2 and 3M2 on vein Cux by a broad

hyaline area. In a few specimens an almost imperceptible thread of pigment exists along vein Cux, but

never a broad band as in other species. Anthrax delila

is partially sympatric with simson habrosus in southern

California and Baja California. However, delila apparently is indigenous to wetter, forested areas, whereas

s. habrosus occurs in drier, desert habitats. In addition

to the character given above, s. habrosus may be distinguished by a complete sectoral crossvein or by having the spur at the base of cell R4 at least twice as long

as the r-m crossvein.
NUMBER 43

117

Anthrax xylocopae, new species

Argyr amoeba tigrina.—KertSsz, 1909, p. 67 [part] [not DeGeer, 1776; misidentification].

Anthrax tigrinus.—Maughan, 1935, p. 32 [tigrina (DeGeer)].—Painter and Painter, 1965, p. 432 [part] [not

DeGeer, 1776; misidentification].

Anthrax simson.—Hurd, 1959, p. 57 [not Fabricius, 1805;

misidentification].

MALE.—Setae on front, face, and genae black; scales

on front mostly black, some white ones laterally below;

scales on face white. Fringe of pile on posterior margins

of occiput dark brown with light tips. Pile on prosternum and propleuron black. Setae and scales on mesopleuron, sternopleuron, anterior part of pteropleuron,

and coxae black, few light scales posteriorly on

pteropleuron.

Wing pattern as in Plate 6m; pigment dark brown,

veins slightly darker. Pigment in outer part of wine

expanding along veins to form bands; pigment at base

of cell 3M2 connected to that at bases of cells Mi and

2M2. Vein R4 with short spur at base, not connected to

vein R2+3 to form sectoral crossvein.

Scales on dorsum of abdomen entirely black except

for patches of white scales along posterior margins of

segments. Pile on lateral margins of first tergum black

on anteroventral three-fourths, white posterodorsally.

Venter of abdomen with black setae and scales, few

white scales present along margins of posterior terga.

MALE GENITALIA.—See group description and Figure 127.

FEMALE.—Similar to male. Black scales medially on

face. Scales on mesonotum entirely black.

FEMALE GENITALIA.—Similar to simson habrosus

(Figure 121). Ninth tergum with about 28 spines on

each side. Middle sections of ducts of spermathecae

lightly sclerotized; smaller bulbs of spermathecae obovate in outline, about twice maximum width; third

bulb distinctly larger than other two.

VARIATION.—The white scales on the front may be

more extensive than in the types. White scales may be

absent from the abdominal venter in females, or may

be present across the apices of all segments in males.

The pile on the sides of the first abdominal tergum

may be as much as nine-tenths black. The pigment on

the m crossvein may be narrowly broken between the

bases of cells Mx and 3M2.

DISTRIBUTION AND ECOLOGY.—Anthrax xylocopae

occurs in areas of desert grassland and probably also

in pinion-juniper forests from western Texas to central

Arizona (Map 26). It also occurs in northwestern

Mexico, possibly farther south on the central plateau

than shown on the map. A specimen from Zion National Park, Utah (Maughan, 1935), may be either

xylocopae or simson habrosus. In either case, the distribution of the species must extend into southern Utah.

HOLOTYPE.— <S, 7 mi. SE Rodeo, Hidalgo Co., N.

Mex., VIII-21-1958 (M. A. Cazier) (UCAL).

ALLOTYPE — $, 7 mi. SE Rodeo; Hidalgo Co., N.

Mex., VIII-21-1958 (C. G. Moore) (UCAL).

PARATYPES.—Arizona: Cochise Co., 3cf, Carr Canyon, Huachuca Mts., VIII-1905 (H. Skinner)

(ANSP); 2c?, Huachuca Mts. (ANSP); ? , Huachuca

Mts. (Oslar) (ANSP); d1, Huachuca Mts., VII-15-

1934 (Ar. E. D. Ball) (ARIZ): <f, Huachuca Mts.,

VIII-22-1935 (Jean Russell) (RHP); 2d\ Ramsey

Canyon, Huachuca Mts., IX-5-1955 (F. G. Werner

and G. D. Butler) (ARIZ); ? , Portal (N. Mex.), IX2-1958 (H. V. Weems, Jr.) (FSPB); 3d\ 5 mi.

W Portal (N. Mex.), VIII-6, 12-1958 (P. Opler)

(UCAL, LINSDALE). Coconino Co., $, Bright

Angel Trail, Grand Canyon VIII-4-1917 (R. C. Shannon) (Cornell U. Lot 869, Sub. 2) (CNLL); 3d1

, Oak

Creek Canyon, VIII-17, 22-1954 (K. Doering)

(KANS, NLM) 5 2c?,?, Oak Creek Canyon, VIII-5,

9-1961 (G. C. Eickwort, R. G. Tucker) (MSU,

NLM). Gila Co., ?, Globe, VIII-9-1933 (Parker, 33)

(CAS); $, Wheatfields, near Globe, August (D. K.

Duncan) (AMNH). Pima Co., d1, Ajo Mts., X-17-

1935, 2600 feet (O. Bryant) (CAS); o", Apache

Camp, Santa Catalina Mts., VIII-25-1917, 5500 feet

(MCZ); cf, Baboquivari Canyon, W side Baboquivari

Mts., VII-25-27-1952 (H. B. Leech and J. W. Green)

(CAS); $, Esterero Canyon, Santa Catalina Mts., VI19-1959 (Radford, Patterson, and Samuelson)

(ARIZ); $, Greaterville, X-9-1954 (F. G. Werner)

(Bacchant) (ARIZ); <$, Kits Peak, VIII-1-^1916

(Rincon) (AMNH); <f, Sabino Canyon, VII-20-

1917 (CNLL); cf, Sabino Canyon, Santa Catalina

Mts., VIII-23 (J. Bequaert) (MCZ); 4d", Santa Rita

Mts., VIII-18-1935 (Jean Russell, Jack Beamer)

(KANS, NLM); <f, Santa Rita Mts., VII-17-1932

(J. D. Beamer) (KANS); tf, Santa Rita Mts., VII19-1935 (JeanRussell) (KANS); ?,Tucson,VI1-5-

1960 (F. Werner) (ARIZ). Santa Cruz Co., ? , CanCanyon, VIII-8-1962 (S. L. Wood and J. B. Karren)

(KANS); Garces, VIII (Biedermann). 3 <?, 2 ? ,

Palmerlee, July, August (N. Banks collection)

(UCAL, MCZ).

