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Vol. 56

APRIL 1980

No. 2

THE

Pan-Pacific Entomologist

NELSON—Nomenclatural Changes in the Family Buprestidae with Descriptions of

Previously Unknown Sexes (Coleoptera)_ 81

KIMSEY—Notes on the Biology of some Panamanian Pompilidae, with a Description

of a Communal Nest (Hymenoptera) _ 98

BLICKLE—A New Oxyethira (Hydroptilidae, Trichoptera) of the Aeola Ross Group;

with a Key to Separate the Five Males of the Group_ 101

HAAS et al.—Fleas from some Alaskan Birds (Siphonaptera) _ 105

KENNETT—Occurrence of Metaphycus bartletti Annecke and Mynhardt, a South
African Parasite of Black Scale, Sciissetici oleae (Olivier) in Central and Northern
California (Hymenoptera: Encyrtidae; Homoptera: Coccidae)_ 107

DESHEFY—Anti-predator Behavior in Swarms of Rhagovelia obesa (Hemiptera:

Veliidae)_ 111

CHEMSAK—Revision of the Genus Stenobatyle Casey (Coleoptera: Cerambycidae) 113

DOYEN and KITAYAMA—Review of the North American Species of Apocrypha
Eschscholtz, with a Description of the Immature Stages of Apocrypha anthicoides
(Coleoptera: Tenebrionidae) _ 121

GROGAN and WIRTH—Two New Species of Macrurohelea from Chile with a Key

to the Neotropical Species (Diptera: Ceratopogonidae)_ 137

EICKWORT and ABRAMS—Parasitism of Sweat Bees in the Genus Agapostemon
by Cuckoo Bees in the Genus Nomada (Hymenoptera: Halictidae, Antho-
phoridae)_ 144

MANLEY —Dasymutilla phoenix (Fox), a New Synonym of D. foxi (Cockerell)

(Hymenoptera: Mutillidae) _ 153

SCIENTIFIC NOTES

Arnaud and Davies—Note on Collembolan Household Pest_ 155

Stone—Notes of Life History of Three Conoderus Species_ 157

NOTICES_ 143, 152

ZOOLOGICAL NOMENCLATURE_ 97

SAN FRANCISCO, CALIFORNIA • 1980

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

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T. D. Eichlin, A. R. Hardy, Co-Editors
S. Kuba, Editorial Asst.

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OFFICERS FOR 1980

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Title of Publication: The Pan-Pacific Entomologist.

Frequency of Issue: Quarterly (January, April, July, October).

Location of Office of Publication, Business Office of Publisher and Owner: Pacific Coast Entomological Society,
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This issue mailed 17 April 1980

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The Pan-Pacific Entomologist (ISSN 0031-0603)

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PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 81-97

NOMENCLATURAL CHANGES IN THE FAMILY BUPRESTIDAE
WITH DESCRIPTIONS OF PREVIOUSLY UNKNOWN SEXES

(COLEOPTERA) 1

G. H. Nelson 2

College of Osteopathic Medicine of the Pacific, Pomona, California 91766

Investigation of the status of published scientific names in the family
Buprestidae has indicated the need to cite new synonyms and/or indicate
changes in the status of others. Some are included here with descriptions
of previously unknown sexes. The genera and species are listed in alpha¬
betical order for easier reference. Unless otherwise indicated, specimens
are in the collector’s collection. Abbreviations of collections [brackets] are
as published in Arnett and Samuelson (1969). Collections not in that work
include: British Museum (Natural History) = BMNH; Museum National
d’Historie Naturelle, Paris = MNHP; Narodni Museum, Prague = NMPC;
Hungarian Natural History Museum, Budapest = HNHM; Zoological Mu¬
seum, Academy of Sciences, Leningrad = ZMAS; and Zoological Museum,
University of Moscow = ZMUM. My name is abbreviated GHN. For some
type specimens label data is given. The hash mark (/) separates data on
individual labels and the abbreviations (h) = handwritten and (p) = printed.

Since several of the late Josef N. Knull’s species are involved in this
paper, it seems worthwhile to mention that his collection, including types
of the many species of Buprestidae he described, was willed to the Field
Museum of Natural History, Chicago, IL [FMNH]. Much of the overflow
material is included in the insect collection of the Ohio State Univ. [OSUC],
where Professor Knull taught.

Thanks are extended to the following individuals and institutions, either
for helpful suggestions, for the loan of material, or for comparing specimens
with types in their care: Anatoly Alexeev, S. Keleinikova, ZMUM, and
Mark Volkovitsch, ZMAS, U.S.S.R.; W. F. Barr, UIMC; Svatopluk Bily
and J. Jelinek, NMPC; A. Descarpentries, MNHP; C. M. F. von Hayek,
BMNH; H. A. Hespenheide, Los Angeles, Calif; Z. Kaszab, HNHM; D.
H. Kavanaugh, CASC; J. M. Kingsolver, USNM; Eric H. Smith, FMNH;
Margaret K. Thayer, MCZC; G. C. Walters, Jr., Los Angeles, Calif.; R. L.
Westcott, ODAC; and the publications committee of the Division of Plant
Industry, Florida Department of Agriculture and Consumer Services. Spe¬
cial thanks are due Ms. Jodelle Alexander for typing the manuscript.

PAN-PACIFIC ENTOMOLOGIST

Agrilus cliftoni Knull
Agrilus cliftoni Knull, 1941:382.

This species was described from a unique male. Others have since been
collected, usually on Juglans nigra L.

Description, female. —Differs from male in being slightly more robust;
front of head aeneous instead of bluish green; first 2 abdominal sternites
convex; tibiae not toothed on inner margin at apex.

Length 6.1 mm; width 1.7 mm.

Indiana, Tippecanoe Co., 14 July 1963, N. M. Downie, on Juglans nigra
L. [GHNC] (New State Record).

The 16 males are from 4.3 to 5.8 mm long; the 40 females are from 4.3 to
6.3 mm.

Agrilus duncani Knull
Agrilus duncani Knull, 1929:270.

This species was described from a unique male lacking metatarsi. Many
specimens have been taken during the intervening years (Nelson, 1965;
Knull, 1970). The claws are similarly toothed on all tarsi.

Description, female. —Differs from male in being slightly more robust;
pro sternal pubescence shorter and less dense; first 2 abdominal sternites
convex, without midline groove.

Length 7.3 mm; width 2.0 mm.

Arizona, Cochise Co., E of Coronado Nat. Monument, 24 August 1964,
GHN and family, on Chrysothamnus nauseosus (Pall.) Britton [GHNC].

The 53 males are from 5.4 to 6.9 mm long; the 51 females are from 5.4 to
7.5 mm.

Agrilus fisherellus Obenberger

Agrilus fisherellus Obenberger, 1936a:92 (replacement name for costipennis

Fisher, 1928:182).

Agrilus tarahumarae Cazier, 1951:32 (NEW SYNONYMY).

When Agrilus tarahumarae Cazier was described from a single male from
Mexico, Chihuahua, 80 km N of Chihuahua City, only a few specimens of
Agrilus fisherellus Obenberger were available for study. More have since
been collected in California, Arizona, and Texas, and the holotypes of both
A. costipennis and A. tarahumarae have beemstudied and compared. The
characteristics indicated by Cazier (1951) to distinguish A. tarahumarae
from A. fisherellus are the concave head, deep median pronotal impression,
and larger size. These are variable in a series, with specimens from Cali¬
fornia having the general pubescence more pronounced and the concavity

VOLUME 56, NUMBER 2

Figs. 1-5. Fig. 1. Agrilus nevadensis Horn, male genitalia, dorsal view (left) and ventral
view (right), (line = 2 mm). Fig. 2. Brachys apachei Knull, male genitalia, dorsal view (left)
and ventral view (right). Fig. 3. Brachys cephalicus Schaeffer, male genitalia, dorsal view
(left) and ventral view (right), (line = 1 mm for Figs. 2, 3). Fig. 4. Buprestis ( Buprestis )
parmaculativentris Knull, last visible abdominal sternite, female. Fig. 5. B. ( B .) parmacula-
tiventris Knull, last visible abdominal sternite, male, (line = 2 mm for Figs. 4, 5).

PAN-PACIFIC ENTOMOLOGIST

of the head less pronounced than those from Texas and Mexico, and spec¬
imens from Arizona being somewhat intermediate. The 21 specimens stud¬
ied are from 6.5 to 10.5 mm long, and since the male genitalia are similar,
as pointed out by Cazier, I consider the 2 names to represent variants of a
single widespread species.

Agrilus fisherellus Obenb. was based on 2 males from Pasadena and Los
Angeles in southern California without biological data (Fisher, 1928). This
species was recorded by Knull (1944) as on Celtis reticulata Sargent in the
Davis and Chisos Mts., Texas and the Chiricahua Mts., Arizona. Wellso
(1973) recorded it as A. tarahumarae and described the female from Texas,
Barksdale, on flowers of Stillingia sp. Adults of this species are found on
diverse hosts as indicated by the foregoing and the following records. Cal¬
ifornia: San Bernardino Co., 12 mi NW of Essex, 16 June 1962, GHN, on
Acacia greggii Gray; Riverside Co., above Palm Desert, elev. 610 m, High¬
way 74, 10 June 1965, GHN, on Baccharis sergiloides Gray; same locality
and plant, 15 June 1968, G. C. Walters; same locality, 8 June 1974, D. S.
Verity, flying about young growing tips Prosopis glandulosa var. torreyana
L. Benson. Arizona: Chiricahua Mts. 9.7 km above SW Res. Sta., 23 July
1969, GHN, on Quercus hypoleucoides A. Camus. Texas: 3.2 km NW of
Fort Davis, Highway 118, 17 June 1965, GHN, on Celtis laevigata Willd-
enow: N of Fort Davis, Highway 118, 16 June 1963, GHN, on Celtis pallida
Torrey; Comal Co., Canyon Lake, 13 July 1975, GHN, on Celtis mississip-
piensis var. reticulata Torrey.

Agrilus nevadensis Horn
Agrilus nevadensis Horn, 1891:303.

The unique female type was the only specimen available to Fisher (1928)
when he revised the genus. Carlson & Knight (1969) regarded the type of
A. nevadensis Horn as deformed and a synonym of Agrilus quadriguttatus
niveiventris Horn. More specimens are now available and indicate that it
should be considered a valid species.

Description, male. —Differs from female as follows: more slender; head
with front blue-green, vertex cupreous, pronotum aeneocupreous, elytra
darkly cupreous, and beneath cupreous with blue-green tints anteriorly to¬
ward midline and on legs; abdominal sternites 1 and 2 with pronounced
median groove, broad and punctured toward base, narrowed and smooth
apically. Genitalia, Fig. 1.

Length 9.0 mm; width 2.0 mm.

California, Trinity Co., Trinity Nat. Forest, 23 June 1919, R. Hopping
[CASC]. Others from California: Trinity Co., Carrville, 19 June 1931; 22
May 1934 [CASC, GHNC]; Plumas Co., Meadow Val., elev. 1220-1524 m,

VOLUME 56, NUMBER 2

10 June 1924, E. C. VanDyke [CASC, GHNC] (New State Record). The
type from western Nevada is in the Horn collection [MCZC].

While A. nevadensis resembles A. q. quadriguttatus Gory and A. q.
niveiventris superficially, it is readily distinguished from both by the follow¬
ing features: pronotum and elytra more coarsely sculptured; pronotum with
median depressions more pronounced; apices of elytra acutely produced;
male with abdominal sternites 1 and 2 with broad median groove. The gen¬
eral color of nevadensis resembles q. niveiventris, but the male genitalia
resemble q. quadriguttatus. Fisher (1928) stated that the scutellum is not
carinate, but as Carlson & Knight (1969) stated, the scutellum on the type
does appear to be transversely carinate as it is on the other 6 examined. No
significant variation was noted in this small series.

Agrilus nigricans Gory
Agrilus nigricans Gory, 1841:257.

Agrilus auricomus Frost, 1912:250 (NEW SYNONYMY).

In lists or revisions of North American Agrilus (Leng, 1920; Fisher, 1928;
Obenberger, 1936b) Agrilus nigricans Gory has been considered a synonym
of Agrilus obsoletoguttatus Gory. That they are not the same was first
indicated by A. Descarpentries (in litt.). However, comparison of the male
lectotype of A. nigricans Gory [MNHP] with the male holotype of Agrilus
auricomus Frost [MCZC] reveals that they represent the same species. The
type of A. nigricans is a small example of the species (6.0 mm) with a bluish
tint on some parts of the body, possibly due to discoloration. The color of
the hair is whitish and not golden as in the holotype of A. auricomus, but
is similar in that respect to 1 paratype in the latter. In other salient features,
including male genitalia, they compare favorably, and I consider A. auri¬
comus Frost a junior synonym of A. nigricans Gory.

There are 2 A. nigricans, 1 male and 1 female, in the MNHP. The male,
here designated as lectotype, has the following labels: Red label with
“TYPE” (p)/white label with “ nigricans Gory” (h).

The type locality for A. nigricans Gory is “Amerique Boreale”; for A.
auricomus Frost, “Framingham, Mass.”

Agrilus palmerleei Knull
Agrilus palmerleei Knull, 1944:76.

Description, female .—Differs from male in being more robust; head and
pronotum more deeply reddish cupreous; pro- and mesotibiae not toothed
on inner margin at apex.

Length 10.3 mm; width 2.8 mm.

PAN-PACIFIC ENTOMOLOGIST

Arizona, Chiricahua Mts., 9.6 km above S. W. Res. Sta., 24 July 1969,
GHN and D. E. Nelson, on Quercus hypoleucoides A. Camus [GHNC].

The 8 males are from 8.7 to 10.0 mm long; the 9 females are from 9.3 to
11.3 mm.

Anthaxia ( Anthaxia ) prasina Horn
Anthaxia ( Anthaxia ) prasina Horn, 1882:108.

This species was characterized by Horn, under his broad concept of An¬
thaxia aeneogaster Laporte and Gory, as the brilliantly green form from
“Yosemite Valley of California,” which “for convenience” was called pra¬
sina. He indicated having seen 8 specimens. There are 5 specimens now in
the Horn collection [MCZC], 4 males and 1 female. Each has the same 2
labels, “Cal” (p) and “Horn Coll, H.” (p). Apparently no type was ever
designated, so I designate 1 male as lectotype and the other 4 specimens as
paralectotypes. The lectotype is 4.5 mm long and 2.0 mm wide. Two males
in the LeConte collection, which were also part of the series before Horn,
are designated as paralectotypes. One is labelled—“Cal” (p)/“A. prasina
Horn” (h )/“aeneogaster 25” (h); the other—“Cal” (p )/“aeneogaster 26”

(h).

