NORMAN MARST Taxonomic Study of

the Known Pupae

of the Genus Anthrax

(Diptera: Bombyliidae)

in North and South A

America

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 100

Norman Marston TaXOIlOmic Stud y of

the Known Pupae

of the Genus Anthrax

(Diptera: Bombyliidae)

in North and South

America/

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Norman Marston. Taxonomic Study of the Known Pupae of the Genus Anthrax

(Diptera: Bombyliidae) in North and South America. Smithsonian Contributions to

Zoology, number 100, 18 pages, 1971.—The pupae of fourteen species in the cephus,

oedipus, and tigrinus groups of the genus Anthrax are described, and a key is given to

separate these and four other species in the albofasciatus and trimaculatus groups.

Photographs illustrate the cephalic and anal tubercles of sixteen species.

Contribution No. 896, Department of Entomology, Kansas Agricultural Experiment

Station, Kansas State University, Manhattan.

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 : 1971

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

Washington, D.C. 20402 - Price 35 cento (paper cover)
Norman Marston Taxonomic Study of

the Known Pupae

of the Genus Anthrax

(Diptera: Bombyliidae)

in North and South

America

Introduction

For several years I have been studying the taxonomy

of adults of the genus Anthrax Scopoli. During this

time a number of reared specimens associated with the

pupal skins have been assembled. Since very little information is available in the literature on the taxonomy

of Bombyliid pupae, descriptions of the available pupae

and a key for their separation are here published.

I am grateful to the various institutions and individuals listed in my previous papers (1962, 1970) for

the loan of specimens for study, and to Dr. Carl W.

Rettenmeyer for photographic help. The work was

supported in part by National Science Foundation

Grant G21291 to Dr. R. H. Painter and by Kansas

Agricultural Experiment Station Project 546.

A previous study (Marston, 1964) dealing with the

biology of the Anthrax species of the albofasciatus and

trimaculatus groups included descriptions and keys to

the pupae of four species. Otherwise, the only key to

pupae of Anthrax is that of Malloch (1917) which

separates four species (under Spogostylum Macquart

and Argyramoeba Schiner) along with species in other

genera. Several species have been described incomNorman Marston, Biological Control of Insects Research

Laboratory, Entomology Research Division, Agricultural Research Service, United States Department of Agriculture,

Post Office Box A, Columbia, Missouri 65201

pletely (Townsend, 1893; Malloch, 1915; Malloch,

1917; Brooks, 1952) due to lack of closely related

species for comparison.

In this study descriptions are given for fourteen

species in the cephus, oedipus, and tigrinus groups,

together with a key to these and the four species of

the albofasciatus and trimaculatus groups described

previously. Photographs illustrate the cephalic and

anal tubercles of sixteen of the species.

Taxonomic Characters

Pupae of species of Anthrax possess many excellent

diagnostic characters. Particularly, consistent variation

between each species has been noted in the shapes of

cephalic and anal tubercles (see plates). Interspecific

variations involve primarily forms of tubercles and

their spatial relationships. No attempt was made to

compare relative lengths of tubercles, as specimens

emerging through hard clay or similar materials have

abraded tubercles. The anterior and posterior facial

tubercles (on the long, narrow sclerite on the ventral

surface of the head) also vary between species.

The cephalic tubercles are peculiarly modified in the

two species representing the tigrinus group. The two

sets of tubercles are broadly separated mesally, rather

than united on a common, raised cephalic area as in

other groups. Also, the tubercles are broad basally and

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Table 1.—Spines and Setae on Pupal Skins of Anthrax (Number of Skins Examined

Shown in Parentheses).

Segment

Tergal spines

Tergal spines

Tergal setae

Pleural setae

Sternal setae

Tergal spines

Tergal setae

Pleural setae

Sternal setae

Tergal spines

Tergal setae

Pleural setae

Sternal setae

Tergal spines

Tergal setae

Pleural setae

Sternal setae

Tergal spines

Tergal setae

Pleural setae

Sternal setae

Tergal spines

Tergal setae

Pleural setae

Sternal setae

Tergal spines

Tergal setae

Pleural setae

Sternal setae

I

Anthrax

-

25-29

13-17

Anthrax

-

27-30

13-14

-

Anthrax

-

22-28

15-16

-

Anthrax

-

15-31

12-25

-

II III IV

tigrinus (DeGeer)

16-21

17-20

1

12-18

17-18

1

13-16

17-18

1

xylocopae Marston

14

26-27

4-5

3-4

11-14

27

3-5

3-4

pluricellus

12-16

11-12

16-22

6-9

11-14

12-16

17-20

10-12

11-12

24-25

4

2-4

V

(3)

10-14

14-18

1-2

(2)

11

21-23

3-5

3-4

VI

10-12

14-16

1-3

11-13

21

4-5

4-5

Williston (11)

9-13

12-15

15-20

8-10

aterrimus (Bigot)

14-18

10-17

12-30

7-20

14-20

11-21

10-33

10-26

Anthrax argyropygus

-

11-20

6-16

-

12-16

10-11

7-16

5-10

13-14

9-12

7-17

7-12

12-17

11-15

10-34

6-19

8-11

10-12

11-20

6-11

(26)

10-13

11-19

10-24

6-16

8-11

9-13

12-17

7-10

9-11

9-17

11-24

6-12

argyropygus Wiedemann

11-13

8-12

7-15

4-8

Anthrax hyalacrus Wiedemann

-

11

5-10

-

12-13

9-14

15

6-7

10-12

10-11

7-9

8

Anthrax analis Say

-

8

3

-

11

5-6

3

5

11

6

3

4-5

10

10-13

12-15

12

(1)

11

7

3

4

8-11

8-10

6-14

3-10

(2)

8

10-12

15-16

8-10

6

6

3

3

6-11

8-11

6-13

4-8

6-8

8-10

11-15

7-9

4

6

3

3

VII

5-7

14

5

2-3

7-8

17-18

4

4-5

10-11

10-14

13-16

8-12

7-10

8-12

7-20

3-7

(27)

8-10

8-11

5-15

4-10

7-8

7-9

10-11

6-8

1

5-6

3

3

VIII

-

1

-

-

1

-

4-9

-

2-4

2-5

3-4

-

2-7

2-3

4

-

1-6

3

3-4

1

3-4

2-3

1-2

-

1-2

-

I

Anthrax

-

10-15

5-6

Anthrax

-

9-15

7-13

-

Anthrax

-

13-15

10-13

-

Anthrax

-

14-15

10-12

-

Anthrax

-

9-10

10-11

-

Anthrax

-

15

16

-

II III IV

koebelei Marston

10-11

7

4-5

3-5

9-11

6-7

5-7

3-5

8

7

5-7

2-3

irroratus irroratus

11-13

9-14

8-16

3-4

11-12

9-13

8-16

3-7

10-12

9-14

7-13

3-8

cintalpa Cole (4)