New Mexico: Hidalgo Co., d, Rodeo, IX-17-1958

(P. D. Hurd) (reared from Xylocopa californica ariz-
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

onensis Cr.) (UCAL); d", 1.5 mi. S Rodeo, IX-15-

1958 (P. D. Hurd) (reared from Xylocopa californica

arizonensis Cr.) (NLM); d\ ° , 7 mi. SE Rodeo,

VIII-21-1958 (J. M. Marston, C. G. Moore) (UCAL,

NLM); d\ 18 mi. N Rodeo, VIII-25-1958 (P. M.

Marsch) (UCAL).

Texas: San Saba Co., ? , Mouth of Brady Creek,

VII-23-1931 (RHP). Brewster Co., S, Lajitas, IX6-1960 (J. E. Gillaspy) (CAS).

BIOLOGY.—According to Hurd (1959), xylocopae

has been reared from nests of Xylocopa californica arizonensis Cresson at a locality 1.5 miles south of Rodeo,

Hidalgo Co., N. Mex., and 40 miles southeast of

Nuevas Casas Grandes, Chihuahua, Mexico. The bee

nests in dead-flower stalks of Yucca and related plants.

DISCUSSION.—Anthrax xylocopae may be distinguished from tigrinus by the black hairs anterodorsally

on the first abdominal tergum. It differs from delila by

the spot at the base of cell 3M2 being broadly connected

along vein Cu! with the spot at the base of cell 2M3

(Plate 6m). Anthrax simson differs from xylocopae by

having the wing pigment light brown and the veins yellowish brown and distinctly lighter than the pigment.

The scales on the abdominal sterna are largely white in

males of simson, whereas males of xylocopae have

broad areas of black scales.

Anthrax xylocopae and simson habrosus have been

taken together in several localities in central Arizona.

It seems, however, that s. habrosus prefers drier habitats, although their habitats probably broadly overlap

at edges of desert valleys. Besides characters given,

s. habrosus differs from xylocopae by having a

complete sectoral crossvein in most specimens. The

few specimens of s. habrosus in which the sectoral

crossvein is incomplete have the spur at the base of cell

R4 at least twice as long as the r-m crossvein, whereas

in xylocopae it is seldom more than 2.0 times as long.

The relationship between xylocopae and tigrinus is

discussed under the latter.

Anthrax tigrinus (DeGeer)

Nemotelus tigrinus DeGeer, 1776, p. 206.

Argyramoeba tigrina.—Kerte'sz, 1909, p. 67 [part].—Rau,

1926, p. 231.

Anthrax tigrinus.—Painter and Painter, 1965, p. 432 [part].

Anthrax scripta Say, 1823, p. 43; 1869, p. 59.

Anthrax simson.—Wiedemann, 1828, p. 259.—Macquart,

1840, p. 59.—Walker, 1849, p. 251.—Osten Sacken, 1858,

p. 41 [part].—Johnson, 1925, p. 108.—Curran, 1927.

p. 85.—Brimley, 1938, p. 341.—Hurd, 1959, p. 56 [not

Fabricius, 1805; misidentification].

Argyramoeba simson.—Osten Sacken, 1877, p. 241.—Osten

Sacken, 1878, p. 90 [part].—Coquillett, 1894, p. 95.—

Johnson, 1895, p. 325 [not Fabricius, 1805; misidentification].

Spongostylum simson.—Aldrich, 1905, p. 223 [part] [Spogostylum].—Johnson, 1913, p. 55 [Spogostylum].—Malloch,

1917, p. 393 [Spogostylum].—Cole, Malloch, and McAtee,

1924, p. 186 [Spogostylum] [not Fabricius, 1895; misidentification].

MALE.—Setae on front and face black, or setae on

lower half of front and face partly or entirely reddish

brown. Scales on upper part of front black, on lower

half mixed gold and yellowish white, often some black

scales present; scales on face mixed white and yellowish white, often some black scales present. Setae on

occiput reddish brown, some longer black ones at

vertex; scales gold or brown; fringe of pile on posterior

margin light or dark brown basally, white or light

brown apically, lighter below.

Setae on mesonotum black; black scales intermixed

with lighter ones along meson. Disc of scutellum with

spot of gold scales anteromedially (sometimes reduced) . Setae on sternopleuron, mesopleuron, and anterior part of pteropleuron black, or with some reddishbrown setae intermixed above; scales on sternopleuron

and pteropleuron white, brown or mixed, on mesopleuron black anteriorly, brown or white posteriorly;

upper margin of mesopleuron with white pile. Pile on

prosternum and propleuron black, reddish brown or

mixed. Setae and scales on coxae black.

Wing pigmented as in Plate 6n. Pigment dark

brown; veins dark reddish brown, not contrasting with

pigment. Pigment at base of cell 3M2 broadly connected to spot at base of cell 2M along vein Cui, and

usually connected along m crossvein to spot at base of

cell Mi. Spots at apices of veins M2 and Cui broadly

connected to spots at bases of cells Mt and 3M2. Spur

at basal angle of vein R4 not connected to vein R2+3j

usually less than 2.0 times longer than r-m crossvein.

Lateral margins of first abdominal tergum with black

or reddish brown pile ventrally and white pile dorsally,

sometimes with some white hairs posteroventrally, but

without black hairs anterodorsally. Dark scales on terga

mostly black, some light-brown scales usually intermixed along posterior margin and medially on second

tergum. Setae on dorsum black on discs, reddish along

margins. Setae on venter reddish brown; scales black

on discs, white and yellow posterolaterally and along

posterior margins, light scales more numerous

posteriorly.
NUMBER 43 119

MALE GENITALIA.—See group description and Figure 127.