Anthaxia ( Haplanthaxia ) cyanella Gory
Anthaxia ( Haplanthaxia ) cyanella Gory, 1841:285.

Anthaxia kaszabiana Pochon, 1967:282 (NEW SYNONYMY).

An examination of the type of Anthaxia kaszabiana Pochon [HNHM],
collected in Fundort, Ohio, reveals it is a male of Anthaxia cyanella Gory.
The latter was described from a female, which is deep blue throughout while
the male is green and bronzy-brown.

Brachys apachei Knull
Brachys apachei Knull, 1952:359.

This species was described from 2 females collected in Arizona, Chiri¬
cahua Mts., 17 June 1939, by D. J. & J. N. Knull, on oak foliage.

Description, male. —The male is much like the female holotype in general
appearance, but is less robust; and apical margin of last visible abdominal
sternite is roundly truncate, without row of deflexed teeth and submarginal
ridge without row of longer hairs. Genitalia, Fig. 2.

Length 3.8 mm; width 1.8 mm.

Arizona, Chiricahua Mts., 6.4 km above Portal, 12 July 1977, GHN, on
foliage Quercus hypoleucoides A. Camus.

VOLUME 56, NUMBER 2

In the Knull collection [FMNH] 17 females are from 3.1 to 4.3 mm long
and 1.4 to 2.2 mm wide, 13 males from 3.2 to 4.0 mm long and 1.4 to 1.8
mm wide.

In the original description this species was compared with Brachys
cephalicus Shaeffer and Brachys aeruginosas Gory. The latter is an eastern
species with the front of the head less obliquely retracted than in B. apachei.
B. cephalicus, sympatric with B. apachei, differs in being more robust, in
having more strongly convex tubercles above the eyes, and in the shape of
the male genitalia, Fig. 3.

Buprestis ( Buprestis) parmaculativentris Knull
Buprestis {Buprestis) parmaculativentris Knull, 1958:154.

This species was described from a unique male.

Description, female. —Differs from male as follows: more robust; yellow-
red on front of head confined to spots near medial margin of eyes; antennae
shorter, not reaching hind angles of pronotum when laid along side; protibia
without internal recurved spine at apex; disk of last visible abdominal ster-
nite more convex toward apex, apical margin convexly truncate in female,
Fig. 4, concavely truncate in male, Fig. 5.

Length 20.0 mm; width 8.0 mm.

Texas, Chisos Mts. Basin, 21 June 1965, GHN, on Pinus cembroides
Zuccarini.

The 5 males available for study are from 16.0 to 17.5 mm long and from
6.5 to 7.0 mm wide; the 15 females from 16.5 to 21.5 mm long and from 7.0
to 8.5 mm wide.

Buprestis (Stereosa) Salisburyensis Herbst
Buprestis (Stereosa) salisburyensis Herbst, 1801:174.

Buprestis salisburyensis cazieri Heifer, 1946:100 (NEW SYNONYMY).

Buprestis salisburyensis cazieri Heifer, a dark form, was described as a
subspecies of B. salisburyensis Herbst. The more common color is green
with cupreous elytral margins while “ cazieri ” is dark green to purplish.
Since color is the only distinguishing feature and since the 2 forms occur
sympatrically it seems advisable to regard B. cazieri as a dark color variant
of B. salisburyensis. The occurrence of dark or cupreous color phases of
green forms is not rare among Buprestidae. Others of this genus in which
this color variation occurs include: Buprestis striata Fabricius, in which the
dark color seems more common; and Buprestis decora Fabricius in which
dark forms are uncommon.

PAN-PACIFIC ENTOMOLOGIST

Chalcophorella langeri (Chevrolat)

Chalcophorella langeri (Chevrolat), 1853:308 ( Chalcophora).

Chalcophorella strandi (Obenberger), 1936c: 109 (Texania) (NEW SYNON¬
YMY).

When Chalcophorella strandi (Obenberger) was described, Chalcopho¬
rella langeri (Chevrolat) was not considered valid, and C. strandi was com¬
pared to Texania fulleri (Horn), Texania campestris (Say) and Texania
serriger (Casey). The last is a synonym of C. campestris. Knull (1956)
recognized C. langeri as a valid species and presented a key for separating
the 3 North American species: C. fulleri, C. campestris, C. langeri. Ex¬
amination of the female type of C. strandi and comparison of it with homo¬
types of C. langeri reveal it is a junior synonym of the latter.

The type locality for Chalcophora langeri Chevrolat is “Greenville, pres
de la Nouvelle—Orleans” [Louisiana], female type [BMNH]; for Texania
strandi Obenberger, “Fort Madison” [Iowa], type [NMPC].

Chrysobothris chlorocephala Gory
Chrysobothris chlorocephala Gory, 1841:161.

Chrysobothris concinnula LeConte, 1860:238 (NEW SYNONYMY).

A color slide of the type of C. chlorocephala Gory [MNHP] and sketches
of its clypeus and last visible abdominal sternite made by H. A. Hespenheide
reveal it is the same species as C. concinnula LeConte. The male type of
C. chlorocephala is 5.5 mm long. Fisher (1942) had not seen the type of C.
chlorocephala and, following Horn (1886), erroneously placed it as a syn¬
onym of Chrysobothris harrisi (Hentz).

This species is closely related to Chrysobothris scitula Gory, and further
study may show the two to be conspecific, as suggested by Fisher (1942).
In C. chlorocephala the violaceous brown color predominates above with
iridescent blue or blue-green spots, but in C. scitula the latter color pre¬
dominates with the violaceous brown reduced. A color slide of the type of
C. scitula [MNHP] indicates Fisher assessed its characteristics accurately.

Chrysobothris cribraria Mannerheim
Chrysobothris cribraria Mannerheim, 1837:77.

Chrysobothris floricola Gory, 1841:179 (NEW SYNONYMY).

Chrysobothris calcarata Melsheimer, 1845:146 (NEW SYNONYMY).

The name Chrysobothris cribraria Mannerheim has been considered in¬
correctly a synonym of Chrysobothris femorata (Olivier) by American
workers for many years (see Fisher, 1942). An examination of the lectotype

VOLUME 56, NUMBER 2

(Nelson, 1976) revealed it to be the same species that Gory later described
as floricola and Melsheimer as calcarata. This species is widely distributed
in the pine-growing regions of eastern North America.

Chrysobothris georgei Nelson

Chrysobothris georgei Nelson, REPLACEMENT NAME for Chrysobothris
vogti Nelson, 1975:18, preoccupied by Chrysobothris ephedrae vogti
Knull, 1964:376.

I overlooked the subspecies described by Knull, so the new name C.
georgei is proposed for the junior homonym, named in honor of Mr. George
B. Vogt.

Chrysobothris purpurifrons Motschulsky
Chrysobothris purpurifrons Motschulsky, 1859:183.

Chrysobothris pubescens Fall, 1907:238 (NEW SYNONYMY).

Chrysobothris subpubescens VanDyke, 1937:112 (error for pubescens ).

Four species of Buprestidae were described from California in the above
paper by Motschulsky. The author of Polycesta californica, and Chryso¬
bothris subcylindrica was Menetries, while Chrysobothris purpurifrons and
Belionota californica were described by Motschulsky. In Horn (1886), Fish¬
er (1942), and Barr & Westcott (1976), Motschulsky was listed as the author
of C. subcylindrica.

The holotype of C. purpurifrons Motschulsky [ZMUM] compares well
with the type of C. pubescens Fall in most features including shape and
sculpture of pronotum and elytra, length of body hairs, shape of clypeus,
and emargination of last visible abdominal sternite of females. The lateral
margins of the elytra are more weakly serrate than in most C. pubescens
Fall seen, but I feel the two represent the same species. Thus C. purpurifrons
Motsch. is not synonymous with Chrysobothris semisculpta LeConte, as
indicated by Horn (1886) and Fisher (1942).

The female holotype of C. purpurifrons Motschulsky is in poor condition.
Ventral surface is coated with mold. Left front and hind legs missing; right
protarsus and all but basal segment of metatarsus missing. Antennae missing
except for 2 basal segments on right. Abdomen is glued upside down on
card under specimen and 2 antennal segments are glued to same card. Labels
on the type are as follows: gray-green disk with C. (h)/green with Chryso¬
bothris purpurifrons Motsch California (h)/blank red label/red holotype label
det. G. H. Nelson ’79. Dr. S. Keleinikova [ZMUM] kindly loaned the holo-
types of C. purpurifrons Motsch. and C. subcylindrica Menetr., for which
I am deeply thankful.

PAN-PACIFIC ENTOMOLOGIST

Chrysobothris subcylindrica Menetries
Chrysobothris subcylindrica Menetries, in Motschulsky, 1859:182.

Chrysobothris deleta LeConte, 1860:255 (NEW SYNONYMY).

Chrysobothris delecta Obenberger, 1934:619 (typographical error for de¬
leta).

The female holotype of Chrysobothris subclylindrica Menetries compares
well with the female holotype of Chrysobothris deleta LeConte but is larger
(9.0 mm compared to 8.0 mm for C. deleta) and is slightly more cupreous.
Since LeConte’s paper appeared in 1860, C. subcylindrica Menetries has
priority.

The holotype of C. subcylindrica is in poor condition. Antennae missing
except for right basal segment which is attached near its normal position.
Left front leg, and meso- and metatarsi missing; right front leg, part of
mesotarsus, and metatibia and metatarsus missing. Labels on the type are
as follows: white with 60 (h)/white with Californ. Sept (p)/green with Chry¬
sobothris subcylindrica Menetr California (h)/blank red label/red holotype
label, det. G. H. Nelson, ’79.

Chrysobothris ulkei LeConte
Chrysobothris ulkei LeConte, 1860:240.

This species was described from a single female. It has since been col¬
lected in large numbers, and notes on its biology have been recorded (Nel¬
son, 1962; Barr & Westcott, 1976).

Description, male. —Iridescent green above and below, disk of elytra with
3 pairs of violaceous spots, faint pair at basal one fourth, distinct pairs at
middle and apical one fourth. Externally similar to female, but differs in
having apex of last visible abdominal sternite arcuately emarginate, Fig. 6,
instead of sinuate. Genitalia, Fig. 7.

Length 12.5 mm; width 5.5 mm.

Arizona, Cochise Co., 2 mi E of Portal, 28 August 1959, GHN, on Ephed¬
ra trifurca Torrey.

The iridescent bronzy-green general color is more common in males with
the tendency toward bluish hues more common in females. The discal vi-

Figs. 6-10. Fig. 6. Chrysobothris ulkei Horn, last visible abdominal sternite, male. Fig. 7.
C. ulkei Horn, male genitalia, dorsal view (left) and ventral view (right), (line = 1 mm for Figs.
6,7). Fig. 8. Melanophila ( Phaenops) abietis Nelson, a) clypeus; b) dorsal view,
female. Fig. 9. M. (P.) lecontei Obenberger, a) clypeus; b) dorsal view, female. Fig. 10. M.
( P .) drummondi (Kirby), a) clypeus; b) dorsal view, female (line = 5 mm for Figs. 8, 9, 10).

VOLUME 56, NUMBER 2

PAN-PACIFIC ENTOMOLOGIST

olaceous elytral spots vary from 3 distinct pairs to no indication of spots.
Partial reduction in the number of spots usually proceeds from anterior to
posterior. The 24 males studied are from 11.0 to 13.5 mm long and the 31
females from 11.0 to 15.0 mm.

Dystaxia murrayi LeConte
Dystaxia murrayi LeConte, 1866:385.

Dystaxia murrayi cuprea Knull, 1947:72 (NEW SYNONYMY).

Dystaxia murrayi cuprea Knull is a coppery color phase of D. murrayi
LeConte, as suggested by Nelson (1960). An examination of the type of D.
murrayi cuprea and a series of 18 green and 26 cupreous color forms sup¬
ports this conclusion. The 2 forms occur sympatrically.

Melanophila {Melanophila) cuspidata Klug
Melanophila ( Melanophila) cuspidata Klug, 1829:34.

Melanophila pecchiolii (Laporte & Gory), 1839:33, ( Anthaxia ) (NEW

SYNONYMY).

In the literature, Melanophila pecchiolii (Laporte & Gory) has consis¬
tently been listed as a synonym of Melanophila acuminata (DeGeer). A.
Descarpentries indicated (in litt.) that the type of M. pecchiolii [MNHP]
represents the same species as Melanophila cuspidata Klug rather than M.
acuminata. Since M. cuspidata is a European species the synonym M.
pecchiolii should not appear on North American lists.

Melanophila ( Melanophila ) notata notata (Laporte & Gory)

Melanophila ( Melanophila ) notata notata (Laporte & Gory), 1837:4, ( Apa-

tura).

Melanophila opaca LeConte, 1860:213. Melanophila hungarica Csiki, 3
1905:579 (NEW SYNONYMY).

Fisher (1925) established Melanophila opaca LeConte as a synonym of
Melanophila notata (Laporte & Gory) but Sloop (1937), in his revision of
the genus, distinguished M. notata from M. opaca primarily on the basis
of color, the former yellow-maculate, the latter immaculate. In a series of
M. notata, variations from distinctly maculate to immaculate occur and no
constant morphological character has been found to separate them, so Fish¬
er’s concept is valid. Melanophila hungarica Csiki was described from a
single specimen from Hungary [HNHM], An examination of the type re¬
veals it is an immaculate example of M. notata, evidently introduced into
Hungary, and should not be considered indigeous to Europe. Dr. S. Bily

VOLUME 56, NUMBER 2

[NMPC] first recognized that M. hungarica = M. opaca and had so labelled
the type.

Melanophila ( Phaenops) abietis Nelson, (NEW STATUS)

Melanophila {Phaenops) drummondi var. abietis Champlain & Knull,
1923:105.