15-16

13-16

16-24

4-7

13-14

12-18

13-22

6-8

13-14

11-21

IS-24

6-9

peruvianus Marston

13

10

15-18

4-7

13

11-12

16-17

7

12-14

11-12

16-20

6

V

[4)

6-8

6-7

5-6

3

Say

9-10

8-14

6-13

2-7

9-10

11-21

14-31

5-8

(3)

10-12

10-11

16

6-7

VI

6

6

5-8

3

(6)

-

10-13

7-14

2-8

-

14-26

14-27

5-8

-

14

15

7

cordillerensis Marston (1)

15

10-12

15-19

7

15

8-12

16

7-8

melanopogon

15

11

18

9

15

14

19

10

14

9-10

12-15

4-6

(Bigot)

11

13

15

9

Anthrax atriplex Marston

-

20-24

25-27

-

16-17

13

24-27

9-15

14-15

11-14

25-27

12-14

13-15

12-15

23-24

16-17

12

8-9

13-14

6

(1)

7

12

14

10

(2)

11-12

13-17

19-23

11-13

8

9-10

12-14

5-7

-

13

11

9

-

15-17

21

10-11

VII

6-7

5-6

4-8

3-4

-

7-11

5-12

3-7

-

11-31

13-27

6-12

-

11

12-13

6

-

10

11

6

-

10

12

6

-

12-16

16-17

8-10

VIII

2

3

2-3

-

0-4

3-7

-

-

6-15

4-13

10

-

4

4-6

1-3

-

4

3

1

-

2

6

-

-

5-7

4-5

3
NUMBER 10 0

broadly rounded apically, except for the anterior pair

in A. xylocopae Marston. The anal tubercles in the

tigrinus group are quite simple, represented by broadly

separated, spurlike structures, whereas in species in

other groups the tubercles show various combinations

of denticles and do not taper evenly to the apices.

Several excellent characters have been noted on the

posterior abdominal terga. Species of the cephus group

and A. trimaculatus (Macquart) have several small,

simple spines between setae on the sixth and seventh

terga. Most species of the albofasciatus and oedipus

groups have no such spines, although some specimens

of A. limatulus Say have a few, especially on the sixth

tergum. (Tables in my 1964 paper on the albofasciatus

and trimaculatus groups omitted these spines and those

on the eighth tergum). On the eighth tergum of A.

trimaculatus and on all species of the cephus group

except A. analis Say and A. koebelei Marston, are two

pairs of triangular, spinelike processes, absent in other

groups of the genus, which occur on each side of the

dorsolateral tumidities (rounded tumorlike irregularities in the integument). In koebelei the mesal pair of

processes is absent, while in analis both are absent.

Members of the oedipus and albofasciatus groups

have a transverse row of setae on each side of the

eighth tergum in front of the dorsolateral tumidities.

In the species representing the cephus group, setae

are present on the tergum only in A. hyalacrus Wiedemann, and are on a line between the dorsolateral

tumidities.

The numbers of sections (membranous parts) of the

spiracles vary widely, but with considerable intraspecific variation. The flange around the spiracles varies

from broad and well developed to entirely absent. In

general, species of the cephus group have the least

developed flanges, while species of the albofasciatus

and oedipus groups have the broadest. In the oedipus

group, the two species representing the pluto subgroup,

A. melanopogon and A. atriplex, have the flanges

broadly lobed posterodorsally on the anterior spiracles,

while species of the oedipus subgroup have them evenly

rounded.

Numbers of abdominal setae and spines differ greatly

among species, as can be seen in Table 1, but there is

considerable intraspecific variation. In general, smaller

specimens have fewer setae and spines than larger

specimens. The ranges of setae and tergal spines are

tabulated. The number of tergal spines in the table is

the total on the segment, but the number of setae is

given for one side only. Only males have setae on the

eighth sternum.

Key to Known Pupae of Anthrax in North and South America

1. Anterior cephalic tubercles broadly separated mesally (Plate 2H) ; anal tubercles,

acuminate, without denticles (Plate 3D) (tigrinus group) 2

Anterior cephalic tubercles broadly connected mesally (Plate 2i); anal tubercles

with at least mesal, preapical denticles 3

2(1). Anterior cephalic tubercles broad, dorsoventrally flattened, broadly rounded distally

(Plate 2H) • A. tigrinus (DeGeer)

Anterior cephalic tubercles more or less terete, acuminate apically (Plate 2G).

A. xylocopae Marston

3(1). Seventh abdominal tergum with numerous small, simple spines between setae; eighth

with one or two sharp processes on each side (except in analis); setae absent or,

between dorsolateral tumidities (cephus and trimaculatus groups) 4

Seventh abdominal tergum without spines; eighth without sharp processes, with

setae in transverse line in front of dorsolateral tumidities (albofasciatus and

oedipus groups) 10

4(3). Dorsal and ventral denticles at base of each anal tubercle, when present, not connected

by sharp lateral ridge (Plate 3G), occasionally indistinctly joined by low ridge;

mesal process on eighth tergum, when present, usually tapering to sharp apex

(cephus group) 5

Dorsal and ventral denticles at base of each anal tubercle connected by sharp lateral

ridge (Plate 3E ); mesal process on eighth tergum bifid apically (trimaculatus

group) A. trimaculatus Macquart

5(4). Anal tubercles without denticles basally, long, narrow, and broadly separated mesally

(Plate 3A,B) 6

Anal tubercles with more or less distinct denticles basally, separated by less than width

of one tubercle mesally 7
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Known Pupae of Anthrax in North and South America—Continued

6(5). Eighth tergum with one spine on each side lateral to dorsolateral tumidities; anterior

facial tubercles sharply rounded apically A. koebelei Marston

Eighth tergum without spines; anterior facial tubercles long and acuminate.

A. analis Say

7(5). Eighth tergum without setae between innermost processes; dorsal, basal denticles on

each anal tubercle arising at basal fourth or fifth (Plate 3F,O) 8

Eighth tergum with a seta on each side mesad to innermost process; dorsal, basal

denticles on each anal tubercle arising at basal third or half. (Plate 3H) .

A. hyalacrus Wiedemann

8(7). Each anal tubercle with sharp denticle ventrally on mesal margin about one-third

of distance to apex; medial facial tubercle formed on a line between anterior and

posterior facial tubercles or slightly outside it (Plate 2F) 9

Each anal tubercle with poorly denned, blunt-tipped ridge running about one-third

of way to apex; medial facial tubercle formed slightly inside a line between anterior

and posterior facial tubercles (Plate 2D) . . A. argyropygus argyropygus Wiedemann

9(8). Each anal tubercle about six times longer than medial width (Plate 3F).

A. pluricellus Williston

Each anal tubercle about five times longer than medial width or shorter.

A. aterrimus (Bigot)

10(3). Anal tubercles with more or less distinct mesal denticles about two-thirds of distance

to apices (Plate 4F) 11

Anal tubercles without sharp denticles mesally, each with sharp ridge curving

ventromesally from tip and then recurving dorsally to meet oblique ridge at sharp

point two-thirds of distance to apex (Plate 4E) A. nidicola Cole

11(10). Medial and posterior cephalic tubercles not widely separated from anterior pair,

united less than halfway to apices (Plate 1H) 12

Medial and posterior cephalic tubercles widely separated from anterior pair, united

about two-thirds of distance to apices, about equal in length (Plate 2c).