FEMALE.—Setae on head mostly or entirely black.

Light scales on mesonotum and pleura reduced in

number or entirely absent. Spot of brown scales anteriorly on scutellum absent. Setae on abdomen black

(rarely some reddish setae on venter) ; white scales on

venter restricted to posterior sterna and lateral margins

of first sternum.

FEMALE GENITALIA.—Similar to simson habrosus

(Figure 121). Ninth tergum with about 32 spines on

each side. Ducts of spermathecae with middle sections

sclerotized; bulbs of spermathecae obovate in outline,

about twice as long as maximum diameter and broadest medially.

DISTRIBUTION AND ECOLOGY.—Anthrax tigrinus may

be found in forested areas from extreme southeast Canada throughout eastern United States to central Texas

(Map 27). The distribution appears to be limited on

the north by coniferous forests and on the west by the

Great Plains. In western Texas tigrinus seems to be

replaced by xylocopae, the latter being adapted to areas

of grassland with xerophytic shrubs.

TYPES.—The two cotypes of Nemotelus tigrinus

DeGeer are in the Naturhistoriska Riksmuseet, Entomologiska Avdelningen, Stockholm, Sweden. Dr. KarlJohan Hedqvist (personal communication) has compared a photograph of the wing of a species of the

tigrinus group with the type and found them closely

similar- He also stated that the pile on the lateral margin of the first abdominal tergum is white above and

black below, a characteristic of tigrinus. Neither specimen carries any label. The type-locality is Pennsylvania.

The type of Anthrax scripta Say has apparently been

destroyed with the remainder of Say's collection. The

excellent description leaves no doubt that the specimen

is a member of the tigrinus group, and since the typelocality is Pennsylvania, it is a synonym of tigrinus.

BIOLOGY.—Anthrax tigrinus has been recorded as a

parasite of Xylocopa virginica by Angus (1868) and

Rau (1926). Hurd (1959) gives an excellent review

of the information they compiled, as well as information on the biology of other species in the tigrinus

group.

DISCUSSION.—Anthrax tigrinus is characterized by

having the pile laterally on the first abdominal tergum

black below (sometimes a few light hairs posteriorly)

and entirely white above. In most specimens of other

species, dark pile occupies the anteroventral part of

the margins with few or many dark hairs anteriorly on

the dorsal part. In a few specimens of other species,

particularly delila, the pile laterally on the first tergum

is largely or entirely white or yellowish, but there are

no dark hairs dorsally.

Anthrax tigrinus is most similar to xylocopae and

the two seem to have been relatively recently derived

from a single population. The two species apparently

now have different ecological preferences, however,

xylocopae parasitizing carpenter bees nesting in flower

stalks of Yucca and related plants, tigrinus parasitizing

bees utilizing hardwood billets and structural beams for

nesting sites. Anthrax xylocopae may have become

adapted to a drier climate during the preceding interglacial period when forests retreated from between

central Texas and southeast Arizona. Further collecting may show hybridization between the two populations, although, with their apparent ecological isolation, that seems unlikely.

Anthrax simson simson Fabricius

Anthrax simson Fabricius, 1805, p. 19—Osten Sacken, 1858,

p. 41 [part].—Hurd, 1959, p. 58 [part].

Argyramoeba simson.—Osten Sacken, 1878, p. 90 [part].—

Osten Sacken, 1886, p. 100.

Spongostylum simson.—Aldrich, 1905, p. 223 [part]

[Spogostylum].

Argyramoeba tigrina.—Kert6sz, 1909, p. 67 [part] [not

DeGeer, 1776].

Anthrax tigrinus.—Painter and Painter, 1965, p. 432 [part]

[not DeGeer, 1776].

MALE.—Setae on front and face reddish brown and

black; scales on front black above, mixed black and

yellowish white below; scales on face gold and yellowish white; setae on genae black. Fringe of pile on posterior margin of occiput brown or yellow basally,

white apically.

Setae on mesonotum black or partly reddish brown;

area of light scales more extensive than area of black

scales. Scutellum often with some gold scales

anteromedially. Setae on mesopleuron, sternopleuron,

and anterior part of pteropleuron reddish brown and

black; scales mixed gold and white, some black scales

above on mesopleuron; upper part of mesopleuron

with white pile above and black pile below. Pile on

prosternum and propleuron light or dark brown. Setae

on coxae black or partly reddish brown; scales gold,

some white scales on anterior pair.

Wing pigmented as in Plate 6k. Pigment dark

brown, veins light brown and contrasting with pig-
120

ment. Pigment at base of cell 3M2 broadly connected

with spot at base of cell 2M2 along vein Cuj, usuallyseparated from pigment at base of cell Mi on m crossvein. Spots at apices of cells M2 and Cui broadly connected to spots at bases of cells Mj and 3M2. Spur

at base of cell R4 not connected to vein R2+3 to form

sectoral crossvein, less than 2.0 times as long as r-m

crossvein.

Abdominal terga with reddish-brown setae along

margins, black setae anteriorly. Pile on lateral margins

of first tergum light reddish brown to reddish black on

anteroventral half to three-fourths, some white hairs

below in lighter specimens. Setae on venter reddish

brown; scales white laterally and along posterior margins, dark brown anteromedially.

MALE GENITALIA.—See group description and Figure

127.

FEMALE.—Setae on face, front, thorax, and abdomen

black. Pile on prosternum and propleuron black; scales

on mesopleuron mostly black. Dark pile on first

abdominal tergum black; white scales less extensive

on sterna.

FEMALE GENITALIA.—Similar to simson habrosus

(Figure 121).

DISTRIBUTION AND ECOLOGY.—The few specimens

available indicate that simson simson occurs along the

east coast of Mexico from southern Tamaulipas southward throughout Central America, and along the

Caribbean coast to Venezuela (Map 27). It may also

occur southward into eastern and southern Brazil,

although no specimens have been seen from there.