This form was described as a bright green color variety of Melanophila
drummondi (Kirby). As indicated in the following tabular comparison of
Melanophila abietis Nelson, Melanophila lecontei Obenberger, and M.
drummondi, the differences in body form, color, and elytral margins
show that M. abietis should be regarded as a valid species. It is most similar
to M. lecontei including the male genitalia.

abietis

lecontei

drummondi

(Fig. 8)

(Fig- 9)

(Fig. 10)

Body form

broadly oval

moderately oval

Color

blue-green

bronzy-black to

black, rarely with

bluish tint

Elytral

broadly expla-

moderately ex-

moderately expla-

margins

nate, apically not

planate, apically

nate, apically not

or finely serrate

coarsely serrate

or finely serrate

Pronotal

coarsely rugose,

finely rugose, punc¬

disk

punctures large

tures not evident

Clypeus

deeply emar-

shallowly

ginate

emarginate

Distribution

eastern Canada &

western North

New Hampshire

America

Nanularia {Nanularia) pygmaea (Knull) (NEW COMBINATION)

Hippomelas pygmaea Knull, 1941:386.

An examination of the type reveals it has the characteristics of the genus
Nanularia (s. str.), as defined by Barr (1970).

Pachyschelus laevigatus (Say)

Pachyschelus laevigatus (Say), 1839:164, ( Trachys ).

Pachyschelus oblongus (Motschulsky), 1860:54, ( Metonius ) (NEW SYN¬
ONYMY).

PAN-PACIFIC ENTOMOLOGIST

Specimens of Pachyschelus laevigatus (Say) were compared with the type
of Pachyschelus oblongus (Motschulsky) [ZMUM] by Dr. Alexeev (pers.
comm.) and were found to be conspecific. P. oblongus, listed erroneously
from Venezuela by Obenberger (1937), was described from “Etats Unis.
Tenessee” [sic], while P. laevigatus was described from “New York.”

Pachyschelus purpureus purpureus (Say)

Pachyschelus purpureus purpureus (Say), 1839:164, (Metonius).

Pachyschelus biimpressus (Motschulsky), 1860:54, {Metonius), (NEW

SYNONYMY).

Comparison of specimens of Pachyschelus p. purpureus (Say) with the
type of Pachyschelus biimpressus (Motschulsky) [ZMUM] by Dr. Alexeev
(pers. comm.) indicates they both represent the same species. The latter
was listed by Obenberger (1937) from “America bor.,” but no locality was
given with the original description. P. p. purpureus (Say) was described
from “Indiana.”

Polycesta deserticola Barr
Polycesta deserticola Barr, 1974:6.

Polycesta californica Menetries (not LeConte), in Motschulsky, 1859:180

(NEW SYNONYMY).

This species from southwestern U.S. and northern Mexico was known as
Polycesta velasco Laporte & Gory until Barr (1974) recognized it was dif¬
ferent. An examination of the specimen of Polycesta californica Menetries
[ZMAS], kindly loaned by Mark Volkovitsh and here designated as lecto-
type, reveals it is the same species as Polycesta deserticola Barr. Since
LeConte (1857) described another species as Polycesta californica, the
name P. californica Menetries is not available and thus must be treated as
a synonym of P. deserticola Barr.

The lectotype of Polycesta californica Menetries is labelled as follows:
white label with “California sept. Leconte” (p)/white label with “ Polycesta
californica ex Calif.” (h). This female, 17.0 mm long and 6.0 mm wide,
lacks antennae except for 2 basal segments on left and 3 on right, lacks front
legs, right middle leg and metatarsi except for 2 segments on left. The type
locality listed in the original description is “Col Ross,” a Russian settlement
near San Francisco. It was undoubtedly collected from much farther south.

Literature Cited

Arnett, R. H., Jr., and G. A. Samuelson (eds.). 1969. Directory of Coleoptera collections of
North America (Canada through Panama). Purdue Univ., Lafayette, Ind., 123 pp.

VOLUME 56, NUMBER 2

Barr, W. F. 1970. The subgenera of Nanularia and Hippomelas (Coleoptera: Buprestidae).
Occas. Pap. Biol. Soc. Nev., No. 25, pp. 1-9.

Barr, W. F. 1974. New genera and species of North American Buprestidae (Coleoptera).
Occas. Pap. Biol. Soc. Nev., No. 39, pp. 1-13.

Barr, W. F., and R. L. Westcott. 1976. Taxonomic, biological and distributional notes of
North American Chrysobothris, with the description of a new species from California
(Coleoptera:Buprestidae). Pan-Pac. Entomol., 52(2): 138-153.

Carlson, R. W., and F. B. Knight. 1969. Biology, taxonomy, and evolution of four sympatric
Agrilus beetles (Coleoptera:Buprestidae). Contrib. Amer. Entomol. Inst., 4(3):i-vi, 1-
105.

Cazier, M. A. 1951. The Buprestidae of north central Mexico (Coleoptera). Amer. Mus. Nov.,
No. 1526, 56 pp.

Champlain, A. B., and J. N. Knull. 1923. A new variety of Melanophila drummondi Kby.
(Buprestidae, Coleoptera). Can. Entomol., 55(5): 105.

Chevrolat, A. 1853. Description d’une nouvelle espece de Buprestide. Rev. Mag. Zool., 2nd
Ser., 5:308-309.

Csiki, E. 1905. Coleoptera nova ex Hungaria. Ann. Hist.-Natur. Mus. Nat. Hung., 3:575-582.

Fall, H. C. 1907. New Coleoptera from the Southwest-III. Can. Entomol., 39:235-243.

Fisher, W. S. 1925. A revision of the West Indian Coleoptera of the family Buprestidae. Proc.
U.S. Nat. Mus., 65(9): 1-207.

Fisher, W. S. 1928. A revision of the North American species of buprestid beetles belonging
to the genus Agrilus. U.S. Nat. Mus., Bull. 145, 347 pp.

Fisher, W. S. 1942. A revision of the North American species of buprestid beetles belonging
to the tribe Chrysobothrini. U.S. Dep. Agr., Misc. Pub. No. 470, 275 pp.

Frost, C. A. 1912. New species of Coleoptera of the genus Agrilus. Can. Entomol., 44:245-
252.

Gory, H. L. 1841. Histoire naturelle et iconographie des insectes coleop teres, Suppl. (livrai-
sons 43-52). Paris, pp. 107-111; 126-356.

Heifer, J. R. 1946. A new subspecies of Buprestis (Coleoptera, Buprestidae). Pan-Pac. Ento¬
mol., 22(3): 100-102.

Herbst, J. F. W. 1801. Natursystem aller bekannten in-und auslandischen insecten . . . . , Kafer,
vol. 9. Berlin, pp. 1-344.

Horn, G. H. 1882. Revision of the species of some genera of Buprestidae. Trans. Amer.
Entomol. Soc., 10:101-163.

Horn, G. H. 1886. A monograph of the species of Chrysobothris inhabiting the United States.
Trans. Amer. Entomol. Soc., 13:65-124, 7 plates.

Horn, G. H. 1891. The species of Agrilus of boreal America. Trans. Amer. Entomol. Soc.,
18:277-336.

Klug, J. C. F. 1829. Symbolae physicae .... Vol. 1, p. 34, pi. 3.

Knull, J. N. 1929. Three new species of Agrilus (Coleop.:Buprestidae). Entomol. News,
40:270-272.

Knull, J. N. 1941. Nine new Coleoptera (Plastoceridae, Buprestidae and Cerambycidae). Ohio
J. Sci., 41(5):381-388.

Knull, J. N. 1944. Notes on Agrilus with descriptions of two new species (Bupresti¬
dae: Coleoptera). Ann. Entomol. Soc. Amer., 37(l):75-83.

Knull, J. N. 1947. New North American Coleoptera (Buprestidae, Schizopodidae and Cer¬
ambycidae). Ohio J. Sci., 47(2):69-73.

Knull, J. N. 1952. Two new species of North American Brachys (Coleoptera: Buprestidae).
Ohio J. Sci., 52(6):358-359.

Knull, J. N. 1956. A new Texas Agrilus with notes on Chalcophorella (Coleoptera: Bupres¬
tidae). Entomol. News, 67(2):47—48.

PAN-PACIFIC ENTOMOLOGIST

Knull, J. N. 1958. Two new North American Buprestidae (Coleoptera). Ohio J. Sci., 58(3): 153-
154.

Knull, J. N. 1964. Two new Buprestidae and distribution records of others (Coleoptera). Ohio
J. Sci., 64(5):376—378.

Knull, J. N. 1970. Notes and synonymy on Buprestidae (Coleoptera). Entomol. News, 81:263-
264.

de Laporte, F. L., and H. L. Gory. 1837. Histoire naturelle et iconographie des insectes
coleopteres (livraisons 12-16), Vol. 1, part. Paris, genera paged separately.

de Laporte, F. L., and H. L. Gory. 1839. Histoire naturelle et iconographie des insectes
coleopteres (livraisons 25-36); Vol. 2, part. Paris, genera paged separately.

LeConte, J. L. 1857. Reports on the U.S. Pacific railroad exploration. 47th parallel. Insects,
12:1-72.

LeConte, J. L. 1860. Revision of the Buprestidae of the United States. Trans. Amer. Phil.
Soc. (1859), 11 (n.s.): 187-258.

LeConte, J. L. 1866. Additions to the coleopterous fauna of the United States. No. 1. Proc.
Acad. Natur. Sci. Phila., 18:361-394.

Leng, C. W. 1920. Catalogue of the Coleoptera of America, north of Mexico. J. D. Sherman,
Mt. Vernon, New York, x + 470 pp.

von Mannerheim, C. G. 1837. Enumeration des buprestides, et description de quelques nou-
velles especes de cette tribu de la famille des sternoxes, de la collection de M. le Comte
Mannerheim. Bull. Soc. Imp. Natur. Moscou, 10(8):3—126.

Melsheimer, F. E. 1845. Descriptions of new species of Coleoptera of the United States. Proc.
Acad. Natur. Sci. Phila. (1844), 2:134-160.

von Motschulsky, V. 1859. Coleopteres nouveaux de la Californie. Bull. Soc. Imp. Natur.
Moscou 32, part 2:122-185.

von Motschulsky, V. 1860. Insectes de Indes orientales, et de contrees analogues. Etud.
Entomol., part 8 (1859), pp. 25-118.

Nelson, G. H. 1960. Notes on Buprestidae and Schizopodidae. Bull. Brooklyn Entomol. Soc.,
55(3):70-74.

Nelson, G. H. 1962. Notes on the Buprestidae: Part III. Bull. Brooklyn Entomol. Soc.,
57(2): 56-60.

Nelson, G. H. 1965. Notes on the Buprestidae: Part IV with a new synonym in Chrysobothris.
Bull. Brooklyn Entomol. Soc., 59—60:37^41.

Nelson, G. H. 1975. A review of the basalis group of the genus Chrysobothris
(Coleoptera:Buprestidae). Coleopt. Bull., 29(1): 1-30.

Nelson, G. H. 1976. Lectotype designations in the Mannerheim collection of Buprestidae at
the Universitetets Zoologiska Museum, Helsingfors, Finland. Coleopt. Bull., 30(2): 167-
168.

Obenberger, J. 1934. Coleopterorum catalogus, Vol. 12, pars 132, Buprestidae III. W. Junk,
Berlin, pp. 569-781.

Obenberger, J. 1936a. Synonymia Agrilorum (Col. Bupr.) II. Cas. Ceske Spolecnosti Entomol.,
33:91-92.

Obenberger, J. 1936b. Coleopterorum catalogus, Vol. 13, pars 152, Buprestidae. V. W. Junk,
‘s-Gravenhage, pp. 935-1246.

Obenberger, J. 1936c. Eine festarbeit zum sechzig jahrigen jubilaum meines freundes Univ.—
Prof. Dr. Embrik Strand. Festschr. z. 60 geburtstage v. Prof. Dr. Embrik Strand, 1:
97-145.

Obenberger, J. 1937. Coleopterorum catalogus, Vol. 13, pars 157, Buprestidae VI. W. Junk,
‘s-Gravenhage, pp. 1247-1714.

Pochon, V. H. 1967. Beschreibung neuer buprestiden aus der sammlung des Naturwissen-
schaftlichen Museums, Budapest. 2 Teil. Ann. Hist.-Natur. Mus. Nat. Hung., 59:279-
282.

VOLUME 56, NUMBER 2

Say, T. 1839. Descriptions of new North American insects, and observations on some already
described. Trans. Amer. Phil. Soc. (1836), 6 (n.s.): 155-190.

Sloop, K. D. 1937. A revision of the North American buprestid beetles belonging to the genus
Melanophila (Coleoptera, Buprestidae). Univ. Calif. Publ. Entomol., 7(1): 1-20.

VanDyke, E. C. 1937. Notes and descriptions of North American Buprestidae and Ceram-
bycidae (Coleoptera). Bull. Brooklyn Entomol. Soc., 32(3): 105—116.

Wellso, S. G. 1973. A new species of Anthaxia with notes on other buprestids
(Coleoptera:Buprestidae). Coleopt. Bull., 27(4): 165-168.

Footnotes

1 Contribution No. 383, Bureau of Entomology, Division of Plant Industry, Florida Depart¬
ment of Agriculture and Consumer Services, Gainesville, FL 32602.

2 Research Associate, Florida State Collection of Arthropods, Division of Plant Industry,
Florida Department of Agriculture and Consumer Services.

3 Bfly, 1979. Acta Zool. Bulgarica, 13:48, records Melanophila lecontei Obenberger as a
synonym of M. hungarica Csiki. Dr. Bfly (in lift.) indicated that he erred in relating hungarica
with lecontei and agrees that it = opaca.

ZOOLOGICAL NOMENCLATURE

AN(S) 111 1st November 1979

The Commission hereby gives six months notice of the possible use of its
plenary powers in the following cases, published in Bull. zool. Nom. Vol¬
ume 36, part 3, on 1st November 1979, and would welcome comments and
advice on them from interested zoologists. Correspondence should be ad¬
dressed to the Secretary.

2240 Anaspis Muller, 1764; Luperus Muller, 1764; Lampyris Muller, 1764;
and Clerus Muller, 1764 (Insecta: Coleoptera): proposed designation
of a type species.