A. albofasciatus albofasciatus Macquart

12(11). Mesal process on ninth tergum small, broad and low, or sharply tipped apically 13

Mesal process on ninth tergum well developed, about as long as broad, bifid, truncate

or broadly rounded apically subspecies of A. limatulus Say

13(12). Flange on anterior abdominal spiracles lobed posterodorsally 14

Flange on anterior abdominal spiracles evenly rounded posterodorsally 15

14(13). Posteromesal ridge on each medial cephalic tubercle extending to posterior margin

of cephalic area A. atriplex Marston

Posteromesal ridge on each medial cephalic tubercle becoming indistinct near base

of tubercle A. melanopogon (Bigot)

15(13). Dorsal and ventral denticles at base of each anal tubercle connected by low ridge

or unconnected (Plate 4C,D) ; ventrobasal denticles not connected by flanges to

ventral denticles on mesal margins (when present) 16

Dorsal and ventral denticles at base of each anal tubercle connected by high flange

(Plate 4B); ventrobasal denticle connected by sharp flange to ventral denticle

on mesal margin A. peruvianus Marston

16(15). Sixth abdominal tergum with several spines between setae. A. cordillerensis Marston

Sixth abdominal tergum without spines between setae A. irroratus irroratus Say

and A. cintalpa Cole
NUMBER 100

Anthrax tigrinus (DeGeer)

Hurd (1959) has reviewed available biological information on A. tigrinus, a common parasite of Xylocopa

virginica virginica (Linnaeus) in the eastern United

States. Three pupal skins have been examined.

Integument of pupa brownish hyaline with yellow

setae and reddish brown, black tipped tubercles and

spines. Cephalic tubercles (Plate 2H ) formed in

straight line, anterior pair not joined mesally; anterior

tubercle broad, flattened dorsoventrally, broadly

rounded apically, little longer than medial tubercle,

with sharp ventromesal and lateral edges; medial

tubercle flattened dorsoventrally, broadly rounded

apically, with sharp mesal and lateral edges, and ventral and ventrolateral ridges; posterior tubercle somewhat flattened dorsoventrally, with sharp mesoventral

and dorsolateral edges, and anteromesal and posterior

ridges. Length of facial sclerite about one and a half

times width, posterior edge truncate; anterior facial

tubercles simple, acuminate to blunt apices,

broadly separated mesally; posterior facial tubercle

undeveloped.

Thoracic spiracle with about twenty-seven sections,

posterior ones undeveloped; flange undeveloped.

Abdominal spiracles with about twelve to seventeen

sections, anteroventral ones reduced; flange poorly

defined, forming thick lobe anteroventrally. Setae on

abdomen broad, flat, and finely pubescent.

Eighth tergum without setae or spines. Ninth tergum

without mesal process. Anal tubercles (Plate 3D)

without denticles, rounded basally, tapering to acute

apices.

The three specimens examined do not vary significantly in basic structure. A. tigrinus may be readily

separated from xylocopae by the structure of the

anterior cephalic tubercles. In tigrinus they are broad,

dorsoventrally flattened, and broadly rounded apically, whereas in xylocopae they are narrow, more or

less terete, and acuminate apically.

Numerous characters separate tigrinus, xylocopae,

and probably other species in the tigrinus group, from

other groups of Anthrax. Probably the most readily

observable is the distinct separation mesally between

the anterior cephalic tubercles of tigrinus and

xylocopae. In other groups the bases of the tubercles

are broadly united mesally.

Anthrax xylocopae Marston

Two pupal skins of this species have been examined

(see table). Adults were reared from nests of

Xylocopa californica arizonensis Cresson in flower

stalks of Yucca elata Engelmann at Rodeo, New

Mexico. Hurd (1959) discussed the biology of the

species (as Anthrax simson Fabricius).

Integument of pupa brownish hyaline with yellow

setae and yellowish brown black-tipped tubercles. Bases

of anterior cephalic tubercles (Plate 2G) not joined

mesally; medial tubercle situated in nearly straight

line between apices of anterior and posterior tubercles;

anterior tubercle narrow, acuminate, curved posteromesally at apex, with high mesoventral and dorsolateral ridges; medial tubercle flattened dorsoventrally,

broadly rounded apically, with sharp dorsal, ventral,

and ventrolateral ridges; posterior tubercle compressed

on dorsal side, with sharp mesal, lateral, and ventral

ridges. Facial sclerite slightly less than twice its narrowest width, with broad, shallow emargination posteriorly; anterior facial tubercles simple, acuminate to

blunt apices, broadly separated mesally; posterior

facial tubercle undeveloped. Anterior facial setae arising slightly in front of level of anterior facial tubercles.

Thoracic spiracle with about fifteen sections, posterior ones reduced; indistinct flange present around

anterodorsal fourth. Abdominal spiracles with about

thirteen sections, anteroventral ones reduced, flange

represented by low ridge around posterodorsal half,

expanding as broad lobe anteroventrally.

Eighth tergum without setae or spines. Ninth tergum

without medial process. Anal tubercles (Plate 3c),

acuminate, without denticles; bases of tubercles

broadly separated, mesal margin tapering outward to

apices, then slightly curved inward at tips.

The two pupal skins examined do not vary significantly in diagnostic characters. Characters for separating A. xylocopae from tigrinus and other species in

the genus are discussed under tigrinus.

Anthrax pluricellus Williston

Seven pupal skins of A. pluricellus have been examined

(see table). These were taken by Rau from nests of

Chalybion zimmermanni Dahlbom. (Hymenoptera:

Sphecidae) (Rau, 1940). The specimens are in the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

collection of Dr. R. H. Painter and the National

Museum of Natural History.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles and

spines. Cephalic tubercles (Plate 2F) evenly spaced, in

a straight line; anterior tubercle of moderate length,

compressed dorsoventrally toward apex, with poorly

defined mesal and lateral ridges; medial tubercle nearly

as long as anterior one, slightly bent forward, acuminate, with indistinct dorsolateral and ventromesal

ridges, and sharp ventrolateral ridge connecting to

posterior tubercle; posterior tubercle short, slightly

curved anterodorsally, concave on anteromesal surface, with sharp anterolateral and posteromesal ridges

and indistinct posterolateral ridge. Facial sclerite about

eight times longer than minimum width, posterior end

with broad, semicircular emargination; anterior facial

tubercles flattened anteroposteriorly, bent forward

slightly, broad, short, bluntly rounded apically; posterior facial tubercle undeveloped, represented by

rounded node. Facial setae arising just anterior to level

of anterior facial tubercles.

Spiracles a little above level of integument, with

out distinct flanges; thoracic spiracle with about 12

sections, posteroventral ones reduced; abdominal spiracles with about ten sections, anteroventral ones

reduced.