In southeastern Mexico s. simson is apparently

allopatric to s. habrosus, although few specimens are

available from that area. They may occur together in

Chiapas and eastern Oaxaca, although s. simson

probably is largely restricted to wetter habitats along

the Atlantic side, whereas s. habrosus occurs in the

drier Pacific side.

TYPE.—According to Painter and Painter (personal

notes) the type male of Anthrax simson Fabricius is in

the Universitets Zoologiske Museum in Copenhagen.

It carries the red type label and another label "A.

simson ex Am. Mer. Schmidt." The antennae are missing and the specimen is dirty and somewhat greasy.

A photograph of the type indicates that it is the species

described here. Fabricius gave the type-locality as

"Habitat in America Meridionali."

BIOLOGY.—The only reference to the biology of s.

simson is that of Hurd (1959, p. 58) :

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Many cells of this species [Xylocopa augusti Lepcletier] located on January 13, 1956, in a hardwood fence post at Araucaria, Parana, Brasil, were found l>y Dr. C. D. Michener

and Padre J. S. Moure to contain larvae and pupae of an

undetermined bombyliid (Hurd, 1958). This is the first record

of beefly parasitism of a South American Xylocopa. Since Anthrax simson is known to occur in the vicinity where the nest

was located, it quite possibly may have been the species involved.

Since no specimens have been seen from south of

Venezuela, validity of this record cannot be verified.

DISCUSSION.—One specimen of s. simson, from

Tamazunchale, San Luis Potosi, Mexico, lacks the

crossvein between the in crossvein and vein Cu! which

is characteristic of species of the tigrinus group.

Anthrax simson simson differs from (ielila, xylocopae, and tigrinus by having the veins of the wing light

reddish brown, contrasting with the surrounding pigment, rather than dark brown or red and not contrasting. The pigment also is distinctly lighter in s.

simson. The relationship of s. simson to s. habrosus is

discussed under the latter.

Anthrax simson habrosus, new subspecies

Argyramoeba simson.—-Williston, 1901, p. 275 [not Fabricius,

1805; misidentification].

Spongostylum simson.—Aldrich, 1905, p. 223 [part] [Spogostylum] [not Fabricius, 1805; misidentification].

Argyramoeba tigrina.—Kertesz, 1909, p. 67 [part] [not DeGeer, 1776, misidentification].

Anthrax tigrinus.—Painter and Painter, 1965, p. 432 [part]

[not DeGeer, 1776, misidentification].

MALE.—Setae on face and front reddish brown.

Scales on front mostly yellowish white and tan, darker

above; scales on face yellowish white and tan. Setae

and scales on genae tan. Fringe of pile on posterior margin of occiput yellowish brown basally, white apically.

Setae on mesonotum reddish brown. Scutellum with

large anteromedial patch of gold scales. Gold and white

scales more extensive than black scales on mesonotum.

Pile on prosternum and propleuron light brown. Setae

on mesopleuron, sternopleuron, and anterior part of

pteropleuron light reddish brown; scales white, upper

part of mesopleuron with dense white pile. Coxae with

dark reddish-brown setae and gold scales.

Wing pigmented as in Plate 6/. Pigment brown,

veins light brown, contrasting. Spur at base of cell R+

connected to vein R2+3 to form complete sectoral crossvein. Spot at base of cell 3M2 broadly connected along

vein Cux to spot at base of cell 2M2, but not connected

to spot at base of cell Mj.
NUMBER 4 3 121

Pile on lateral margins of first abdominal tergum

reddish brown posteroventrally to black anterodorsally

on anteroventral nine-tenths. Dark setae on terga reddish brown; few light brown scales intermixed with

black scales. Setae on venter reddish brown. Scales

light brown basally and white apically on proximal

segments, entirely white on distal segments.

MALE GENITALIA.—See generic description and Figure 127.

FEMALE.—Similar to male. Setae on front mostly

black; some black setae intermixed on face. Setae on

mesonotum, thoracic pleura, and coxae black. Pile on

prosternum and propleuron dark reddish brown and

black. Scales on mesopleuron, sternopleuron, and anMAP 27.—Distribution of Anthrax tigrinus (triangles and vertical lines), simson simson (squares

and diagonal lines), and A. simson habrosus (circles and horizontal lines).
122

terior part of pteropleuron mixed gold and white. Dark

pile on lateral margins of first abdominal tergum black:

setae black on anterior parts of terga. Setae on venter

dark reddish brown; scales mostly black, some white

scales laterally along posterior margins of sterna.

FEMALE GENITALIA (Figure 121).—Ninth tergum

with about 22 spines on each side. Middle sections of

ducts of spermathecae lightly sclerotized, slightly

darker than other sections; smaller bulbs slightly asymmetrical, obovate in outline, about 2.0 times longer

than maximum diameter, larger bulb only slightly

broader than smaller ones.

VARIATION.—Males may have numerous black setae

intermixed with reddish brown ones on the front,

mesonotum, and abdomen. The setae on the pleura

may be partly or entirely black and there may be

numerous black scales, especially on the mesopleuron.

The pile on the prosternum and propleuron varies

from light brown to black. The dark pile on the lateral

margins of the first abdominal tergum varies from

light tan to black, being darker on females. Gold or

brown scales may be absent on the scutellum and

abdominal terga. The white scales on the abdominal

sterna may be less extensive in both sexes. The sectoral

crossvein on the wings is incomplete in some specimens,

in which case the spur at the base of cell R4 is about

twice as long as the r-m crossvein. The pigmented

area at the base of cell 3M2 often is connected to that

at the base of cell Ma.

DISTRIBUTION AND ECOLOGY.—Anthrax simson habrosus occurs in desert and semidesert areas from Arizona, southern California, and Baja California south

along the Pacific coast to Chiapas, Mexico, and possibly into western Guatemala (Map 27). It is allopatric to delila in southern California. Although the

two have been collected together at several localities,

A. s. habrosus and A. xylocopae are generally allopatric in east-central Arizona. In southeastern Mexico,

s. habrosus appears to be allopatric to s. simson, the

former occupying drier areas on the Pacific side; the

latter, wetter areas on the Atlantic side.