2244 Ptilium Gyllenhal, 1827 and Ptenidium Erichson, 1845 (Insecta: Co¬
leoptera): proposed conservation.

2246 Chrysomela flavicornis Suffrian, 1851 and C. tibialis Suffrian, 1851
(Insecta: Coleoptera): proposed conservation.

2146 Rhodesiella plumigera (Loew, 1860) (Insecta: Diptera): proposed
suppression.

Secretary, International Commission on Zoological Nomenclature,

% British Museum (Natural History), Cromwell Road,

London, SW7 5BD, United Kingdom.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 98-100

NOTES ON THE BIOLOGY OF SOME PANAMANIAN POMPILIDAE,
WITH A DESCRIPTION OF A COMMUNAL NEST

(HYMENOPTERA)

Lynn Siri Kimsey

Department of Entomology, University of California, Davis 95616

The large amount of work done by Evans (1966, 1973) on the taxonomy
of the Central American spider wasps has made most pompilids in this
region identifiable, at least to genus, although there are probably many
species yet to be described. Very little is known about the biology of the
neotropical pompilids. This is not a problem unique to these wasps, since
knowledge of most groups of insects in the tropics is inadequate. In many
cases a biologist making a study of a group must become a taxonomist as
well.

The nest of Auplopus esmeralda (Banks) described below represents the
second record of a communal pompilid nest. The first record was the nest
of Auplopus argentifrons (F. Smith) (as Paragenia ), described by F. X.
Williams (1919) in the Philippines. The argentifrons nest consisted of about
24 cells tended by approximately eight females and was built in a bamboo
stump.

The remainder of this paper contains behavioral notes on four species of
spider wasps. Voucher specimens of wasps and prey have been deposited
in the museum of the Department of Entomology, University of California
at Davis.

Auplopus esmeralda (Banks). A nest was found on Barro Colorado Island
in the Canal Zone on W. M. Wheeler 8 trail. It was first observed in August
1976 when four females were seen flying in and out of the base of a fallen
Scheelea palm frond, 2 m south of the trail. The palm frond was examined
at the island field station on September 22, 1976. At this time many of the
adults had already emerged, and most cells were empty.

The nest consisted of 95 mud cells arranged in an elongate mass along the
curved inner wall of the palm frond base (Fig. la, b). Each cell was ovoid
with one flattened end, which usually contained the exit hole of the emerging
wasp. The external dimensions of the cells were 13-16 mm by 10-11 mm
(Fig. Id). The cell walls were smooth and unlined inside and rough exter¬
nally, 0.5-1.0 mm thick. The bottom of the cell was not completely covered
with mud medially (Fig. lc).

The cocoons were papery and ranged from pale yellow to dark brown

VOLUME 56, NUMBER 2

13-16mm

10-11 mm

Fig. 1. Communal nest of Auplopus esmeralda: a, nest diagram; b, nest in situ; c, cross
section of cell; d, dorsal view of cell.

depending on their age. Each cocoon was attached to the wall of the cell
opposite the flattened end.

At least 18% of the cells in this nest were parasitized. Fungus accounted
for at least five cells and two cells had well developed Corticeps fruiting
bodies. Other parasites included one unidentified muscoid fly, and a female
Ephuta sp. (Mutillidae) reared from one cell.

Two other nests of this species were collected from traps which were
made from sections of 10 cm diameter bamboo cut with the joint walls
forming end walls and a 1 cm diameter hole bored in the middle. One nest
had three cells and three adult Auplopus were reared from these, one male
and two females. The other nest consisted of two cells and two females
were reared from these. The construction of these cells was the same as the
cells in the palm frond.

Two females of this species were observed collecting mud from a Nasu-
totermes sp. termite nest on the side of a dead tree, one in July 1977 and
the other in August 1978. Each female apparently secreted saliva on a small
area on the mud wall of the termite nest. After wetting the material the
female worked it with her mandibles into a ball almost the size of her head.

100

PAN-PACIFIC ENTOMOLOGIST

She flew away from the tree with the ball held beneath the mandibles,
apparently supported with the palpi.

The following observations were also made on Barro Colorado Island:

Mystacagenia elegantula Evans. The type female was observed collect¬
ing mud from the same Nasutotermes nest as the Auplopus females.

Priochilus splendidulum splendidulum (Fabricius). A female was col¬
lected carrying a spider, Ctenus sp. (Ctenidae), along the top of a log. The
spider’s legs were not amputated.

Priocnemella rufothorax (Banks). One female was collected carrying the
spider, Acanthoctenus sp. (Acanthoctenidae), in the leaf litter. The wasp
had not amputated the spider’s legs.

Ageniella anconis Banks. This female was collected while she was ‘rest¬
ing’ on a blade of grass. She was carrying the spider, Sarinda sp. (Clubio-
nidae) by the spinnerette and had not amputated the spider’s legs.

Acknowledgments

I would like to thank the Smithsonian Tropical Research Institute for
allowing me to stay and work at their field station on Barro Colorado Island,
Panama. Also I would like to thank Drs. Marius S. Wasbauer and Howard
E. Evans for identifying the pompilid wasps, Willis Gertsch for identifying
the spiders and Richard M. Bohart and Marius Wasbauer for comments and
criticisms.

Literature Cited

Evans, H. E. 1966. A revision of the Mexican and Central American spider wasps of the
subfamily Pompilinae. Mem. Amer. Entomol. Soc., 20:1-442.

Evans, H. E. 1973. Studies on neotropical Pompilidae, IX: the genera of Auplopodini. Psyche,
80:212-226.

Williams, F. X. 1919. Philippine wasp studies: descriptions of new species and life history
studies. Exp. Sta. Hawaiian Sugar Planters Assoc. Entomol. Ser., 14:19-186.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 101-104

A NEW OXYETHIRA (HYDROPTILIDAE, TRICHOPTERA) OF THE
AEOLA ROSS GROUP; WITH A KEY TO SEPARATE THE
FIVE MALES OF THE GROUP 1

R. L. Blickle

University of New Hampshire, Durham 03824

Oxyethira allosi, new species

Holotype, male. —3.3 mm long, specimens from the type series vary from
3.0 to 3.5 in length. Genitalia, Fig. 1A, B. Seventh sternal process short,
thin, pointed. Eighth tergite deeply and widely incised, sides of the incision
regular. Ninth tergite, the projecting rods apparent in the other species of
the group are lacking; subgenital plate arcuate, deeply pigmented, emargin-
ate apically, the apico-lateral corners rounded. Claspers below subgenital
plate round, pigmented; a short cylindrical process arises near the clasper
base, process tipped with a seta. Aedeagus, Fig. lae, long tubular the tip
membranous, expanded; a curved structure extends into the membranous
part; aedeagal length 0.5 mm.

Holotype, male. —Oroville, Butte Co., California, 1-24-1975 T. D. Eichlin
and T. Kono collectors. The holotype was taken on concrete fish ladders
at fish hatchery. Paratypes, 18 males, same data as above. One male, Sunol
Regional Park, Alameda Co., California, X-15-1974, D. G. Denning collec¬
tor. 14 males, Mendocino Co., California, VI-24-1978, D. G. Denning col¬
lector, creek at Brook Trail near Willits. Four males, Nimrod Warm Springs,
Route 1-90, Granite Co., Montana, V-10-1970, D. S. Potter collector. One
male, Goshen, Utah Co., Utah, 1-27-1973, R. N. Winget collector.

The paratype from Utah was taken at springs and ponds 1.5 km east of
Goshen from a slow running stream (about 10 m in length) connecting two
small ponds each less than half a hectare in extent. These ponds are fed by
warm water springs and are characterized by heavy growths of submerged
and floating vegetation. The connecting stream has a sand-gravel-silt sub¬
strate with significant amounts of water cress and Chara and other algae.

Allotype, female. —Fig. IF, same size and color as male. Two sclerotized
plates exterior to the cerci, two small setae between cerci. Ninth sternite
rounded, covered with setae on anterior part. Bursa copulatrix with three
“chambers.” Fig. lbc. There appears to be an inverted Y structure at the
base.

Allotype, female .—Mendocino Co., California, creek at Brook Trail near
Willits, VI-24-1978, D. G. Denning collector.

102

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Oxyethira allosi n.sp. Genitalia; A, male, lateral; B, male dorsal; ae, aedeagus,
dorsal; F, female, ventral; be, bursa copulatrix. Line = 0.25 mm. Fig. 2. Oxyethira ahacatia
Denning; A, male, lateral; ae, aedeagus. Fig. 3. Oxyethira anabola Blickle; B, male, dorsal;
ae, aedeagus; ael, apex of aedeagus lobes expanded. Fig. 4. Oxyethira barnstoni Harper; B,

VOLUME 56, NUMBER 2

103

The two plates and the two small setae near the cerci plus the three
chambered bursa sets this apart from related species.

Deposition of type material .—Holotype male, allotype female and 13 male
paratypes to the California Academy of Science, San Francisco, California.
Two male paratypes to California State Department of Agriculture, Insect
Taxonomy Laboratory, Sacramento, California. Remaining paratype males
to Dr. D. G. Denning, Moraga, California.

This species belongs to a group of hydroptilids characterized by the males
having an arcuate subgenital plate, a large tubular aedeagus having no spiral
process, and (usually) rods projecting posteriorly from the 9th tergite.

This species differs from abacatia Denning (1947) and anabola Blickle
(1966) by a lack of aedeagal lobes, from barnstoni Harper (1976) by having
the incision of the 8th tergite deep and from aeola Ross (1938) by having
the sides of the 8th tergal incision regular, in addition allosi n. sp. lacks rods
projecting posteriorly from the 9th tergite as possessed by the others of the
group. The subgenital plate posteriorly is also more irregular than those of
the other species.

The known geographic distribution of the five described species of the
aeola group is: Oxyethira abacatia Denning, Florida, Georgia; O. aeola
Ross, British Columbia to Minnesota: O. anabola Blickle, New Jersey to
Quebec to Minnesota; O. barnstoni Harper, Quebec; O. allosi n. sp., Cal¬
ifornia, Montana, Utah.

Key to Males of aeola Ross Group

1. Apex of aedeagus 3 or more lobes (Figs. 2, 3). 2

Apex of aedeagus simple or expanded membranous mass .... 3

2. Apex of aedeagus 3 lobed; rods of 9th segment do not extend beyond

subgenital plate (Fig. 2). abacatia Denning

Apex of aedeagus 4 lobed; rods of 9th segment extend beyond
subgenital plate (Fig. 3). anabola Blickle

3. Incision of 8th tergite narrow (Fig. 4) . barnstoni Harper

Incision of 8th tergite broad, deep . 4

4. Sides of 8th tergite incision irregular (Fig. 5) . aeola Ross

Sides of 8th tergite incision regular (Fig. 1) . allosi n.sp.

The key above replaces the couplets 18 to 19, pages 39-40 of the Oxy-

male, dorsal; ae, aedeagus. Fig. 5. Oxyethira aeola Ross; B, male, dorsal; ae, aedeagus. Fig.
2, redrawn from Denning (1947); Fig. 3, redrawn from Blickle (1966); Fig. 4, redrawn from
Harper (1976); Fig. 5, redrawn from Ross (1938).

104

PAN-PACIFIC ENTOMOLOGIST

ethira key in the Hydroptilidae (Trichoptera) of America North of Mexico,
Blickle (1979).

Literature Cited

Blickle, R. L. 1966. A new Hydroptilidae (Trichoptera). Entomol. News, 67:185-187.
Blickle, R. L. 1979. Hydroptilidae (Trichoptera) of America North of Mexico. New Hamp.
Agr. Exp. Sta. Bull. 509, 98 pp.

Denning, D. G. 1947. Hydroptilidae from Southern United States. Can. Entomol., 79:12-20.
Harper, P. P. 1976. Oxyethira barnstoni n.sp. un noveau trichoptere de Radisonie Quebec
(Hydroptilidae). Ann. Entomol. Soc. Quebec, 21:35-38.

Ross, H. H. 1938. Descriptions of Nearctic Caddisflies with special reference to Illinois
species. Ill. Natr. Hist. Surv. Bull., 21:101-183.

Footnote

1 Published with the approval of the New Hampshire Agricultural Experiment Station as
Scientific Contribution No. 962.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 105-106

FLEAS FROM SOME ALASKAN BIRDS
(SIPHONAPTERA)

Glenn E. Haas, Tim Rumfelt

State of Alaska Department of Health and Social Services,
Environmental Health Section, MacKay Building,

338 Denali Street, Anchorage 99501

Richard E. Barrett

United States Department of Agriculture, Animal, Plant Health
Inspection Service, Veterinary Service, Box 720,

Palmer, Alaska 99645

and

Nixon Wilson

Department of Biology, University of Northern Iowa, Cedar Falls 50613

Records of fleas collected from birds are much fewer than records from
mammals. Not as many birds are examined for fleas, birds are seldom in¬
fested, and in Alaska species of mammal fleas outnumber species of bird
fleas 2 to 1. By thoroughly examining dead birds found by chance since
1973, we obtained four fleas. Three of these were true bird fleas, one being
a new record for the United States. The fourth was a mammal flea not
previously recorded from birds. The specimens are deposited in the collec¬
tion of the senior author. We thank F. G. A. M. Smit, London, for his
comments on Ceratophyllus adustus and Dr. G. P. Holland, Ottawa, for his
comments on C. diffinis.

Ceratophyllus adustus Jordan

Record. —Palmer, 8.8 km SE: 1 female, Picoides tridactylus, 27.X.75, G.
E. Haas.

No records of this species have been published since Jordan (1932) de¬
scribed it from a single female collected from a porcupine in the Atlin region
of British Columbia. He noted the reduced chaetotaxy and concluded that
it presumably resulted from a bird flea changing hosts, the porcupine prob¬
ably becoming the true host. Our studies in Alaska indicate, however, that
C. adustus is a woodpecker nest flea. The infested porcupine probably
climbed a tree in which a woodpecker had nested and approached the nest
close enough for the flea to make contact.

106

PAN-PACIFIC ENTOMOLOGIST

Ceratophyllus diffinis Jordan

Record .—Palmer, 4 km E: 1 female, Ixoreus naevius, 2.V.76, I. Sims.
This Nearctic flea is transcontinental on a wide variety of birds, especially
Galliformes and Passeriformes (Holland, 1963; Benton & Shatrau, 1965).
Holland (in litt. 1975) has specimens from I. naevius in Alaska.

Ceratophyllus gallinae (Schrank)

Record .—Talkeetna, 19.2 km S: 1 male, Dendroica coronata, 1.VI.76,
G. E. Haas.