Sixth and seventh terga with numerous, small spines

between setae. Eighth tergum with two spines on each

side, one on each side of dorsolateral tumidities, additional small spines sometimes present; setae absent.

Medial process on ninth tergum broad, low, sharply

rounded apically. Anal tubercles (Plate 3F) about

six or seven times longer than medial width, separated

medially by one-fourth or less of width of one lobe.

Each tubercle with dorsal and ventral, sharp denticles

near base unconnected or joined by low, poorly denned

ridge; mesal margin with sharp denticle ventrally

about one-third of distance to apex and large mesal

denticle about three-fourths of distance to apex; apex

narrow, straight, and acuminate beyond preapical

denticle.

The pupae of A. pluricellus, aterrimus, argyropygus

argyropygus, and hyalacrus are quite similar and probably are typical of most species of the cephus group

except for species closely related to analis. The pupa

of koebelei seems to be intermediate between analis

and the first four, sharing some common characters

with each, as does the adult. The pupa of koebelei,

and those of the first four, may be readily separated

from pupae of all other species studied, except trimaculatus, by the presence of spines on the eighth tergum.

A. trimaculatus may be readily distinguished by the

high flange connecting the dorsal and ventral denticles

on the anal tubercles.

In the cephus group, A. analis is unique in that it

lacks spines on the eighth tergum, while koebelei is

distinctive in that it has only one spine on each side,

lateral to the dorsolateral tumidities. Other species

have an additional pair mesad to the tumidities. In

pluricellus, the anal tubercles are distinctly longer than

in argyropygus, hyalacrus, and aterrimus, about six or

seven times the medial width of one lobe rather than

five times the medial width of one lobe or less. A.

hyalacrus may be distinguished from pluricellus,

argyropygus, and aterrimus by a pair of setae between

the innermost spines on the eighth tergum. The setae

are in a direct line between the dorsolateral tumidities

rather than in front of them as on pupae of other

groups. The cephalic tubercles of A. argyropygus are

distinctive in that the medial one is set slightly inside

a line between the tips of the anterior and posterior

ones, whereas in pluricellus, aterrimus, and hyalacrus

they are in a straight line or the middle one is set

slightly outside the line.

Anthrax aterrimus (Bigot)

Twenty-six pupal skins of this species from the National Museum of Natural History have been studied

(see table). According to Krombein (1967), they

were taken from nests of Trypargilum striatum (Provancher), T. clavatum (Say), T. collinum rubrocinctum (Packard), and Isodontia auripes (Fernald)

(Hymenoptera: Sphecidae) ; Euodynerus megaera

(Lepeletier), Monobia quadridens (Linnaeus), and

Ancistrocerus spinolae (Saussure) (Hymenoptera:

Vespidae). An additional specimen was reared from a

"mud dauber wasp."

Integument of pupa yellowish hyaline with yellow

setae and reddish brown, black-tipped tubercles and

spines. Anterior cephalic tubercle straight, tapering,

with dorsomesal and ventrolateral ridges distally, without well-defined ridges basally; medial tubercle formed

on or slightly outside a line between anterior and posterior tubercles, closer to posterior tubercle, slightly

shorter than anterior tubercle, acuminate, with sharp

ventrolateral ridge connecting to posterior tubercle;
NUMBER 100

posterior tubercle short, slightly curved dorsomesally

with sharp dorsal, ventromesal, and ventrolateral

ridges. Facial sclerite about five and a half times longer

than its narrowest width, with shallow, rounded emargination posteriorly. Anterior facial tubercles short,

broad, anteroposteriorly flattened, tapering slightly to

broadly rounded apices; posterior facial tubercle not

developed.

Sixth and seventh terga with small spines between

setae. Eighth tergum with two spines on each side, one

on each side of dorsolateral tumidity (lateral spines

sometimes bifurcate); setae absent on tergum (1 seta

present adjacent to lateral spine on one specimen).

Medial process on ninth tergum short, broad to narrow basally, acuminate to sharp or narrowly bifurcate

apex. Each anal tubercle three to five times longer than

medial width of one tubercle, straight, separated

mesally one-fourth to one-third of width of one tubercle; each tubercle with sharp dorsal and ventral, unconnected denticles near base; mesal margin with

broad, sharp, preapical ridge, forming dorsal and ventral denticles, and sharp ventral denticles about onethird of distance to apex; each tubercle tapering outward from preapical ridge to sharp tip.

The pupae of A. aterrimus and pluricellus are quite

similar. Differences are discussed under the latter

species.

Anthrax argyropygus argyropygus Wiedemann

Twenty-seven pupal skins of A. argyropygus have been

examined (see table). The species has been reared

from Trypargilum collinum collinum Smith, T. c.

rubrocinctum (Packard), T. striatum (Provancher),

and T. tridentatum archboldi (Krombein) (Hymenoptera: Sphecidae); and from 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) (Krombein, 1967). An additional specimen was

reared from the nest of a "trypoxylonid" (Hymenoptera: Sphecidae). Specimens are in the National

Museum of Natural History and the R. H. Painter

collection.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles and

spines. Anterior cephalic tubercle (Plate 2D) narrow,

straight, and without definite ridges; medial tubercle

closer to anterior one than to posterior one, formed

slightly inside line between them, narrow, straight, with

sharp posterolateral ridges connecting to posterior

tubercle; posterior tubercle short, slightly curved

anteriorly, with poorly defined anteromesal, anterior,

and posterolateral ridges. Facial sclerite about eight

times longer than narrowest width, with shallow,

rounded emargination posteriorly. Anterior facial

tubercles short, broad, anteroposteriorly • flattened,

produced slightly laterad and evenly rounded apically;

posterior facial tubercle very short, flat, bent posteriorly, acute apically.

Spiracles produced slightly above margin of integument; thoracic spiracle with about thirteen sections,

the posterior ones reduced; abdominal spiracles with

eight to ten sections, anteroventral ones reduced.

Sixth and seventh terga with numerous small spines

between setae. Eighth tergum with two spines on each

side, one on each side of each dorsolateral tumidity

(one lateral spine bifurcate from base); no setae

present or tergum. Medial process on ninth tergum

short, broad basally, acuminate to sharp apex. Anal

tubercles (Plate 3o) short, thick basally, straight,

separated mesally by about one-third to one-fourth

width of one lobe; each tubercle with sharp dorsal and

ventral, unconnected denticles near base; mesal

margin with sharp dorsal and weak ventral denticles

connected by a sharp ridge about three-fourths of distance to apex; apex broadly acuminate, slightly

curved mesad apically.

The pupa of A. argyropygus argyropygus is most

similar to pluricellus, aterrimus, and hyalacrus. Relationships of the four species are discussed under

pluricellus.