HOLOTYPE—cf, Tucson, Pima Co., Arizona, VI-17-

1954 (R. S. Beal) (A taken in copulation with specimen "B") (UCAL).

ALLOTYPE.— 9 , Tucson, Pima Co., Arizona, VI-17-

1954 (R. S. Beal) (B taken in copulation with specimen "A") (UCAL).

PARATYPES.—Arizona: Coconino Co., 2cf, Bright

Angel Trail, Grand Canyon, VIII-4-1917 (E. C.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Shannon) (Cornell U. Lot 869, Sub. 2) (CNLL,

RHP); d, Oak Creek Canyon, VIII-4-1961, 3500

feet (G. C. Eickwort) (MSU) ; <f, 15 mi. N Sedona,

IX-14-16-1955 (G. D. Butler) (ARIZ). Gila Co., 9,

CapitanMt.,VIII-3-1933 (R.Anderson) (CAS) ; $,

Globe (D. K. Duncan) (AMNH) ; <?, Midland Pioneer Camp, Pinal Mts., YIII-16-1950, 5000-6100

feet (T. Cohn, P. Boone, and M. Cazier) (AMNH) ;

cT, base of Pinal Mts., August (D. K. Duncan)

(AMNH); d\ Roosevelt, VIII-22-1933 (Parker)

(CAS); 9 , Tonto National Mon., VIII-26-1932

(R. H. Painter) (RHP) ; rf, Wheatfields, near Globe

(D. K. Duncan) (AMNH). MaricopaCo., J, Phoenix, 1934 (R. J. Crandell) (AMNH). Pima Co., 9,

Arivaca, VII-17-1934 (L. P. Wehrle) (ARIZ); d\

Baboquivari Mts., VII-19-1932 (R. H. Beamer)

(KANS) ; 9 , Esterero Canyon, Santa Catalina Mts.,

VI-19-1959 (Radford, Patterson and Samuelson)

(ARIZ) ; 2d\ VII-15, IX-30-1939 (Bryant) (CAS,

NLM); d\ Tucson (F. H. Snow) (C. H. Curran

collection, Ace. 31144) (AMNH) ; 9, Tucson, VII5-1946 (R. H. Crandall) (NLM) ; d\ Tucson, VI11-1961 (F.Werner) (ARIZ); 9 , 18 mi. NE Tucson,

VI-14-1961, 2900 feet (R. H. and E. M. Painter)

(RHP). Santa Cruz Co., 2 9, Patagonia, VIII-6-

1958 (C. W. O'Brien) (Linsdale). Yuma Co. <S,

Yuma, IX-11-1959 (D. Mose) (ARIZ).

California: Los Angeles Co., 9 , Downey, VII-8 -

1940 (M. M. Barnes) (CAS). Orange Co., d, Brea,

VIII-26-1923 (A. J. Basinger) (UCAL); 9 , Newport Bay, VI-31-1940 (P. D. Hurd) (UCAL). Riverside Co., 2d, Riverside, VIII-17, IX-1-1947 (T. E.

Leigh) (UCAL); rf, Riverside, VII-19-1936 (G. P.

Englehardt) (RHP); tf, Riverside, VIII-27-1962

(H. E. Evans) (RHP) ; d\ Riverside, August (A. L.

Melander) (USNM). San Bernardino Co., 2<S, Redlands, IX-13 (UCAL, NLM).

Mexico: Baja California Sur., c?, Cerralbo Island,

Gulf Calif., VI-5-1921 (E. P. VanDuzee) (CAS);

cf, Las Animas, Sierra Laguna, X-12-1941 (Ross and

Bohart) (CAS) ; 2 9 , La Paz, VI-3-1921 (E. P. Van

Duzee) (CAS, NLM) ; tf, 5 mi. S Miraflores, VII10-1938 (Michelbacher and Ross) (CAS); <?, 5 mi.

W San Bartolo, VII-13-1938 (Michelbacher and

Ross) (CAS) ; 9 , 3 mi. N San Pedro, VII-6-1938

(Michelbacher and Ross). Chiapas, tf, 4 mi. SE

Soyalo, 111-15-1953 (R. C. Bechtell, E. I. Schlinger)

(UCAL). Durango, 9 , El Saldado, VI-22-1961, 7600

feet (R. A. Scheibner) (MSU). Guerrero, cf, between
NUMBER 4 3 123

Cajones and Rincon, S of Chilpancingo, VII-1-1932

(H. Smith) (RHP); d, 25 mi. S Iguala, IX-14-1963,

2400 feet (R. H. and E. M. Painter) (RHP).

Michoacdn, d, Apatzingan, VII-21-1947 (T. H.

Hubbell) (MU). Morelos, d, 3 mi. N Alpuyeca, IV18-1959, 3400 feet (H. E. Evans) (RHP) ; d, Canyon

de Lobos, Yautepec, IV-13-1959, 4000 feet (H. E.

Evans) (RHP); d, Cuernavaca (R. Muller)

(USNM); d, Cuernavaca, III-1O-1959, 5500 feet

(H. E. Evans) (RHP); d, Huajintlan, IV-11-1959,

2800 feet (H. E. Evans) (NLM). Nayarit, d, nr.

Compostela, V-7-VIII-17-1937 (E. Rosenbauer)

(RHP). Oaxaca, d, 4 mi. S Tehauntepec, VII-18-

1952 (E. E. Gilbert and C. D. MacNeil) (UCAL);

d, 14 mi. NW Tehauntepec, VI-26-1961, 700 feet

(U. Kans. Exped.) (KANS); d, Tequistlan, IV-S1953 (R. C. Bechtel and E. I. Schlinger) (UCAL).

Sinaloa, d, 40 mi. S Culiacan, VII-22-1954 (M.

Cazier, W. Gertsch and Bradts) (AMNH). Sonora,

d, 10 mi. E Navajoa, VI11-13-1959 (W. L. Nutting

and F. G. Werner) (ARIZ) ; 9 , San Bernardo, VIII25-1935 (CAS).