This widespread Old World flea is a natural passerine parasite that became
a domestic fowl pest in Europe and supposedly was introduced into North
America (Holland, 1963). Holland (1949) recorded it ranging from New
Brunswick to Saskatchewan and later (Holland, 1963) from western Alaska,
one of several localities in which Smit (1969) considered it an introduction.

Tarsopsylla octodecimdentata coloradensis (Baker)

Record .—Matanuska Glacier, 1 km N: 1 female, Perisoreus canadensis,
27.IX.78, T. Rumfelt.

This is a tree squirrel nest flea (Holland, 1963). The jay might have become
infested while feeding on a roadkill red squirrel ( Tamiasciurus hudsonicus )
or while visiting a tree hole frequented by this mammal. This is the first
record of T. o. coloradensis parasitizing a bird.

Literature Cited

Benton, A. H., and V. Shatrau. 1965. The bird fleas of eastern North America. Wilson Bull.,
77:76-81.

Holland, G. P. 1949. The Siphonaptera of Canada. Canada Dept. Agr. Tech. Bull. No. 70,
306 pp.

Holland, G. P. 1963. Faunal affinities of the fleas (Siphonaptera) of Alaska with an annotated
list of species. Proc. 10th Pacific Sci. Congr. (1961), 45-63.

Jordan, K. 1932. Siphonaptera collected by Mr. Harry S. Swarth at Atlin in British Columbia.
Novit. Zool., 38:253-255.

Smit, F. G. A. M. 1969. A catalogue of the Siphonaptera of Finland with distribution maps
of all Fennoscandian species. Ann. Zool. Fenn., 6:47-86.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 107-110

OCCURRENCE OF METAPHYCUS BARTLETTI ANNECKE AND
MYNHARDT, A SOUTH AFRICAN PARASITE OF
BLACK SCALE, SAISSETIA OLEAE (OLIVIER)

IN CENTRAL AND NORTHERN CALIFORNIA
(HYMENOPTERA: ENCYRTIDAE;

HOMOPTERA: COCCIDAE)

C. E. Kennett

Division of Biological Control, University of California, Berkeley 94720

From 1976 through 1978 I conducted periodic field surveys to assess the
natural enemy complex of the black scale, Saissetia oleae (Olivier), in cen¬
tral and northern California where the scale is a sporadic economic pest in
commercial olive groves and on several ornamental trees and shrubs. The
surveys were made in conjunction with a continuing program of periodic
colonization of parasites newly introduced from the western Cape Province,
South Africa, where S. oleae is thought to have originated (De Lotto, 1976).

Of special interest among the several parasite species recovered during
the surveys was the presence of Metaphycus bartletti Annecke and Myn-
hardt, an African encyrtid not recorded previously from central and north¬
ern California.

Although the reports on colonization of beneficial organisms in California
by the Division of Biological Control, University of California, Riverside,
for the years 1958-1960, show that an encyrtid then identified as Metaphy¬
cus inviscus Compere was colonized on S. oleae infestations in six southern
California counties (Fig. 1), it was not reported to have been released in
central or northern California. The stock colonized as M. inviscus was orig¬
inally obtained from S. oleae infesting olive near Stellenbosch, South Africa
by B. R. Bartlett in 1958. Regarding colonization of the parasite, Bartlett
(see Annecke and Mynhardt, 1972) stated, “recoveries were abundant the
first year of release but it was not taken thereafter . . . Annecke and
Mynhardt (1972) studied specimens of the stock colonized in southern Cal¬
ifornia and concluded that they differed from M. inviscus as described by
Compere (1940). The California material was then described as M. bartletti
by Annecke and Mynhardt (1972) who illustrated the cryptic characters
which separate the two species.

During the 1976-1978 surveys various host plants of S. oleae (principally
olive) were examined at 225 sites in 18 central and northern California coun¬
ties. Light to heavy infestations of S. oleae were sampled at 73 sites. At all
other sites the scale was either absent or too scarce to permit sampling.

108

PAN-PACIFIC ENTOMOLOGIST

Table 1. Recoveries of Metaphycus bartletti from black scale samples according to counties
during 1976-1978.

County

No. sample sites

Parasite recov¬
eries—no. sites

No. parasites

Alameda 3

107

Butte

Contra Costa 3

167

Fresno

204

Glenn

Madera

Marin 3

Merced

Placer

Sacramento

San Joaquin

San Luis Obispo 3

Solano

Stanislaus

937

Sutter

Tehama

Tulare

Yolo

1577

a Coastal counties.

Results

As shown in Table 1, M. bartletti was recovered at 27 (37%) of the 73
sample sites. The parasite was recovered from 18 sites in 10 Central Valley
counties and from 9 sites in 3 coastal counties. Thirty-one percent of the
samples from the Central Valley counties and 53% of the samples from
coastal counties yielded M. bartletti. Chronologically, M. bartletti was re¬
covered at 7 of 15 sites in 1976, at 4 of 12 sites in 1977, and at 16 of 46 sites
in 1978.

The frequency of recovery for M. bartletti (27 sites—13 counties) was
second only to that for M. helvolus (Compere), another African species
which was recovered at 42 sites in 17 counties. Total recoveries of M.
bartletti (1577) were also second to those for M. helvolus (2477). Together
they represented 71.2% of all parasite recoveries. Coincidental recoveries
of M. bartletti and M. helvolus occurred at 20 sites. At these sites recoveries
of M. helvolus were numerically superior to M. bartletti at 14 sites whereas
M. bartletti was dominant in numbers at 6 sites.

Laboratory emergence records for M. bartletti showed that this species
develops gregariously in nearly mature (rubber stage) scales. A lot of 50

VOLUME 56, NUMBER 2

109

Fig. 1. Distribution of Metaphycus bartletti in central and northern California. Solid circles
indicate areas where M. bartletti was recovered during 1976-1978 surveys. Open circles in¬
dicate areas where M. bartletti was colonized during 1958-1960.

mummified S. oleae (collected at Oakdale, Stanislaus County) yielded an
average of 4 parasites/host (range, 2-8).

The wide distribution of M. bartletti in central and northern California
(Fig. 1) suggests that the parasite has probably been expanding its range
over a number of years, very likely throughout the period since it was first
colonized in southern California in 1958-1960. The fact that Bartlett’s re¬
leases some twenty years ago were confined to southern California localities
suggests that M. bartletti is a recent arrival in central and northern Cali¬
fornia. Also, earlier sporadic samples of S. oleae taken from various local¬
ities in central and northern California by the author during the 1950’s did
not reveal the presence of Metaphycus species similar to bartletti or invis-

PAN-PACIFIC ENTOMOLOGIST

cus. Thus the possibility that M. bartletti became established in central and
northern California prior to 1958 by some unexplained means is not indi¬
cated.

The source of colonizing stocks of M. bartletti responsible for its estab¬
lishment in central and northern California is uncertain. Of several possible
sources, transport by man of live plant material infested with parasitized
scale hosts appears to afford the most logical explanation. Although rela¬
tively rapid natural dispersal of parasitic Hymenoptera is known to exist
(DeBach, 1974), the relatively barren mountain ranges between southern
and central California (Fig. 1) provide a formidable barrier to natural mi¬
gration. While there remains a possibility that M. bartletti was released in
central California during 1958-1960, this seems unlikely since it is a policy
of the Biological Control divisions of the University of California to report
all colonizations of beneficial organisms.

Acknowledgments

I thank Dr. D. P. Annecke, Plant Protection Research Institute, Pretoria,
South Africa, who made the initial identification of Metaphycus bartletti
from material recovered during the early stages of the surveys. I also thank
Dr. K. S. Hagen, who reviewed the manuscript and Mr. J. Hamai, who
made the figure.

Literature Cited

Annecke, D. P. and M. J. Mynhardt. 1972. The species of the insidiosus -group of Metaphycus
Mercet in South Africa with notes on some extra-limital species (Hymenoptera-
Encyrtidae). Revue Zool. Bot. Afr., 85(3-4):227-274.

Compere, H. 1940. Parasites of the black scale, Saissetia oleae in Africa. Hilgardia, 13(9):
387-425.

DeBach, P. 1974. Biological control by natural enemies. Cambridge Univ. Press, London.
323 pp.

DeLotto, G. 1976. On the black scales of southern Europe (Homoptera: Coccoidea: Coccidae).
J. Ent. Soc. Sth. Afr., 39:147-149.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 111-112

ANTI-PREDATOR BEHAVIOR IN SWARMS OF RHAGOVELIA OBESA

(HEMIPTERA: VELIIDAE)

G. Scott Deshefy

Department of Zoology, Clemson University, Clemson, SC 29631

The broad-shouldered water strider or riffle bug, Rhagovelia obesa (Uhl-
er, 1871), is an aquatic hemipteran commonly found along edges of lotic
habitat. Its prey principally consists of small insects and crustaceans,
trapped within the water’s surface film. The median legs, equipped with fan¬
shaped, setal plumes at apical segments of the tarsi, provide locomotion,
beating alternately beneath the water surface (Bueno, 1907; Coker et al.,
1936; Cheng and Fernando, 1971). Rhagovelia obesa is active in low tem¬
peratures and undergoes four larval stages (Cheng and Fernando, 1971).
First instar nymphs have been observed in late May in South Carolina.
Adults have been observed in November.

Behavioral experiments were conducted on Rhagovelia obesa popula¬
tions on 18-mile Creek in Pendleton, South Carolina. Both nymphs and
adults were observed to aggregate into swarms, disperse when disturbed
and eventually reassociate. Insects were observed in groups ranging from
4 to 80 individuals. Mating pairs were occasionally included in swarms and
were observed as early as July.

Ten large swarms were investigated at midday along various points of 18-
mile Creek. Experiments were conducted during sunny days over a two
week period using sub-surface and overhead “approach models” to stim¬
ulate dispersal. Sub-surface models, resembling salmonid dorsal silhouettes,
consisted of metal ovals, approximately 25 cm x 6 cm and painted flat black
and dark green. Sub-surface models were pulled by monofilament line along
the sandy creek bottoms and were slowly drawn upstream towards riffle
bug swarms.

Overhead models were constructed from crow and blue jay skins,
stretched across 15 cm x 8 cm x 5 cm wooden frames. These models were
suspended from fishing line and lowered by bamboo pole directly above
Rhagovelia obesa swarms. Twenty trials at five minute intervals were per¬
formed for each experiment and for each insect group. Riffle bug responses
to shadow movements and water surface disturbances were also noted.

Both moderate and rapid introductions of overhead models resulted in
immediate dispersals of tested swarms. The same reactions occurred when
gentle, irregular wave action was generated by randomly splashing a wooden
ladle within a few feet of the insects. Both experiments revealed that, fol-

112

PAN-PACIFIC ENTOMOLOGIST

lowing 8 to 10 trials, repeated disseminations led to reduced swarm cohesion
and partial extinction of dispersal behavior. Aggregates also exhibited small
increases in time elapsed prior to regrouping.

However, insect swarms, approached by slowly moving shadows cast on
the streambed, maintained tight formations and avoided the projections with
coordinated group movements. Riffle bug swarms also behaved as organized
units while moving in advance of and at the same speed as approaching sub¬
surface models.

The described results suggest that swarms of Rhagovelia obesa, which
may facilitate mate availability, could also function as an indirect deimatic
anti-predator defense. Coordinated avoidance of sub-surface images appears
to function as a primary defense mechanism. Supplementing this behavior,
riffle bug swarms, dispersing at the instant of attack, may further create a
secondary “flash” stimulus. Such rapidly scattering assemblies of insects
could effectively distort or obliterate search images of approaching fish or
fowl.

Acknowledgment

I wish to thank the Deshefy-Sekora clan.

Literature Cited

Bueno, J. R. de la Torre. 1907. On Rhagovelia obesa, Uhler. Can. Entomol., 39:61-64.
Cheng, L., and C. H. Fernando. 1971. Life history and biology of the riffle bug, Rhagovelia
obesa Uhler (Heteroptera: Veliidae) in Southern Ontario. Can. J. Zool., 49:435-442.
Coker, R. E. et al. 1936. Swimming plume and claws of the broad-shouldered water-strider,
Rhagovelia flavicinta Bueno (Hemiptera). Bull. Brooklyn Entomol. Soc., 31:81-84.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 113-120

REVISION OF THE GENUS STENOBATYLE CASEY
(COLEOPTERA: CERAMBYCIDAE)

John A. Chemsak

201 Wellman Hall, Department of Entomology,
University of California, Berkeley 94720

The purpuricenine genus Stenobatyle was proposed by Casey (1912) to
include Entomosterna prolixa Bates and a new species, cribrata, from Mex¬
ico. Linsley (1935) described the species inflaticollis in a new genus, Lep-
tobatyle, observing its close resemblance to cribrata Casey. The remaining
species herein placed into Stenobatyle were in the genus Entomosterna
Chevrolat until Chemsak and Linsley (1974) transferred them into Paraba-
tyle Casey. Subsequent studies indicate that the type species of Parabatyle
is not congeneric with the other species. Therefore, sanguiniventris Chev¬
rolat is retained as Parabatyle and the others, including inflaticollis, placed
into Stenobatyle.

The species of Stenobatyle are all presently known from Mexico, with
one species, eburata (Chevrolat), ranging to Costa Rica. Adults are diurnal
and are found on the flowers of various trees and shrubs. No host plant data
have been seen for any of the species.

Genus Stenobatyle Casey

Stenobatyle Casey, 1912:326; Linsley, 1935:101; Chemsak and Linsley,

1974:182 (synonymy).

Leptobatyle Linsley, 1935:101; Chemsak and Linsley, 1974:182 (synonymy).

(Type: Leptobatyle inflaticollis Linsley, monobasic).

Form moderate sized, slender. Head small, front oblique; palpi short,
apical segments short, truncate at apices; mandibles short, slightly curved
at apices, apices acute; eyes finely faceted, deeply emarginate, upper lobes
small; antennal tubercles divergent, moderately elevated; antennae slender,
eleven-segmented, slightly longer than body in males, third segment usually
longer than scape, fourth shorter than third. Pronotum convex, sides round¬
ed; apex narrower than base; apex and base narrowly margined, hind mar¬
gins usually flaring laterally; prosternum not or very feebly impressed, in-
tercoxal process abruptly declivous, not expanded at apex, coxal cavities
broadly open behind; mesosternum with intercoxal process arcuate to de¬
clivous, coxal cavities open to epimeron; metasternum with episternum sub¬
parallel to slightly tapering posteriorly. Elytra slender, tapering, lateral mar-

114

PAN-PACIFIC ENTOMOLOGIST

gins feebly to strongly impressed at basal V3; eburneous ridges or elevated
costae usually present; apices crenulate, outer angles dentate. Legs slender;
hind femora linear, arcuate, usually extending to apices of elytra; posterior
tarsi slender, usually elongate, third segment cleft to middle. Abdomen nor¬
mally segmented.