Anthrax hyalacrus Wiedemann

Two specimens of A. hyalacrus were reared by Dr. Carl

W. Rettenmeyer from nests of a trypoxylonid wasp

(Hymenoptera: Sphecidae) on Barro Colorado Island,

Canal Zone, Panama. The wasps were nesting in a

cardboard carton in a clearing at the biological

station.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles and

spines. Cephalic tubercles (Plate 2E) evenly spread,

almost in a straight line; anterior tubercle short, com414-282 O - 71 - 2
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pressed dorsoventrally, bluntly rounded apically, and

with poorly defined dorsomesal and ventrolateral

ridges toward apex; medial tubercle short, acuminate

to blunt apex, with well-defined dorsolateral, ventrolateral, and mesal ridges; posterior tubercle short,

curved slightly forward and bluntly rounded apically,

connected to medial tubercle by sharp dorsolateral

ridge, and with less distinct ventral and mesal ridges.

Facial sclerite about eight times longer than narrowest

width, with deep, square, narrow emargination at

posterior end. Anterior facial tubercles short, broadly

acuminate, flattened anteroposteriorly, narrowly separated or connected mesally. Posterior facial tubercle

undeveloped, represented by rounded node.

Spiracles produced little above surrounding integument, without flanges; thoracic spiracle with about

fifteen sections, posterior ones reduced; abdominal

spiracles with about twelve sections, anteroventral ones

reduced.

Sixth and seventh terga with numerous, small, single

spines between setae. Eighth tergum with two spines

on each side, one on each side of dorsolateral tumidities, one of dorsal pair sometimes absent; one seta also

present on each side between tumidities rather than

anterior to them. Medial process on ninth tergum

short, narrow, acuminate. Anal tubercles (Plate 3H )

short and thick, slightly curved upward apically, separated mesally by about two-fifths of medial width of

one lobe; each tubercle with short, spinelike, dorsal

and ventral denticles about one-third of way to apex,

ventral one sometimes reduced; ventromesal margin

with sharp ridge extending about two-thirds of distance to apex, without denticles; dorsomesal margin

with sharp denticle about three-fourths of distance to

apex; apex bluntly acuminate beyond preapical

denticle.

The pupa of A. hyalacrus is most similar to pluricellus, aterrimus, and argyropygus argyropygus. Relationships of the four species are discussed under

pluricellus.

Anthrax analis Say

Shelford (1913) gave an excellent account of the life

history and habits of A. analis, which he reared from

burrows of "Cicindela scutellaria Say, var. lecontei

Hald." (Coleoptera: Cicindellidae). Malloch (1915)

described the larva and pupa, utilizing specimens collected by Shelford. The specimen described here is

from the R. H. Painter collection (see table). It agrees

well with Malloch's description, although the latter is

rather incomplete, due to a lack of closely related

species for comparison.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles and

spines. Anterior cephalic tubercle (Plate 2A) broadly

separated from middle and posterior ones, long, narrow, acuminate, and with sharp lateral, mesal, and

ventral ridges; medial and posterior tubercles united

about halfway to apex of posterior tubercle; medial

tubercle formed well inside line between tips of anterior and posterior ones, narrow, acuminate, and with

sharp dorsal, mesal, and posterolateral ridges; posterior

tubercle slightly curved mesad, acuminate, sharp, with

sharp dorsal, ventral, and lateral ridges. Facial sclerite

about seven times longer than narrowest width, posterior end shallowly emarginate. Anterior facial tubercles

anteroposteriorly flattened, long, broad basally, tapering to sharp apices, meeting at acute angle mesally.

Posterior facial tubercles undeveloped, represented by

rounded node.

Spiracles produced distinctly above surrounding integument; thoracic spiracle with about seventeen sections, posterior ones reduced; abdominal spiracles with

about ten sections, anteroventral ones reduced. Flanges

undeveloped.

Sixth and seventh terga with small, single spines between setae. Eighth tergum without spines or setae.

Medial process on ninth tergum undeveloped, represented by low, acutely pointed node. Anal tubercles

(Plate 3A) long, narrow, curved upward, and separated by about one and a half times width of one tubercle mesally. Each tubercle without denticles at base or

on ventral side; dorsomesal side with poorly defined

denticle about one-half of distance to apex formed by

apical extension of thin, sharp dorsomesal ridge; apex

narrow, curved slightly upward and mesad, sharp, and

with poorly defined lateral, dorsomesal, and ventromesal ridges.

The pupa of A. analis is most similar to that of

koebelei. A. analis may be readily recognized, however,

by absence of spines on the eighth abdominal tergum

and by acuminate rather than rounded anterior facial

tubercles. These two species may be separated from

all other species studied, except xylocopae and tigrinus,

by the absence of denticles at the bases of the anal

tubercles, and from the latter by being much smaller
NUMBER 100

and having acuminate rather than rounded cephalic

tubercles.

Anthrax koebelei Marston

Three pupal skins and one dried pupa associated with

adults of this species are in the California Academy of

Sciences collection. No host data were recorded with

the specimens, however.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles. Anterior cephalic tubercle (Plate 2B) broadly separated

from posterior ones, long, narrow, acuminate, slightly

curved ventrad, and with indistinct lateral, mesal, and

ventral ridges. Medial and posterior tubercles united

basally about halfway to apices of posterior one; medial

tubercle formed slightly mesad of line between apices

of anterior and posterior ones, slightly curved anterodorsally, acuminate, and with sharp posterior and less

distinct lateral and mesal ridges; posterior tubercle

about two-thirds as long as medial one, curved anteromesally, flattened posterolaterally, and with distinct

anteromesal, anterolateral and posteromesal ridges.

Facial sclerite about six times longer than its narrowest width, shallowly emarginate posteriorly. Anterior

facial tubercles broad, flattened anteroposteriorly,

bluntly rounded apically, not connected by mesal ridge.

Posterior tubercle undeveloped, represented by broad,

low, transverse node.

Thoracic spiracle slightly produced above integument, without flange, composed of about fifteen sections, posterior ones reduced. Abdominal spiracles like

thoracic, but with ventral sections reduced.

Sixth and seventh terga with small, single spines between setae. Eighth tergum without setae dorsally,

with one spine on each side laterad of tumidities.

Medial process on ninth tergum acuminate. Anal tubercles (Plate 3B) long, narrow, broadly separated

mesally, and slightly curved upward; tubercles without denticles basally or on ventral surfaces at basal

third; strong dorsal and weak ventral denticles present

mesally two-thirds of distance to apices, connected by

sharp, serrate ridge; tips curving slightly outward and

recurving dorsomesally, narrow and acuminate.

The pupa of A. koebelei appears to be most closely

related to that of analis Say. They may be readily separated, however, by the absence of spines on the eighth

tergum in analis, and by the acuminate rather than

rounded apices of the anterior facial tubercles of analis.

Anthrax irroratus irroratus Say

Townsend (1893) first described the pupal skin of A.

irroratus irroratus (as Argyiamoeba 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" deep in adobe soil." Baker (1895)

recorded 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."