Biology.—No reared specimen of s. habrosus has

been seen and the literature contains no records of the

biology.

Discussion.—Anthrax simson habrosus is similar in

all respects to s. simson, except the former usually has

a complete sectoral crossvein. In view of their occupying ecologically different habitats, however, and because their distributions do not overlap appreciably,

they are regarded here as distinct subspecies. They

probably were isolated by cold uplands in southeastern Mexico during the last glacial period, s. simson

being restricted to the Gulf coast of Mexico and Central America and s. habrosus occurring in warmer

Pacific side areas in south-central and western Mexico.

The subspecies A. s. habrosus may be readily distinguished from both delila and xylocopae by the complete sectoral crossvein on the wing. In the few specimens of s. habrosus with the sectoral crossvein

incomplete, the spur at the base of cell R4 is usually

at least twice as long as the r-m crossvein, whereas it is

rarely more than 2.0 times longer in delila and xylocopae. Also, the veins of the wing are light brown, contrasting with the pigment, in s. habrosus, whereas they

are the same color as the pigment or darker in

xylocopae and delila.

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126 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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NUMBER 43

127

FIGURES 1-6.—Hypothetical origins, points of isolation, and routes of dispersal of the taxa of

the Anthrax cephus group in North and South America: 1, ancestral form of cephus group;

2, analis complex; 3, luctuosus complex; 4, argyropygus complex, I; 5, cephus complex;

6, argyropygus complex, II.

36»-873 O—70 9
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Hypothetical evolutionary scheme for the tigrinus group of the genus Anthrax,

showing postulated point of origin, patterns of dispersal, and locations of isolating barriers.
NUMBER 4 3 129

FIGURES 8-27.—Lateral view of male genitalia of Anthrax (inset shows distal segment of

gonostylus in ventromedial view): 8, cintalpa; 9, irroratus irroratus (9a, anteroventral view of

epiphallus); 10, peruvianus; 11, inordinatus; 12, cordillerensis; 13, oedipus oedipus. Dorsal

view of right antenna of Anthrax: 14, cintalpa; 15, irroratus irroratus; 16, oedipus oedipus.

Representative scales found in Anthrax (not drawn to scale): 17, ovate; 18, ovate-truncate;

19, obovate-truncate; 20, linear; 21, lanceolate; 22, lanceolate truncate; 23, oblanceolate; 24,

oblanceolate truncate; 25-26, curly. 27, Representative hair from a tuft of pile. Abbreviations:

aae, apex of aedeagus; age, apex of gonocoxite; ae, apex of epiphallus; bae, bulb of aedeagus;

bap, basal apodeme of aedeagus; bgs, basal segment of gonostylus; cer, cercus; dgs, distal segment

of gonostylus; drb, dorsal band; lap, lateral apodeme of aedeagus; epm, epandrium.
130

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

37

FIGURES 28-37.—Lateral view of male genitalia of Anthrax: 28, seriepunctatus; 29, cybele; 30,

stellans; 31, atriplex; 32, melanopogon; 33, insulanus; 34, pluto pluto; 35, pluto nigriventris.

Anterior view of middle legs of Anthrax: 36, cintalpa; 37, oedipus oedipus.
NUMBER 4 3 131

49

FIGURES 38-49.—Anterodorsal view of female genitalia of Anthrax: 38, irroratus irroratus; 39,

irroratus striatipennis; 40, cybele; 41, pluto pluto; 42, seriepunctatus; 43, cordillerensis; 44,

insulanus; 45, atriplex; 46, peruvianus; 47, cintalpa; 48, inordinatus; 49, oedipus oedipus.

Abbreviations: fcrr, bristles; spb, bulb of spermatheca; j/>rf, duct of spermatheca; ssc, sclerite on

each side of spermathecal duct; 9 th T, ninth tergum.
132

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

55

FIGURES 50-61.—Male genitalia of Anthrax, lateral view (at right: above, dorsal view of the

apex of epiphallus and below, ventral view of apices of the gonocoxites and gonostyli): 50,

argentatus; 51, nitidus; 52, hylaios; 53, cathetodaithmos; 54, luctuosus; 55, koebelei; 56, aterrimus; 57, hyalacrus; 58, midas; 59, cephus; 60, pluricellus; 61, snowi. Abbreviations: aae, apex

of aedeagus; age, apex of gonocoxite; ags, apical segment of gonostylus; bae, base of aedeagus;

bap, basal apodeme of aedeagus; bgs, basal segment of gonostylus; dae, dorsal part of apex of

epiphallus; drb, dorsal band; lap, lateral apodeme of aedeagus; vae, ventrolateral part of apex

of epiphallus; vrb, ventral band.
NUMBER 43

133

FIGURES 62—74.—Male genitalia of Anthrax. (At right: above, dorsal view of apex of epiphallus

and below, ventral view of apices of gonocoxites and gonostyli): 62, clinopictus; 63, gideon; 64,

analis; 65, repertus; 66, argyropygus argyropygus; 67, argyropygus albosparsus; 68, argyropygus

painteri; 69, angustipennis; 70, macquarti; 71, austrinus; 72, laticellus; 73, innubilipennis; 74,

delicatulus.
134

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

10 T

FIGURES 75-94.—Anthrax, dorsointcrior view of female genitalia: 75, aterrimus; 76, midas; 77,

cephus; 78, hyalacrus; 79, hylaios; 80, cathetodaithmos; 81, luctuosus; 82, costaricensis; 83,

pluricellus; 84, argentatus; 85, angustipennis; 86, argyropygus argyropygus; 87, argyropygus

albosparsus; 88, argyropygus painteri; 89, macquarti; 90, repertus; 91, clinopictus; 92, funebris;

93, analis; 94, gideon. Abbreviations: jg, sclerite on each side of gonopore; J^fc, bulbs of spermathecae; j£;, spines on tenth tergum; 97", ninth tergum; 707", tenth tergum.
NUMBER 43 135

120

FIGURES 95-120.—Anthrax, dorsal (left) and mesal (right) views of antennae: 95, midas; 96,

aterrimus; 97, cephus; 98, hyalacrus; 99, luctuosus; 100, inaquosum; 101, nitidus; 102, argentatus;

103, hylaios; 104, snowi; 105, pluricellus; 106, costaricensis; 107, angustipennis; 108, delicatulus;

109, innubilipennis; 110, argyropygus argyropygus; 111, argyropygus albosparsus; 112, argyropygus painteri; 113, macquarti; 114, austrinus; 115, funebris; 116, analis; 117, gideonj 118,

clinopictus; 119, repertus; 120, laticellus.