Type species: Entomosterna prolixa Bates (by original designation).

This genus may be recognized by the slender form, small head, relatively
short antennae, convex, usually tapering pronotum with distinct margins at
the base and apex, and usually strongly costate or eburneous ridged elytra.

Key to the Species of Stenobatyle

1. Elytra with very distinct longitudinal, often eburneous costae .. 2

Elytra lacking elevated costae, opaque, very coarsely, confluently
punctate; pronotum dull, closely alveolate-punctate; integu¬
ment black, pronotum reddish to orange except for middle.

Length, 13 mm. Mexico and Guerrero . inflaticollis

2(1). Elytra with costae eburneous . 3

Elytra with costae not eburneous. 4

3(2). Each elytron with a single broad costa, punctures rather fine,
subequal to those of pronotum. Length, 9-17 mm. Sonora,

Mexico to Costa Rica. eburata

Each elytron with 2 costae, one dorsal and one extending back
from humerus, punctures coarse, contiguous, much larger than

those of pronotum. Length, 10-11 mm. Guerrero. gracilis

4(3). Pronotum finely, rather sparsely punctate, punctures finer and
sparser at sides; elytra finely, very sparsely punctate at basal

margin. Length, 15-18 mm. Guerrero to Oaxaca . prolixa

Pronotum coarsely, confluently punctate, punctures transverse;
elytra coarsely, contiguously punctate throughout. Length, 7-
12 mm. Sinaloa to Veracruz and Oaxaca . miniaticollis

Stenobatyle inflaticollis (Linsley), NEW COMBINATION

Leptobatyle inflaticollis Linsley, 1935:101.

Parabatyle inflaticollis, Chemsak and Linsley, 1974:182.

Male .—Form moderate sized, slender; integument black, pronotum ex¬
cept apical and basal margins and a median vitta or spot reddish. Head
confluently punctate, sparsely pubescent; antennae extending beyond elytra
by four segments, scape finely, confluently punctate, segments from sixth
longitudinally carinate beneath, outer segments minutely, densely pubes¬
cent, third segment much longer than scape, fourth a little shorter than third,
fifth equal to fourth. Pronotum a little broader than long, sides broadly

VOLUME 56, NUMBER 2

115

rounded; disk strongly convex, alveolate-punctate with a vague median lin¬
ear callus behind middle; pubescence obsolete; prosternum opaque, sparse¬
ly pubescent; meso- and metasternum finely, densely punctate, densely
clothed with silvery recumbent pubescence, mesosternal process subdecli-
vous. Elytra about 2 Vi times as long as broad, epipleura shallowly emargin-
ate at basal one-third; punctures very coarse, deep, contiguous to confluent;
pubescence obsolete; apices sinuate truncate. Legs slender; hind femora
extending a little beyond apices of elytra; hind tibiae arcuate; hind tarsi
elongate. Abdomen very densely clothed with silvery recumbent pubes¬
cence except narrowly at apices of first four segments; last sternite truncate
at apex. Length, 13 mm.

Female. —Form similar. Antennae a little longer than body. Abdomen
with last sternite broadly truncate at apex. Length, 13 mm.

Type locality. —Bejucos, Temascaltepec, Mexico, Mexico.

Range. —Mexico and Guerrero.

Flight period. —July.

The punctation of the pronotum and elytra will distinguish this species.
The black median line of the pronotum varies from a spot over the basal
one-half to extending the entire length. The female specimen at hand has an
indication of narrow, pale sub-marginal vittae on the elytra.

New records. —1<3, 12, Canyon del Zopilote, 34 mi N Chilpancingo,
Guerrero, Mexico, 11 July 1970 (Fisher and Sullivan).

Stenobatyle eburata (Chevrolat), NEW COMBINATION

Entomosterna eburata Chevrolat, 1862:755; Bates, 1880:85.

Parabatyle eburata, Chemsak and Linsley, 1974:182.

Entomosterna trucidata Chevrolat, 1862:755; Bates, 1880:86; 1885:330.

NEW SYNONYMY.

Parabatyle trucidata, Chemsak and Linsley, 1974:182.

Entomosterna unicostata Casey, 1912:332. NEW SYNONYMY.

Male. —Form moderate sized, tapering; integument black, pronotum
often reddish at sides, each elytron with a broad, glabrous, elevated ebur-
neous costa. Head densely, irregularly punctate, sparsely pubescent; anten¬
nae slightly longer than body, scape densely, moderately coarsely punctate,
outer segments opaque, densely clothed with very short, recumbent pubes¬
cence, scape shorter than third segment, fourth equal to scape, fifth equal
to third. Pronotum about as long as broad, sides rounded; disk strongly
convex, broadly impressed behind middle at base, usually with a shallow,
irregular callus before depression; punctures moderately coarse, contig¬
uous, becoming confluent at sides; pubescence rather sparse, depressed,
sides with a few, long, erect hairs; prosternum rugulose, densely pale pu-

116

PAN-PACIFIC ENTOMOLOGIST

bescent; meso- and metasternum finely, densely punctate, densely clothed
with recumbent silvery pubescence, mesosternal process moderately decli¬
vous. Elytra more than 2 Vi times as long as broad, sides shallowly impressed
at middle; eburneous costae sutural, extending from basal margin almost to
apex; basal punctures separated, becoming coarser and contiguous almost
to apex; pubescence short, depressed, rather sparse, with a few erect hairs
near base; apices bi-emarginate or bitruncate, angles usually dentate. Legs
slender; hind femora extending to apices of elytra or a little beyond; hind
tarsi elongate. Abdomen minutely, densely punctate at sides, densely pu¬
bescent at sides; last sternite truncate at apex. Length, 9-16 mm.

Female. —Form more robust. Antennae about as long as body. Abdomen
with last sternite broadly truncate. Length, 10-17 mm.

Type locality. —of eburata, Soleda, Yucatan, Mexico; trucidata, Merida,
Yucatan; unicostata, Guerrero, Mexico.

Range. —Sonora, Mexico to Costa Rica (Fig. 1).

Flight period. —May to August.

Flower records. — Croton, Buddleia, Jatropha, Donneilsmithia.

This species is easily recognized by the broad, eburneous costa on each
elytron.

From Sinaloa, Mexico to Veracruz and Yucatan, all specimens have the
pronotum uniformly black. From Oaxaca and Yucatan to Costa Rica all
specimens at hand have the pronotum reddish at the sides. There appear to
be no other differences but the southern population may eventually prove
to represent a distinct subspecies, trucidata.

Adults are common in western Mexico on flowers of Croton, Buddleia
and Jatropha during July and August. In Costa Rica, adults emerge earlier,
in May. This species is usually found on flowering plants together with
miniaticollis.

Most records are from Mexico (Fig. 1). Others include: Chontales, Nic¬
aragua (BCA); 8 mi NW Bagaces, Guanacaste, Costa Rica.

Stenobatyle gracilis, new species

Male. —Form moderate sized, slender; integument black, pronotum ex¬
cept apical and basal margins reddish, each elytra with a yellowish subsu-
tural and submarginal costa. Head finely, confluently punctate, sparsely
pubescent; antennae extending beyond elytra by about two segments, scape
confluently punctate, outer segments densely clothed with fine, dark, ap-
pressed pubescence, third segment much longer than scape, fourth segment
about one-third longer than scape, fifth equal to fourth. Pronotum as long
as broad, sides broadly rounded; disk with a shallow, elongate, longitudinal,
glabrous callus at middle; punctures coarse, shallow, almost alveolate; pu¬
bescence obsolete; prosternum rugulose, sparsely pubescent; meso- and

VOLUME 56, NUMBER 2

117

Fig. 1. Known distribution of Stenobatyle eburata (Chevrolat) in Mexico.

metasternum minutely, densely punctate at sides, coarsely and more sparse¬
ly to glabrous midline of mesosternum, pubescence silvery, dense at sides.
Elytra more than three times as long as broad, epipleura moderately emar-
ginate at basal one-third; subsutural costae moderately broad, sub-marginal
pair narrow; punctures coarse, contiguous, larger at middle; pubescence
obsolete; apices unevenly truncate. Legs slender; hind femora extending to
apices of elytra, rather sparsely punctate; hind tarsi moderately elongate.
Abdomen minutely, densely punctate, densely clothed with short appressed,
silvery pubescence; last sternite truncate at apex. Length, 10 mm.

Female .—Form similar. Antennae slightly longer than body. Abdomen
with last sternite broadly truncate. Length, 10 mm.

Holotype male (California Academy of Sciences) from Mexcala, Guer¬
rero, Mexico, 29 June, 1951 (P. D. Hurd). One female paratype from Canyon
del Zopilote, 24 mi N Chilpancingo, Guerrero, 11 July, 1970 (Fisher, Sulli¬
van).

This species is easily recognized by the color and small, slender form.
Although the type series possesses double yellowish vittae on each elytron,
it is probable that other individuals would lack one or both of these. How¬
ever, the costae should remain evident.

118

PAN-PACIFIC ENTOMOLOGIST

Stenobatyle prolixa (Bates)

Entomosterna prolixa Bates, 1892:180; Chemsak, 1967:76 (lectotype).
Stenobatyle prolixa, Linsley, 1935:102.

Male. —Form moderate sized, elongate; integument shining, black, prono-
tum bright red except narrowly at apical and basal margins, femora often
reddish. Head finely, irregularly punctate, pubescence very short; antennae
extending about three segments beyond elytra, scape finely, densely punc¬
tate; segments densely clothed with very fine, dark, appressed pubescence,
third segment longer than scape, fourth equal to scape, fifth equal to third.
Pronotum about as long as broad, sides broadly rounded; disk with a me¬
dian, glabrous callus behind middle; punctures fine, somewhat transverse,
densest at middle, becoming very shallow toward sides; fine setae rising out
of punctures depressed, transversely directed toward center of disk; pro¬
sternum transversely rugulose, densely clothed with short, erect, pale pu¬
bescence; meso- and metasternum very finely, densely punctate at sides,
densely clothed with appressed, silvery pubescence, mesosternal process
moderately declivous. Elytra almost three times longer than basal width,
sides feebly impressed at middle; each elytron with an elevated, glabrous
costa near suture and vague one extending from humerus to near apex where
it joins with the other; basal punctures fine, sparse, becoming denser and
coarser toward apex; pubescence short, sparse; apices sinuate-truncate.
Legs slender; hind femora extending to apices of elytra; pubescence fine;
hind tarsi elongate. Abdomen finely, densely punctate and pubescent at
sides; last sternite truncate at apex. Length, 15-16 mm.

Female. —Form more robust. Antennae about as long as body. Abdomen
with last sternite broadly subtruncate at apex. Length, 15-18 mm.

Type locality. —Guerrero, Mexico.

Range. —Guerrero to Oaxaca.

Flight period. —July and August.

The finely punctate, reddish pronotum and sparse punctures at the base
of the elytra will separate this species from miniaticollis.

New records. —Mexico: 35 5, 29 9, Acapulco, Guerrero, 9-14 July, 10
August, 1936 (C. Seevers); 19, Puerto Angel, Oaxaca, 15 July, 1964 (A. B.
Lau); lcJ, 23 mi S Matias Romero, Oaxaca, 14 August, 1963 (Parker,
Stange); 1 9,56 mi NW Tehuantepec, Oaxaca, 27 July, 1963 (W. A. Foster).

Stenobatyle miniaticollis (Chevrolat), NEW COMBINATION

Entomosterna miniaticollis Chevrolat, 1862:756; Bates, 1880:86; 1885:330.
Parabatyle miniaticollis, Chemsak and Linsley, 1974:182.

Stenobatyle cribrata Casey, 1912:331; Linsley, 1935:102; Chemsak and

Linsley, 1974:182 (synonymy).

VOLUME 56, NUMBER 2

119

Fig. 2. Known distribution of S. miniaticollis (Chevrolat).

Male .—Form moderate sized, slender; integument black, pronotum red¬
dish, usually with a longitudinal, median black spot. Head densely, irregu¬
larly punctate, very finely pubescent; antennae slightly longer than body,
scape moderately coarsely, densely punctate, outer segments opaque,
densely clothed with very short, appressed pubescence, third segment much
longer than scape, fourth segment shorter than scape, fifth slightly longer
than scape. Pronotum about as long as broad, sides broadly rounded, as
broad as elytra at base; disk convex, moderately coarsely, closely punctate
at middle, punctures at sides coarse, confluent; pubescence sparse, de¬
pressed, transverse; prosternum rugulose, rather sparsely pubescent; meso-
and metasternum minutely punctate at sides, rather coarsely at middle, pu¬
bescence depressed, silvery, denser at sides, mesosternal process arcuate.
Elytra about three times as long as broad, epipleura deeply emarginate at
basal one-third; each elytron with a glabrous, elevated, subsutural costa
extending from basal margin almost to apex; pubescence very short; apices
sinuate-truncate. Legs slender; hind femora extending to apices of elytra;
hind tarsi elongate. Abdomen minutely, densely punctate at sides, pubes¬
cence denser at sides; last sternite truncate at apex. Length, 7-12 mm.

Female .—Form similar. Antennae about as long as body. Abdomen with
last sternite broadly truncate. Length, 8-12 mm.

120

PAN-PACIFIC ENTOMOLOGIST

Type locality. —of miniaticollis, Oaxaca, Mexico; cribrata, Guerrero,
Mexico.

Range. —Sinaloa, Mexico to Veracruz and Oaxaca (Fig. 2).

Flight period .—June to September.

Flower records. — Buddleia, Acacia.

The reddish, coarsely punctate pronotum and coarse punctures of the
elytra will distinguish this species.

Adults are numerous on the flowers of Buddleia wrightii in Sinaloa, Mex¬
ico, where they commonly occur with eburata.