Collections studied by the author contained specimens reared from Megachile nivalis Friese, M. rotundata (Fabricius), Megachile sp., Osmia rostrata Sandhouse and O. clarescens Cockerell (Hymenoptera:

Megachilidae); Anthophora sp. (Hymenoptera: Apidae); Odynerus sp. and Pseudomasaris coquilletti

Rohwer (Hymenoptera: Vespidae); and from Sceliphron sp. (Hymenoptera: Sphecidae). The species

seems to be a nonspecific parasite whose hosts are

various hymenopterous insects nesting in logs, banks,

and open, flat areas. Females often are observed ovipositing in openings of tunnels in stumps and logs, and

sometimes are attracted to the collector, flipping eggs

at dark spots on the clothing.

Six pupal skins have been examined (see table).

They are in the collections of the author, the University of California at Davis and Riverside, and the

Canadian National Collection. The specimens vary

greatly in size, which accounts for wide variation in

numbers of setae and spines.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles.

Cephalic tubercles (Plate ID) formed almost in

straight line, medial tubercle situated slightly outside

line between apices of anterior and posterior tubercles

and closer to posterior tubercle. Anterior tubercle

straight, acuminate, with sharp ventromesal and dorsolateral ridges and less distinct dorsomesal and ventrolateral ones; medial tubercle curved slightly forward,

acuminate, and with sharp ventromesal and dorsolateral ridges, and sharp ventrolateral ridge connecting

to posterior tubercle; posterior tubercle short, curved

anterodorsally and tapering to acute apex, with sharp

dorsal, ventromesal, and ventrolateral ridges. Facial

sclerite about four and a half times longer than narrowest width, bluntly rounded or shallowly emarginate
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

posteriorly. Anterior facial tubercles compressed anteroposteriorly, connected by sharp mesal ridge, with

acute lateral apices. Posterior facial tubercle formed

of two small anteroposteriorly flattened, apically

rounded nodes. Anterior facial setae arising distinctly

anterior to anterior facial tubercles.

Thoracic spiracle with about seven sections, with

low, sharp flange around anterior two-thirds. Abdominal spiracles with about six sections, anteroventral ones

reduced, sharp flange around anterodorsal threefourths broad dorsally and especially anteroventrally,

narrow otherwise.

Sixth and seventh terga without spines between

setae. Eighth tergum without spines dorsally, with

setae arising along line in front of dorsolateral tumidities. Ninth tergum with short, simple transverse process, broad basally, tapering to bluntly rounded apex.

Anal tubercles (Plate 4c) about four times longer than

medial width of one lobe, separated mesally by about

one-third medial width of one lobe or almost touching.

Each tubercle basally with sharp dorsal and ventral

denticles sometimes connected by low ridge; ventral

denticle on mesal margin poorly defined or absent;

strong dorsal and weak ventral denticles present preapically on meson from which tubercle tapers outward

to sharp, slightly upturned apex.

The pupae of all available species of the oedipus

subgroup show few good diagnostic characters. Also,

in several species a wide intraspecific variation often

serves to mask valid specific differences, particularly

where pupae have been reared from unrelated hosts.

The pupa of A. peruvianus may be most readily

separated from those of irroratus irroratus, cordillerensis, and cintalpa by presence of high lateral flange

between the dorsal and ventral denticles on the anal

tubercles and by presence of a flange between the

ventral denticles at the base and the ventral denticles

on the mesal margins of the tubercles about halfway to

the apices. In irroratus irroratus, cordillerensis, and

cintalpa some specimens may show a low ridge between the dorsal and ventral denticles, but none has

a connection between the mesal ridge and the ventral

denticles. A. cordillerensis may be readily distinguished

from the other species by the presence of spines between the setae on the sixth abdominal tergum.

There are no constant taxonomic characters to

separate all specimens examined of A. i. irroratus and

cintalpa. All specimens of cintalpa have the medial

cephalic tubercle situated farther outside a line between the anterior and posterior tubercles, however. In

cintalpa, a line through the tips of the posterior and

medial tubercles would be nearly parallel to the meson,

whereas, in irroratus irroratus such a line would be at

a distinct angle from the meson. Several specimens of

the latter have fewer tergal setae on the first abdominal segment—only 9 or 10 compared with about 14 in

all specimens of cintalpa examined. Also, two specimens of cintalpa show a greater number of pleural

setae on the eighth abdominal segment—16 tol8 compared with the usual 3 to 7 on specimens of irroratus

irroratus.

Anthrax cintalpa Cole

Cole (1957) stated that the type specimens of cintalpa

were reared from nests of Melitoma euglossoides

Lepeletier & Serville (Hymenoptera: Apidae). One

paratype studied by the author has the pupal skin

pinned with it. Additional specimens with pupal skins

were reared from nests of Megachile concinna and

Dianthidium sp. (Hymenoptera: Megachilidae).

Specimens with pupal skins are in the collections of

R. H. Painter, the author, and the University of

California at Berkeley.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles. Medial

cephalic tubercle (Plate lc) formed outside a line

between tips of anterior and posterior tubercles; line

through tips of medial and posterior tubercles parallel

to or at slight angle to meson; anterior tubercle slender, tapering to sharp apex, with sharp mesoventral

and dorsolateral ridges and with indistinct dorsomesal

ridge; medial tubercle broad basally, tapering to sharp

apex, with sharp ventrolateral ridge; posterior tubercle

short, curved upward, and tapering to sharp apex,

with sharp dorsal and ventromesal ridges and blunt

ventrolateral ridge. Facial sclerite about five times

longer than narrowest width, with shallow emargination at posterior end; anterior facial tubercles anteroposteriorly flattened, with bluntly angled lateral apices,

joined by sharp ridge mesally; posterior facial tubercle

formed as low transverse ridge, rounded or indistinctly

binodal. Anterior facial setae arising distinctly anterior

to bases of anterior facial tubercles.

Thoracic spiracle with about seven sections, posterior ones reduced, with sharp flange around anterior

two-thirds. Abdominal spiracles with about six sections,

anteroventral ones reduced; flange arising posteriorly,
NUMBER 100 11

becoming broad dorsally, narrowing anteriorly and

broadening again ventrally; flange evenly rounded

posterodorsally.

Sixth and seventh terga without spines between

setae. Eighth tergum without spines dorsally, with setae

in front of line between dorsolateral tumidities. Medial

process on ninth tergum high and broadly rounded

apically or low and tapering to sharply rounded apex.

Each anal tubercle (Plate 4H ) four to five times longer

than medial width, narrowly separated mesally or

meeting at preapical denticles. Sharp dorsal and ventral denticles present at base of each tubercle, ventral

denticle not connected by ridge; mesoventral ridge not

forming denticles apically, not connected to ventrobasal denticles; dorsal preapical denticles well developed, sharp or bluntly rounded, ventral preapical denticles poorly developed, blunt; apices tapering outward

from preapical denticles.

The similarities of the pupae of A. cintalpa to the

pupae of irroratus irroratus, cordillerensis, and peruvianus are discussed under irroratus.