369-673 O—70 10
136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 121-127.—Anthrax, female genitalia: 121, simson habrosus; 122, caatingensis; 123,

bellulus; 124, latibasis; 125, trimaculatus; 126, mystaceus. Anthrax simson habrosus; 127, male

genitalia. Abbreviations: sg, sclerite on each side on gonopore; spb, spermathecal bulb; sps, spines

on tenth tergum; spt, spermathecal tubes; 9T3 ninth tergum; 10T, tenth tergum.
NUMBER 4 3 137

epm

FIGURES 128-135.—Anthrax, male genitalia: 128 latibasis; 129, minimaculatus; 130, squalidus;

131, baliopteros; 132, trimaculatus; 133, bellulus; 134, mystaceus; 135, plurinotus. Abbreviations:

aae, apex of aedeagus; age, apex of gonocoxite; ags, apex of gonostylus; bae, base of aedeagus;

bap, basal apodeme of aedeagus; bgs, basal segment of gonostylus; cer, cercus; doe, dorsal part of

apex of epiphallus; drb, dorsal band; epm, epandrium; lap, lateral apodeme of aedeagus; vae,

ventrolateral part of apex of epiphallus; vrb, ventral band.

PLATES

369-673 O—70 11


142

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 2.—Wings of Anthrax: a, stellans, 3 ', h, stellans, $ ; c, seriepunctatus, $ ; d, seriepunctatus, $ ; e, pluto pluto, $ ; f, pluto pluto, $ ; g, pluto nigriventris, $ ; h, pluto nigriventris, $ ;

i, insulanus, $ ; j , insulanus, $ ; k, atriplex, $ ; I, atriplex, 9 ; m, melanopogon, $ ; n, melanopogon, $ ; o, cybele; p, tigrinus; q, varicolor varicolor; r, luctuosus; s, trimaculatus.
NUMBER 4 3 143

PLATE 3.—Wings of Anthrax; a, analis, typical form, 9 ; b, analis, form grossbecki, $ ; c, analis,

form grossbecki, 9 ; d, analis, form cedens, $ ; e, analis, form cedens, 9 ; f, argentatus, $ ; g,

xanthomeros, 9 ; h, argyropygus albosparsus, $ ; i, argyropygus albosparsus, 9 ; j, nitidus, 9 ; k,

inaquosum, 9 •
144

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

g

PLATE 4.—Wings of Anthrax: a, snowi, $ ; b, costaricensis, $ ; c, cephus, $ ; d, aterrimus, $ ;

e, midas, $ ; /, hyalacrus, $ ; g, hylaios, 9 ; h, clinopictus, $ ; i, gideon, $ ; ;, repertus, S ; k,

funebris, $ ; /, luctuosus, 2 .
NUMBER 4 3 145

9

PLATE 5.—Wings of Anthrax: a, delicatulus, S ; b, delicatulus, $ ; c, innubilipennis, $ ; d,

angustipennis, $ ; e, angustipennis, 5 ; f, laticellus, S ; g, argyropygus painteri, $ ; h, argyropygus

painteri, 9 ; i, argyropygus argyropygus, $ ; j , argyropygus argyropygus, $ ; k, macquarti, 9 ;

/, austrinus, $.
g

PLATE 6.—-Wings of Anthrax: a, plurinolus; b, baliopteros; c, trimaculatus; d, caatingensis; e,

minimaculatus; f, latibasis; g, bellulus; h, squalidus; i, mystaceus; j , delila; k, simson simson; I,

simson habfosus; m, xylocopae; n, tigrinus.
Index

Anthrax Species

(Synonyms and homonyms are in italics, pages with principal accounts are in boldface.)