Acknowledgments

This study was carried out in conjunction with a National Science Foun¬
dation sponsored project on North American Cerambycidae through Grant
DEB 76-23849 A01. The authorities of the following institutions and indi¬
viduals are gratefully acknowledged for the loan of specimens: American
Museum of Natural History, New York; California Academy of Sciences,
San Francisco; Canadian National Collection, Ottawa; Cornell University,
Ithaca; Essig Museum of Entomology, Berkeley; Field Museum of Natural
History, Chicago; Los Angeles County Museum of Natural History; Texas
A&M University, College Station; University of California, Davis; Univer¬
sity of Kansas, Lawrence; United States National Museum of Natural His¬
tory, Washington, D.C.; and D. Marqua, Los Angeles, CA.

Literature Cited

Bates, H. W. 1879-1885. Biologia Centrali-Americana, Insecta, Coleoptera, 5:1-436, pis. 1-
25.

Bates, H. W. 1892. Additions to the Longicornia of Mexico and Central America, with remarks
on some of the previously described species. Trans. Entomol. Soc. London, 1892:143-
183, pis. 5-7.

Casey, T. L. 1912. Studies in the Longicornia of North America. Memoirs on the Coleoptera,
3:215-386.

Chemsak, J. A. 1967. Lectotype designations of Cerambycidae in the British Museum (Natural
History). J. Kansas Entomol. Soc., 40:73-81.

Chemsak, J. A., and E. G. Linsley. 1974. Reclassification, synonymy, and descriptions of
some North and Central American Cerambycidae. Coleopt. Bull., 28:181-184.
Chevrolat, A. 1862. Revision des genres Eriphus et Mallosoma Serville, du groupe des Cly-
tides et description de trois nouveaux genres dont un doit etre rapporte au groupe des
Callidiites. Ann. Soc. Entomol. France, (4)2:747-763.

Linsley, E. G. 1935. Studies in the Longicornia of Mexico. Trans. Amer. Entomol. Soc.,
61:67-103, 2 pis.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 121-136

REVIEW OF THE NORTH AMERICAN SPECIES OF APOCRYPHA
ESCHSCHOLTZ, WITH A DESCRIPTION OF THE IMMATURE
STAGES OF APOCRYPHA ANTHICOIDES
(COLEOPTERA: TENEBRIONIDAE)

J. T. Doyen and C. Y. Kitayama
Department of Entomology, University of California, Berkeley 94720

The genus Apocrypha was established by Eschscholtz (1831) for A. an-
thicoides Eschscholtz. Subsequently A. dyschirioides was described from
southern California by LeConte (1851) and A. clivinoides from the Owens
Valley, California by Horn (1870). Recently 5 additional species have been
described from Argentina (Kulzer, 1962; Kaszab, 1969). Lacordaire (1859)
placed Compsomorphus Solier (1851) in synonymy under Apocrypha, and
recognized the distinction of the genus by establishing the tribe Apocry-
phini, which has been recognized ever since. Gebien (1942) listed 6 genera
under Apocryphini, including Melytra Pascoe from Tasmania and Diplo-
cyrtus Quedenfeldt from Morocco. However, knowledge of these uncom¬
monly collected beetles has grown by haphazard accretion, and the tribe
will probably require redefinition when adequate material can be compared.
The purpose of this paper is to review the species of Apocrypha occurring
in North America and to describe the immature stages of A. anthicoides so
that the information will be available for an analysis of the higher classifi¬
cation of Tenebrionidae (Doyen and Lawrence, 1979).

Materials and Methods

For morphological comparison adults and larvae were partially dissected,
cleared in hot KOH and then completely dissected under binocular micro¬
scopes. Mouthparts and other sclerotized structures were mounted in glyc¬
erine jelly so that they could be viewed from various perspectives. Defen¬
sive glands and internal parts of the female reproductive tract were cleared,
stained with chlorazol black and mounted in Canada balsam. Larvae were
obtained by confining adults in containers half-filled with slightly moist sand
and providing a variety of grains, dry dog foods and dead insects as food.
Prepupal larvae were removed to petri dishes with a shallow layer of sand
for pupation.

Apocrypha Eschscholtz

Apocrypha Eschscholtz, 1831:13; LeConte, 1851:137; LeConte, 1862:219;

122

PAN-PACIFIC ENTOMOLOGIST

Lacordaire, 1859:433; Horn, 1871:390; LeConte and Horn, 1883:384;
Kulzer, 1962:98; Kaszab, 1969:328.

Compsomorphus Solier, 1851:206; Lacordaire, 1859:433 (synonymy). Type
species: Compsomorphus elegans Solier (monobasic).

Very small beetles with globular prothorax, constricted, petiolate meso-
thorax and long slender legs with clavate femora. Eyes moderate, nearly
round or with slight anterior emargination; antennae slender, filiform ba-
sally, becoming slightly clavate and serrate apically; labrum transversely
subrectilinear with medial processes of tormae transverse; mandibles elon¬
gate, with prominent retinaculum and smooth (nonstriate) mola; maxillae
with galea densely setose, lacinia with 2 large apical teeth and subapical
brush of stout setae; labrum small, exposing ligula and maxillae; tentorium
with sides short, straight, reaching occipital foramen; tentorial bridge
straight (not arched), situated posteriorly. Prothorax globular, rounded,
without defined angles or lateral carinae; procoxal cavities closed externally
and internally. Mesocoxal cavities closed laterally by apposed sterna, tro-
chantin concealed; mesendosternite with short, slender dorsal arms and
stout, horizontal, apically enlarged anterior arms; elytra oval, inflated,
abruptly constricted basally; epipleuron narrow, extending from humerus
to elytral apex. Metanotum membranous; metendosternite with short, broad
stalk, long stout arms without laminae extending to vicinity of elytral artic¬
ulations and terminating in large apical flanges; tendons inserted medially;
wings absent. Abdomen with external membranes between segments 3-5;
abdominal glands small, saccate, without annulations; ovipositor with gon-
ostyles small, subapical; aedeagus with tegmen dorsal.

Type species: Apocrypha anthicoides Eschscholtz (monobasic).

The North American species of Apocrypha can be separated by the fol¬
lowing key.

1. Pronotum with 2-8 long, projecting setae near each lateral margin

of disk; antennae with 3rd segment about 1.3 times longer than 4th ... 2
Pronotum without long, projecting setae; antennae with 3rd segment
subequal to 4th . anthicoides

2. Pronotum with 2 long projecting setae near each lateral border of

disk; elytra without long, projecting setae. clivinoides

Pronotum with 4-8 long projecting setae near each lateral margin of
disk; elytra with scattered long setae projecting from disk, denser
on humeri and declivity. setosa n.sp.

Apocrypha anthicoides Eschscholtz
(Figs. 1-3)

Apocrypha anthicoides Eschscholtz, 1831:13.

Apocrypha dyschirioides LeConte, 1851:137. NEW SYNONYMY.

VOLUME 56, NUMBER 2

123

Figs. 1-3. Apocrypha anthicoides, male genitalic structures. Figs. 1-2, aedeagus, dorsal
and lateral, respectively. Fig. 3, eighth sternite and spiculum.

This common species is easily recognized by the 3 rows of about 6 long,
erect setae on each elytron; by the absence of long, erect setae on the
pronotum; by the barely emarginate eyes; and by the polished lustrous cu¬
ticle. The head, pronotum, elytra and abdominal sternites are relatively
sparsely punctate, with the punctures separated by about 1.5-2 puncture
diameters. The first 3 abdominal sternites are clearly connate, without in¬
cised intersegmental sutures. The 8th sternite of the male is deeply emar¬
ginate (Fig. 3), and the aedeagus abruptly attenuate to an acutely rounded,
upturned apex (Figs. 1-2).

The characters cited by LeConte (1851) and Horn (1870) as distinguishing
A. anthicoides from dyschirioides involved differences in punctation, body
size and color. Color ranges from pale tan (in obviously teneral individuals)
through brown to black or dark brown with paler elytral humeri. Body length
(measured as elytral length plus pronotal length) varies from 1.9-3.1 mm.
The smallest individuals are dark brown to black and are predominantly
from localities near the seacoast. Larger, brownish individuals and those
with pale humeri are predominantly from interior localities. There is con¬
tinuous variation in both color and size, and large collections from many
localities include a range of phenotypes. This pattern of variation appears
to conform to Gloger’s ecogeographic rule, which has been applied to var¬
ious other insects (Mayr, 1963:324).

Horn (1870) stated that the pronotum was longer than broad in A. dys-
chiriodes, broader than long in anthicoides. Measurement of numerous in¬
dividuals showed that the pronotum is always broader than long, with con¬
tinuous variation in both dimensions. Furthermore, the distinctive male

124

PAN-PACIFIC ENTOMOLOGIST

Fig. 4. Distribution of Apocrypha anthicoides (open circles), A. clivinoides (closed circles)
and A. setosa (squares).

genitalic characters (Figs. 1-3) were exhibited by all individuals, indicating
that a single species is represented.

Apocrypha anthicoides is common below 1200 m from northern Mendo¬
cino County and Butte County, south through the Sierra Nevada and the
coast ranges to San Diego County (Fig. 4). A. anthicoides occurs in coastal
sage scrub, chaparral, and about the margins of salt marshes in the coastal
part of its range. Interior populations inhabit deciduous or mixed deciduous-

Figs. 5-7. Apocrypha clivinoides, male genitalic structures. Figs. 5-6, aedeagus, dorsal
and lateral views, respectively. Fig. 7, eighth sternite and spiculum.

coniferous woodland or chaparral habitats, especially in relatively well-
drained situations. Nearly all individuals are collected by Berlese extraction
of leaf or needle litter.

Apocrypha clivinoides Horn
Apocrypha clivinoides Horn, 1870:391.

The more depressed, elongate form and slightly larger size clearly distin¬
guish this species from anthicoides. It is similar to setosa, but slightly
smaller and has only 2 pairs of long projecting setae on the pronotum (4-8
pairs in setosa).

The elytra are densely covered by pale, appressed setae, but lack long,
erect setae. The eyes are slightly emarginate, and the cuticle is very finely
granulate and dull. The head, pronotum and elytra are densely, almost con-
fluently punctate. The anterior 3 abdominal sternites are separated by in¬
cised intersegmental sutures, and the 8th sternite of the male is longer than
broad and shallowly emarginate (Fig. 7). The aedeagus gradually attenuates
to a sharply rounded apex which is not upturned (Figs. 5-6).

Material examined (Fig. 4). —Calif., Alpine Co., 1 mi. E Woodfords, X-30-
1965, N. Ueshima, ex litter Artemsia (9); Fresno Co., Summit Meadow,
VI-4-1911, R. Hopping (1); Inyo Co., Goodale Creek, N. Lone Pine, IV-3-
1953, R. E. Leech (1); 7 mi. SE Benton Crossing, VI-15-1970, M. S. Was-

126

PAN-PACIFIC ENTOMOLOGIST

bauer and F. G. Andrews (1); Waucoba Spr., IV-30-1975, ex litter Pinus
monophylla, F. G. Andrews (1); Nev., Douglas Co., Topaz Jet., VII-19-1975,
ex litter Pinus monophylla, A. R. Hardy, F. G. Andrews (3). The single spec¬
imen from Summit Meadow, Fresno County, is the only record from west
of the Sierran crest. Since the place name Summit Meadow occurs 3 times
in Fresno County, an exact collection locality cannot be specified.

Apocrypha setosa, new species
(Fig. 8)

Body slender, elongate, slightly depressed dorsoventrally, medium brown
to nearly black; cuticle finely granulate, weakly shining or dull. Head sub-
hexagonal in dorsal aspect, anterior epistomal margin straight; eyes with
anterior margin straight or slightly emarginate; ommatidia coarse, number¬
ing about 30 per eye, and set with fine setae at intersections; vertex, frons,
genae and subgenae closely, almost confluently set with punctures about as
large as ommatidia, and densely clothed with long, pale, appressed setae
directed anterad; longer, erect setae located as follows: 1-3 dorsad of each
eye, 1-4 in irregular submedian row on each side of vertex and frons, 1-3
near anterolateral corners of epistoma.

Pronotum with broadly rounded anterior corners, sides nearly parallel in
anterior half, strongly, evenly converging in posterior half; disk slightly
flattened, closely, nearly confluently set with punctures about as large as
ommatidia, and densely clothed with pale, appressed setae directed poste-
rad; each lateral margin set with 4-8 long setae projecting dorsolaterally;
hypomera and sternum coarsely punctate to punctatorugose, with setal
clothing less dense than on disk; sternum bearing about 10-15 long, erect
setae. Elytra irregularly set with punctures slightly larger than those on
prothorax, separated by about 1-2 puncture diameters, and bearing pale,
posteriorly declined setae producing a sparser clothing than on thorax; long,
erect setae located as follows: 3-10 in each humeral region; 2-8 arranged
in 2 irregular rows on each side of disk; about 25 in irregular patch on
declivity. Thoracic and abdominal sternites closely, almost confluently
punctate to punctatorugose, and densely set with long, pale, appressed setae
interspersed with occasional erect setae; intersegmental sutures incised.
Femora, tibiae and tarsi densely covered with pale, appressed setae. Ae-
deagus gradually attenuate to sharply rounded apex (as in Figs. 5-6); eighth
sternite shallowly emarginate (as in Fig. 7).

Length (pronotum plus elytra), 3.0-3.3 mm.

Holotype female and 12 paratypes from New Cuyama, Santa Barbara
Co., Calif., VI-28-1975, ex litter Quercus wizlizenii, Hobza and Muldowney,
colls.; 1 paratype from S. Fork Kings River Canyon, 5000', Fresno Co.,
Calif., VII-4-1910. The holotype and one paratype are deposited in the Cal-

VOLUME 56, NUMBER 2

127

Fig. 8. Apocrypha setosa, dorsal aspect.

ifornia Academy of Sciences, San Francisco and paratypes in the collection
of the California Department of Food and Agriculture, Sacramento and the
Essig Museum of Entomology, Berkeley, California.

The long setae projecting from the lateral margins of the pronotum dis¬
tinguish A. setosa from anthicoides, which lacks long setae on the thorax.
The larger number of long pronotal setae and the long, erect setae on the
elytra of setosa distinguish it from clivinoides, which averages about 20
percent smaller. Counts of setae vary, even between sides of a single spec-

128

PAN-PACIFIC ENTOMOLOGIST

imen. This variation is partly traumatic, and stubs of setae may be found
on some specimens.