Anthrax peruvianus Marston

A number of pupal skins of this species were collected

by R. H. Painter from nests of bees in ruins at Pachacamac, near Lima, Peru. Although no specimens were

reared, adults were collected ovipositing in the same

area. Pupal skins are in the collections of R. H. Painter

and the author.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles. Medial

cephalic tubercle located slightly outside line between

anterior and posterior tubercles, slightly closer to posterior tubercle (Plate 1B) ; each anterior tubercle narrow, acuminate, with sharp mesoventral ridge curving

ventrally at base, sharp lateral ridge curving dorsally

at base, and less distinct ventral ridge; medial tubercle

acuminate to sharp apex, bent slightly forward at base,

with well-defined posterolateral ridge connecting to

posterior tubercle and poorly defined dorsolateral and

mesal ridges basally; posterior tubercle short, thick,

curved forward to sharp apex, with sharp ventromesal,

ventrolateral, and dorsal ridges. Facial sclerite about

four and a half times longer than narrowest width,

shallowly emarginate at posterior end; anterior facial

tubercles relatively long, flattened anteroposteriorly,

tapering from meson to sharp lateral apices, barely

meeting on meson; posterior facial tubercle composed

of two short, anteroposteriorly flattened, sharply

rounded nodes. Anterior facial setae arising distinctly

anterior to anterior facial tubercles.

Thoracic spiracle with about seven segments, the

posterior ones reduced; sharp, even flange present

around anterior three-fourths. Abdominal spiracles

with about six sections, ventral ones reduced; erect,

broad dorsal flange narrowing to anterior end and to

posterior connection with broad, recumbent, flaplike

ventral flange.

Sixth and seventh terga without spines between

setae. Eighth tergum without spines, with setae on line

in front of dorsolateral tumidities. Ninth tergum with

long mesal process tapering from broad base to sharply

rounded apex. Anal tubercles (Plate 4B) about four

times longer than medial width of one tubercle, separated medially by about one-fourth width of one tubercle. Base of each tubercle with large, acute, dorsal and

ventral denticles connected by high, sharp lateral

flange, ventromesal margin with sharp denticle about

two-fifths of distance to apex which curves basally as

sharp ridge to connect with ventrobasal denticle; mesal

margin with blunt dorsal and ventral preapical denticles from which tubercle tapers laterally to sharp

apex.

The relationships of A. peruvianus, cordillerensis, irroratus irroratus, and cintalpa are discussed under

irroratus.

Anthrax cordillerensis Marston

One specimen reared from nest cells of Hypodynerus

Saussure (Hymenoptera: Vespidae) in Santiago Province, Chile, has been studied (see table). It is in the

author's collection.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles. Medial

cephalic tubercle located distinctly outside line between

anterior and posterior tubercles, closer to posterior

tubercle; each anterior tubercle triangular in cross section; acuminate, with sharp dorsomesal, ventromesal,

and lateral ridges; medial tubercle acuminate to sharp

apex, bent slightly forward, with well-defined posterolateral ridge connecting to posterior tubercle and less

well-defined dorsolateral and mesal ridges basally; posterior tubercle short, thick, acuminate, curving forward

distally, with sharp dorsal ridge and poorly defined

ventral and mesal ridges. Facial sclerite about four

times longer than narrowest width, shallowly emargi-
12

nate at posterior end; anterior facial tubercles flattened anteroposteriorly, about as long as basal width

of one tubercle, tapering from meson to sharp lateral

apices, barely meeting on meson; posterior facial tubercle composed of two short, anteroposteriorly flattened

and sharply rounded nodes. Anterior facial setae arising distinctly anterior to anterior facial tubercles.

Thoracic spiracle with about six segments, posterior

ones reduced; sharp flange present around anterior

half. Abdominal spiracles with about five sections, ventral ones reduced; erect, broad, dorsal flange narrowing to posterior end and to anterior connection with

broad, recumbent, flaplike ventral flange.

Sixth tergum with spines between setae; seventh

tergum with setae, but no spines. Eighth tergum with

row of setae in front of each dorsolateral tumidity.

Medial process on ninth tergum about as long as width

at base, tapering to sharp tip. Anal tubercles about

three and a half times longer than medial width of one

tubercle, separately medially by about one-fourth

width of one tubercle. Base of each tubercle with large

dorsal and ventral denticles, lateral connecting ridge

indistinct; ventromesal margin with sharp ridge extending about two-fifths of distance to apex and ending

in blunt angle; mesal margin with sharp dorsal and

ventral denticles from which tubercle tapers laterally

to sharp apex.

Relationships of A. peruvianus, cintalpa, irroratus

irroratus, and cordillerensis are discussed under

irroratus.

Anthrax melanopogon (Bigot)

One reared specimen of this species has been examined

(see table). It is pinned with a nest cell of Megachile

gentilis Cresson (Hymenoptera: Megachilidae). The

specimen is in the collection of the author.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped tubercles and

spines. Medial cephalic tubercle (Plate 1G) formed

slightly outside line between anterior and posterior

ones, and distinctly closer to posterior one; anterior

tubercle long and acuminate, with sharp mesal ridge

curving ventrally at base and lateral ridge curving

dorsally at base; medial tubercle triangular in cross

section, acuminate, with sharp dorsolateral and dorsomesal ridges, and sharp ventrolateral ridge which extends basad laterally and does not join with dorsal

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ridge of posterior pair; posterior tubercle short,

curved dorsally from base, acute, with sharp dorsal

and mesal ridge and less distinct ventrolateral ridge.

Facial sclerite about five and one half times longer

than narrowest width, trunctate posteriorly; anterior

facial tubercles short, tapering to acute apices, with

mesal ridges meeting indistinctly on meson; posterior

tubercle transverse, low, and indistinctly bifid.

Thoracic spiracle with about seven sections, the

posterior ones reduced, with sharp flange around

anterior half. Abdominal spiracles with six sections,

the anteroventral ones reduced; sharp flange present

around anterodorsal three-fourths, expanded as blunt

lobe posterodorsally on anterior pairs.

Sixth and seventh terga without spines between

setae. Eighth tergum without spines, with setae on line

in front of dorsolateral tumidities. Ninth tergum with

simple, transverse medial process. Anal tubercles

(Plate 4D) short, broad, about four times longer than

medial width of one lobe, separated mesally by about

three-fifths of width of one lobe. Each lobe with

strong dorsal and weak ventral denticles basally which

are indistinctly connected by lateral ridge; bluntly

rounded denticle present about halfway to apex as

apical end of sharp ventromesal ridge; acute dorsal

and blunt ventral denticles present mesally about

three-fourths of distance to apex; tubercle tapering

outward from preapical denticles to acuminate apex.

The pupae of A. atriplex and melanopogon differ

little; probably the best character to separate them is

the extension of the posteromesal ridge on the medial

cephalic tubercle of atriplex to the posterior margin of

the cephalic area. The ridge becomes indistinct near

the bases of the tubercle in the one specimen of melanopogon examined. Differences in the numbers of

setae and spines may be attributed to the small size

of the specimen of melanopogon examined.