acroleucus, 3, 85, 87

aequa, 3, 16, 17

analis, 2, 3, 4, 5, 37, 38, 40, 44, 46, 84, 87, 88, 89, 90, 91, 92,

133, 134, 135, 140

analis, form cedens, 3, 44, 89, 91, 92, 140

analis, form grossbecki, 3, 43, 44,61, 89,91, 92, 93, 140

angustipennis, 3, 40, 41, 42, 65, 66, 67, 68, 77, 80, 133, 134,

135,142

argentatus, 3, 4, 38, 39, 40, 43, 45, 46, 47, 91, 132, 134, 135,

140

argyropygus albosparsus, 3, 40, 42, 70, 71, 72, 73, 74, 133,

134, 135,140

argyropygus argyropygus, 3, 37, 40, 42, 68, 69, 70, 71, 72, 74,

96,133, 134,135,142

argyropygus painteri, 40, 41, 42, 70, 71, 72, 133, 134,135, 142

aterrimus, 3, 39, 42, 56, 59, 60,61, 93, 132, 134, 135, 138, 141

atriplex, 9, 10, 26, 31, 32, 33, 34, 130, 131, 139

austrinus, 40, 41, 42, 68,80, 81,84, 133, 135,142

baliopteros, 95, 96, 97, 105, 106, 107, 137, 143

bellulus, 3, 96, 97, 99, 102, 103, 104, 105, 110, 136, 137, 143

binotatus 3, 65,68

caatingensis, 95, 97, 106,108, 136, 143

calopterus,3,16,17

cathetodaithmos, 37, 39,43,49,50, 132, 134

cephus, 2, 39, 42, 56, 57,58, 59, 60, 89, 91, 93, 132,134, 135,

141

cintalpa, 3, 9,13,14,15,16,129,130,131,138

clinopictus, 40,44, 80, 84, 85, 133,134, 135, 141

contiguus, 3,68, 70

cordillerensis, 7, 8, 21, 22, 23, 129, 131, 138

costaricensis, 38,40,41, 62,63,64, 134, 135, 141

crinitus,3, 109, 110

cybele, 3, 4, 7, 9, 34, 35,130, 131, 139

delicatulus, 3, 37,39,41, 42, 76, 77, 133,135, 142

delila, 3, 5, 111, 113, 114, 115, 116, 118, 119, 120, 122, 123,

143

duodecimpunctatus, 3, 107,108, 110

funebris, 3,5,40,44,88,90,91,92, 134,135, 141

georgicus, 3, 88,89

gideon, 2, 3, 4, 37, 40, 44, 80, 84, 85, 86, 87, 88, 89, 91,

133, 134, 135, 141

gideon, form propinquus, 3,44,87, 88

guianicus, 3, 53,54

hyalacrus, 3, 43, 55, 56, 132, 134, 135,141

hylaios, 39, 43, 50,51, 52, 53, 132, 134, 135, 141

imitans, 3,98, 99

inappendiculatus, 3,102,103

inaquosum, 5, 40,43,93, 94,107, 135, 140

inexactus, 3, 81, 82

innubilipennis, 37,40, 41, 74, 75, 76, 77,133,135,142

inordinatus, 3, 7,8, 22, 23, 129, 131,138

insulanus, 9, 26, 27, 28,130,131, 139

irroratus.3, 10,11

irroratus irroratus, 2, 3, 7, 8, 10, 11, 12, 13, 16, 35, 129, 131,

138

irroratus striatipennis, 8, 11, 12,13,14, 131, 138

koebelei.40,43,91,92,132

latelimbatus, 3, 89, 91

latibasis, 95, 96, 97,99, 100, 101, 102, 104, 105, 136, 137, 143

laticellus, 40,41, 64,65, 133, 135, 142

leucopygus, 3, 54, 77, 78

leucopygus, 3, 54, 77, 78

limatulus fur, 4

luctuosus, 3, 37, 39, 43,47,48,49, 50,132,134, 135,141

macquarti, 3, 40, 41, 42, 68, 77, 78, 79, 80, 82, 84, 133, 134,

135,142

melanopogon, 3, 10, 32, 33, 130, 139

mexicanus, 3, 53,54

midas, 2, 3, 5,43, 53,54, 55,56, 78,132,134,135,141

minimaculatus, 3, 96,97,104,105, 129,143

mystaceus, 3, 96,97,102,107,108, 109,136,137,143

nitidus, 4, 38, 40, 43,46,47,132, 135,140

occidentalis, 3,88,89

oedipus aquilus, 9,17,18,19, 138

oedipus oedipus, 2, 7, 9, 16, 17, 18, 19, 21, 23, 129, 130,

131, 138

pauper, 94

peruvianus, 7, 8, 9, 16, 18,19, 20, 21, 23, 129,131,138

pluricellus, 3, 37,40,41,61,62, 63,64, 132,134

plurinotus, 3, 35, 94,96,97,110, 111, 137, 143

pluto nigriventris, 10, 24, 25, 26, 130, 139

pluto pluto, 3, 10, 23, 24, 25, 26, 27, 29, 32, 34, 35, 130, 131,

139

poecilophora, 3, 22, 23

punctatus, 3,16, 17

punctum, 3,16, 17

repertus, 3, 5, 37, 38, 40, 44, 80, 81, 82, 83, 84, 85, 133, 134,

135, 141

scriptus,3, 111, 118,119

seriepunctatus, 3, 9, 29, 30, 31, 33,34, 130, 131, 139

147
148 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

simson habrosus, 5, 111, 113, 114, 115, 116, 117, 118, 120,

121, 122, 123, 136, 143

simson simson, 2, 5, 111, 113, 114, 115, 118, 119, 120, 121,

122, 123, 143

slossonae, 3, 59, 60

snowi, 40, 41,62, 63, 64, 132, 135, 141

squalidus, 3,96,97,109,110, 137, 143

stellans, 3, 9, 24,28,29, 34,130,139

tigrinus, 2, 3, 111, 113, 114, 115,118,119,120, 139, 143

trimaculatus, 3,5, 94, 95, 96, 97,98, 99, 136, 137, 139, 143

varicolor varicolor, 139

vicinus, 3, 22

vicinus, 3, 22

xanthomeros, 40, 43, 52, 53, 140

xylocopae, 5, 111, 113, 114, 115, 116, 117, 118, 119, 120,

121, 122, 123, 143

Insect Hosts

Ancistrocerus catskill catskill, 70

spinolae, 60

Anthophora species, 11

Ashmediella bucconis denticulata, 11

Chalybion zimmermanni, 62

Cicindela scutellaria, var. lecontei, 89

Diadasia species, 99

Dianthidium heterulkei fraternum, 11

Dianthidium species, 16

Euodynerus megaera, 60

Hylaeus asininus, 11

Isodontia auripes, 60

Megachile chilopsidis, 16

concinna, 16

gentilis, 11,33

nivalis, 11

rotundata, 11

Melitoma euglossoides, 16

Monobia quadridens, 60

Odynerus guadelupensis, 18

Odynerus species, 11

Osmia clarescens, 11

rostrata, 11

Pachodynerus erynnis, 70

Pseudomasaris coquilletti, 11

Sceliphron species, 11

Stenodynerus fulvipcs fulvipcs, 70

pulvinatus surrufus, 70

saecularis rufulus, 70

Trypargilum clavatum, 60

collinum collinum, 70

collinum rubrocinctum, 60, 70

striatum, 60, 70

tridentatum archboldi, 70

Trypoxylon politum, 60

Xylocopa augusti, 5,120

brasiliorum varipuncta, 5

californica arizonensis, 118

tabaniformis orpifex, 116

virginica, 119

U.S. GOVERNMENT PRINTING OFFICE: 1970 O—360-673


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