Both A. melanopogon and atriplex differ from

cintalpa, peruvianus, and irroratus irroratus in that

the flange on the anterior abdominal spiracles is lobed

rather than evenly rounded posterodorsally.

Anthrax atriplex Marston

Two specimens of this species from the National

Museum of Natural History were reared by Krombein

(1967) from the nests of Megachile gentilis Cresson
NUMBER 100 13

(Hymenoptera: Megachilidae). Pupal skins are

pinned with the specimens.

Integument of pupa yellowish hyaline with yellow

setae and reddish brown black-tipped spines and tubercles. Medial cephalic tubercle (Plate IF) formed outside a line between anterior and posterior ones, and

closer to posterior one; anterior tubercle of moderate

length, tapering to acute apex, with sharp mesal ridge

curving ventrally, sharp lateral ridge curving dorsally,

and less distinct dorsal and ventral ridges basally;

medial tubercle bent dorsolaterally from base at slight

angle, broadly triangular basally, tapering to acute

apex, with sharp ventrolateral ridge connecting to posterior turbercles, and sharp ventromesal ridge continuing around to posterior margin of cephalic area;

posterior tubercle short, curving anteromesally to

sharp apex, concave anteromesally, with sharp anterolateral, posterolateral, and posteromesal ridges. Facial

sclerite about seven times longer than narrowest width,

posterior end with shallow, broad emargination; anterior facial sclerites short, indistinctly united mesally,

not flattened anteroposteriorly, with rugose surfaces

tapering to sharp apices; posterior facial tubercle represented by low, binodal, transverse ridge. Bases of

anterior facial setae distinctly in front of bases of anterior facial tubercles.

Thoracic spiracle with about eight sections, posterior

ones reduced, with sharp flange around anterior twothirds. Abdominal spiracles each with about six sections, the anteroventral ones reduced, with broad,

sharp flange anteroventrally, narrowing anteriorly then

broadening anterodorsally and forming sharp spur

posteriorly.

Sixth and seventh terga without spines between

setae. Eighth tergum without spines dorsally, with

about three setae on each side in front of and mesad

of dorsolateral tumidities. Ninth tergum with transverse, bifid, mesal process. Each anal tubercle (Plate

4G) about four times longer than medial width; tubercles separated mesally by about half the medial width

of one tubercle. Each tubercle with strong dorsal and

ventral denticles near base not connected by lateral

flange (indistinct ridge sometimes present); ventromesal margin with denticle about halfway to apex

extending basally as sharp flange; sharp dorsomesal

and blunt, poorly defined, ventromesal denticles present about three-fourths of distance to apex; tubercle

tapering outward from preapical denticles to bluntly

acuminate, slightly incurved apex.

The relationship of A. atriplex to melanopogon and

other species in the genus is discussed under melanopogon.

Literature Cited

Baker, C. F.

1895. Biological Notes on Some Colorado Diptera. Entomological News, 6:173-174.

Brooks, A. R.

1952. Identification of Bombyliid Parasites and Hyperparasites of Phalaenidae of the Prairie Provinces

of Canada, with Descriptions of Six Other Bombyliid Pupae (Diptera). Canadian Entomologist,

84:357-373, 46 figures.

Cole, F. R.

1957. New Bombyliid Flies from Chiapas, Mexico. PanPacific Entomologist, 33:200-202.

Davidson, A.

1894. On the Parasites of Wild Bees in California. Entomological News and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia, 5:170-172.

Hall,J.C.

1954. Notes on the Biologies of Three Species of Bombyliidae, with a Description of One New Species.

Entomological News, 65:145-149.

Hurd, P. D.

1959. Beefly Parasitism of the American Carpenter Bees

Belonging to the Genus Xylocopa Latreille (Diptera: Hymenoptera). Journal of the Kansas Entomological Society, 32:53-58.

Krombein, K. V.

1967. Trap-Nesting Wasps and Bees: Life Histories, Nests,

and Associates. 570 pages. Washington, D.C.,

Smithsonian Institution Press.

Malloch, J. R.

1915. Some Additional Records of Chironomidae for Illinois and Notes on Other Illinois Diptera. Bulletin

of the Illinois State Laboratory of Natural History,

11:327-334.

1917. A Preliminary Classification of Diptera, Exclusive

of Pupipara, Based Upon Larval and Pupal Characters, with Keys to the Imagines in Certain Families.

Part I. Bulletin of the Illinois State Laboratory

of Natural History (1918), 12:161-409, plates

28-57.

Marston, N.

1963. A Revision of the Nearctic Species of the albofasciatus Group of the Genus Anthrax (Diptera; Bornbyliidae). Technical Bulletin of the Kansas Agricultural Experiment Station, Number 127, 79 pages.

1964. The Biology of Anthrax limatulus fur (Osten

Sacken), with a Key to and Descriptions of Pupae

of Some Species in the Anthrax albofasciatus and

trimaculatus Groups (Diptera; Bombyliidae).

Journal of the Kansas Entomological Society,

37(2):89-105.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1970. Revision of New World Species of Anthrax (Dip- Shelford, V. E.

tera: Bombyliidae), Other Than the Anthrax 1913. The Life-History of a Bee-Fly (Spogostylum anale

albofasciatus Group. Smithsonian Contributions Say), a Parasite of the Larva of a Tiger Beetle

to Zoology, number 43, 148 pages. (Cicindella scutellaria Say, var. lecontei Hald.).

Annals of the Entomological Society of America,

Rau> p

- 6:213-225.

1940. Some Mud-Daubing Wasps of Mexico and Their Townsend C. H. T.

Parasites. Annals of the Entomological Society of 1893. The Pupa of Argyramoeba oedipus Fabr. AmeriAmerica, 33:590-595. can Naturalist, 27:60-63.
NUMBER 100 15

PLATE 1.—Anthrax species: A, Lateral view of pupal skin of A. limatulus fur (aft, anterior

facial tubercles; atb, anal tubercle; ctb, cephalic tubercles; pft, posterior facial tubercle).

Ventral views of cephalic tubercles of pupal skins: B, peruvianus; c, cintalpa; D, irrorattu

irroratus; E, nidicola; F, atriplex; G, melanopogon; H, limatulus fur.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 2.—Ventral views of cephalic tubercles of pupal skins of Anthrax species: A, analis;

B, koebelei; c, albofasciatus albofasciatus; D, argyropygus argyropygus; E, hyalacrus; F, pluricellus;

G, xylocopae; H, tigrinus; i, trimaculatus.
NUMBER 10 0 17

PLATE 3.—Dorsal views of caudal tubercles of pupal skins of Anthrax species: A, analis;

B, koebelei; c, xylocopae; D, tigrinus; E, trimaculatus; v, pluricellus; G, argyropygus argyropygus;

H, hyalacrus.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 4.—Dorsal views of caudal tubercles of pupal skins of Anthrax species: A, albofasciatus

albofasciatus; B, peruvianus; c, irroratus irroratus; v, melanopogon; E, nidicola; F, limatulus

fur; G, atriplex; H, cintalpa.

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