A. setosa and clivinoides appear to be largely allopatric, but each is
known from a single individual from Fresno County. Unfortunately, as men¬
tioned above, the exact collection locality for clivinoides cannot be deter¬
mined.

Description of the Immature Stages of A. anthicoides

Mature larva (Figs. 9-18 ).—Body elongate, cylindrical, pale tan in alco¬
hol preserved specimens, creamy in life; cranium, thorax, legs and abdomen
lightly sclerotized, fragile; maxillae, clypeus, labrum and labium moderately
sclerotized, tan; mandibles and claws heavily sclerotized, dark brown to
black.

Cranium (Fig. 9) subquadrate with rounded corners; prognathous, without
ocellar lenses; cuticle finely rugose with few setae scattered dorsally and
laterally and few short spines posterolaterally; labrum evenly arcuate, with
10 long setae dorsally and marginally; epipharynx (Fig. 10) with 6 short,
blunt setae marginally, scattered circular sensillae centrally, and 2 short
spines basally; antennae articulated on prominences just laterad of clypeus
(Fig. 9); basal segment cylindrical, slender; 2nd segment clavate; 3rd seg¬
ment shorter, slender, with 1 long and several shorter setae at apex; man¬
dibles (Figs. 11-12) stout with strongly bidentate incisor lobe and massive
mola with multituberculate anterior prominence; retinaculum prominent,
midway between mola and incisor teeth; basolateral corner of mandible
bearing several long, coarse setae; maxillae (Figs. 13-14) with cardo trian¬
gularly arcuate, poorly differentiated from stipes; mala with row of stout,
sharp setae on medial margin and irregularly scattered finer setae medially
on dorsal surface; maxillary palp with basal 2 segments subequal, bearing
2 long setae or setal sockets; apical segment about 1.3 times length of 2nd,
gradually narrowing to rounded apex bearing minute, spine-shaped sensil¬
lae; labium (Fig. 15) elongate, with submentum, mentum and prementum
not delimited by sutures; bearing 4 long setae basally, 2 apically and 2
anteriorly directed shorter setae on ligula; palp with segments subequal in
length; apical segment finer, bearing minute, spine-shaped sensillae apically;
hypopharynx (Fig. 16) with low, symmetrical, anteriorly concave sclerome;
dorsal surface of ligula bearing about 15 sharp teeth.

Prothorax nearly as long as 1st abdominal segment, flattened, partly en¬
veloping head in contracted specimens, glabrous at lOOx magnification ex¬
cept for few scattered long projecting setae dorsolaterally and laterally;
mesothorax and metathorax similar, about % length of 1st abdominal seg¬
ment; sterna transverse, postcoxales separate on all segments. Legs similar,
but prothoracic pair (Fig. 17) about 1.3 times larger than posterior pairs;
lightly sclerotized except for articulations, with sparse, scattered setae; cox-

VOLUME 56, NUMBER 2

129

Figs. 9-14. Apocrypha anthicoides, larval structures. Fig. 9, dorsal aspect of head. Fig.
10, epipharynx. Figs. 11-12, dorsal aspect of right and left mandibles. Figs. 13-14, ventral
and dorsal aspects of left maxilla.

130

PAN-PACIFIC ENTOMOLOGIST

Figs. 15-18. Apocrypha anthicoides, larval structures. Fig. 15, labium. Fig. 16, hypo-
pharynx. Fig. 17, anterior aspect of left foreleg. Fig. 18, lateral aspect of abdominal apex.

ae directed anteroventrally, forecoxae contiguous, middle and hind coxae
separated by about 1.5 coxal diameters; trochanters short, triangular, with
single tuberculate seta on each foremember, slender setae on posterior pairs;
femora about as long as coxae, stout, bearing single tuberculate seta on
each foremember; tibiae subequal in length to femora, broadest in middle
and bearing 2 large, stout setae on mesal surface of each foremember, slen¬
der setae on posterior pairs; claw stout, sharp, without differentiated base
and apex.

Abdominal segments cylindrical; segments 1-8 subequal, subquadrate in

VOLUME 56, NUMBER 2

131

Figs. 19-20. Apocrypha anthicoides, pupa, dorsal and lateral aspects respectively.

dorsal aspect, glabrous at lOOx magnification except for sparsely scattered
long setae dorsolaterally and laterally; 9th segment (Fig. 18) about half
length of preceding segments, tapering to rounded apex bearing 4 stout,
stiff, erect spines and bearing numerous long slender setae on dorsal, lateral
and ventrolateral surfaces; 10th segment small, subtending 9th, with short,
blunt pygopods posteriorly. Spiracles simple, annular.

Length, 5.5-6.5 mm; head capsule width, 0.49-0.52 mm.

Pupa (Figs. 19-20). —Elongate, subcylindrical with enlarged globular pro¬
thorax, diagonally projecting elytra, and deflexed head; cream colored with
long fine setae.

Head elongate, slightly depressed, exserted and deflexed beneath pro-

132

PAN-PACIFIC ENTOMOLOGIST

thorax (Fig. 19); vertex and epistoma with sparsely scattered, long, erect
setae; antennae free, extending posterolaterally above or below forelegs and
elytra.

Prothorax globular, excavated anteroventrally for reception of head; lat¬
eral margins rounded, without trace of carina; pronotum nearly round in
dorsal aspect, with scattered, long, erect setae peripherally; mesonotum
about l A as long as pronotum, narrower anteriorly, subequal in width to
metanotum posteriorly, and bearing 2 long setae between elytra; elytra di¬
rected diagonally backward and slightly downward or nearly horizontal,
faintly 9-striate, glabrous; metanotum short, transverse, bearing 2 long erect
setae; wing sheaths about 3 A as long as elytra, without apparent venation;
metacoxae projecting ventrally well below level of abdomen. Femora cla-
vate, with short setae on knees.

Abdominal segments 1-6 subcylindrical, about twice as broad as long,
bearing sparsely scattered, long, posteriorly declined setae; lateral margins
without gin traps, but with 2 small tubercles each bearing long, projecting
seta; segment 7 tapering, rounded posteriorly; abdominal apex with appar¬
ently 2-segmented projecting papillae in female, shorter unsegmented pa¬
pillae in male.

Length, 3.0-3.5 mm.

Material examined .—5 larvae, laboratory reared from adults collected in
Calif., S. L. Obispo Co., San Simeon, IV-1976, C. Y. Kitayama; harvested
XII-1976. 20 larvae, 6 pupae, laboratory reared from adults collected in
California, S. L. Obispo Co., 3 mi. S Oceano, 11-23-1975, J. Doyen; har¬
vested IV-18 to VII-24-1975.

The larvae of Apocrypha are superficially very similar to larvae of Tri-
bolinm and other small Ulomini. As elaborated below, this similarity ex¬
tends to several anatomical details. Berlese extraction yielded only adult
beetles, suggesting that the larvae inhabit deeper layers of the soil. The
absence of gin traps on pupae is unusual in Tenebrionidae. Presumably the
tiny lateral abdominal tubercles represent reduced gin traps. Presence of
pupal wing sheaths in brachypterous or flightless Coleoptera is common
(Smith, 1964; Spilman, 1979). In species in which the adult is entirely wing¬
less, the pupal wing sheath is usually much shorter than the elytral sheath,
and the adult wing is absent or only partly fills the sheath, and degenerates
before the adult emerges. Apocrypha is unusual in that the adult is com¬
pletely apterous with the metanotum reduced to a membrane, yet the pupal
wing sheaths are about two-thirds as long as the elytral sheaths and about
half as broad. No imaginal wings are visible within the pupal sheaths, al¬
though wings could have been present and degenerated, since the ages of
the pupae are unknown.

Spilman (1979) examined pupae of various wingless and brachypterous
Coleoptera, and concluded that in general pupal wing sheaths are short or

VOLUME 56, NUMBER 2

133

absent in taxa in which brachyptery or winglessness is widespread. In con¬
trast, in taxa with occasional wingless or brachypterous members, pupal
wing sheaths are long. This might suggest that Apocrypha represents a
specialized, wingless member of some larger taxon containing mostly
winged species. However, the apparent total absence of imaginal wings
suggests derivation from a higher taxon in which wing reduction is wide¬
spread. The only other species of Apocryphini which we have dissected, A.
clivinoides, is apterous.

Apocrypha shows some other interesting differences between the pupa
and adult. In the adults long, erect setae are present on the elytra and head,
absent from the thorax. In the pupa, setae are present on the head and
pronotum but absent on the elytra. The elytra of adult Apocrypha anthi-
coides are without defined striae, although striae are present in some South
American species (Kaszab, 1969). In pupae of A. anthicoides striae are
visible, though very faint in some specimens. These pupal striae are difficult
to count, but apparently are nine in number.

Phylogenetic Relationships

Apocryphini has been placed as a tribe in the subfamily Tenebrioninae
since the work of Lacordaire (1859), who suggested a position close to
Helopini. Watt (1974) treated Apocryphini as a tribe of Lagriinae, but in¬
dicated only a few characters of adults which support this relationship—
notably, the absence of lateral pronotal carinae (shared with Lagriini) and
the presence of 10-striate elytra in some (unspecified) species (universal in
Lagriinae). Similarities resulting from loss of structures (such as carinae)
can easily arise through convergence. The retention of 10 elytral striae is a
primitive feature in Coleoptera, and not very useful in indicating cladistic
relationship. Moreover, the pupal elytra of A. anthicoides bear only 9 striae,
suggesting that this may be the primitive number for the genus.

Watt’s Lagriinae is defined primarily by characters of the larvae, espe¬
cially the structure and position of the antennae. In most Tenebrionidae the
larval antennae have three subequal segments or have the third segment
variably reduced. In Lagriinae the larval antennae have two densely pubes¬
cent segments, with the second much longer than the first. The second
segment bears a distinctive sensorium or group of sensillae at the apex, and
the antennae are clearly separated from the bases of the mandibles by a
strip of cuticle (contiguous in all other Tenebrionidae except Nilionini). In
addition the epipharynx and hypopharyngeal sclerome are highly asymmet¬
rical in Lagriinae (symmetrical in most Tenebrionidae). In all of these fea¬
tures Apocrypha exhibits the generalized character state common to Te¬
nebrioninae and most other subfamilies. In addition, the larva of Apocrypha
is elongate, cylindrical, and weakly sclerotized. Larvae of Lagriinae are

134

PAN-PACIFIC ENTOMOLOGIST

usually much stouter, flattened and moderately sclerotized, especially dor-

sally.

Several features of adult Apocrypha indicate derivation from Tenebrion-
inae. The labrum is short and transverse (elongate in Lagriinae) and the
epipharynx membranous and symmetrical (asymmetrical with sclerotized
plates in Lagriinae). The ovipositor has the gonostyles short and situated
subapically. In Lagriinae, Diaperinae and most other primitive subfamilies
the gonostyles are elongate and apical. The defensive glands are small and
saccate, and the bursa copulatrix bears a single accessory tube which
branches to form the spermatheca and the accessory gland. Diverse glan¬
dular structures occur in Lagriinae, but simple saccate glands without ex¬
trinsic musculature are not represented (Tschinkel and Doyen, in press).
In many Lagriinae multiple spermathecal tubules originate independently
from the bursa. In other Lagriinae the bursa (which may be constricted
to produce 2 subequal chambers) bears a single, unbranched tube which
apparently represents the spermathecal accessory gland.

Precise affinities of Apocryphini within Tenebrioninae cannot be specified
with certainty. Lacordaire’s Helopides contained Adeliini, Misolampini and
Helopini, as well as Apocrypha. These taxa share only a general similarity
in superficial external characters, and are now classified in remote parts of
the Tenebrionidae. Misolampini appear synonymous with Coelometopini
based on structures of defensive glands and the female reproductive tract
(Tschinkel and Doyen, in press), which are very different than those
in Apocrypha. Described larvae of Helopini have long, recurved urogomphi,
frequently opposing sclerotized processes on abdominal tergite eight, and
have the hypopharyngeal sclerome tridentate apically. The larva of Apoc¬
rypha lacks urogomphi, has the sclerome concave anteriorly and the head
and mouthparts differ in many structural details from those of Helopini (see
Byzova and Gilyarov, 1956).

Several of the larval features of Apocrypha are shared with Opatrini and/
or the Tribolium group of Ulomini. For example, the third antennal segment
is relatively large in Tribolium, Palorus and Alphitophagus (Ulomini), as in
Apocrypha. The mandibles of Apocrypha, with large, subequal incisor teeth
and the large anterior molar prominence, are similar to those of many Ulom¬
ini and Opatrini (see Hayashi, 1966). Likewise, the anteriorly concave hy¬
popharyngeal sclerome is shared with many representatives of those tribes.
The enlarged forelegs of Apocrypha, with modified, peg-shaped setae on
the trochanter and femur, are similar to those of many Opatrini and Pedinini
(see Hayashi, 1966; Keleinikova, 1966; Skopin, 1960). Finally, the stiff mar¬
ginal spines on the apex of the ninth abdominal tergite are shared with many
Opatrini and Ulomini.

Some of the features discussed above are probably plesiomorphic and not
indicative of cladistic relationships. For example, the relatively elongate

VOLUME 56, NUMBER 2

135

antennae, with approximately equal segments, are probably primitive to the
subfamily Tenebrioninae, if not the entire family. Other features, such as
the enlarged forelegs with specialized spination, and the elongate, cylindri¬
cal body without urogomphi are shared by soil inhabiting species from sev¬
eral tribes (e.g., the Pedobionta of Skopin, 1964 or the combined tenebrioid
and opatroid lineages of Keleinikova, 1963). Such features may have arisen
convergently in Apocrypha. However, the balance of characters indicates
a position near Opatrini or Ulomini, where Apocryphini should be retained
as a separate tribe until its relationships can be further elucidated.

Acknowledgments

Field research was supported in part by National Science Foundation
Grant BMS74-17924. Peter Churchward, Dune Lakes Limited, Arroyo
Grande, California, kindly gave permission to sample at certain of the field
sites in San Luis Obispo County. The following curators and institutions are
gratefully acknowledged for loan of material under their care: F. Andrews,
California Department of Food and Agriculture, Sacramento; J. Chemsak,
Essig Museum of Entomology, University of California, Berkeley; D. Ka-
vanaugh, California Academy of Sciences, San Francisco; J. F. Lawrence,
Museum of Comparative Zoology, Harvard University, Cambridge, Mas¬
sachusetts. We thank Celeste Greene of the University of California at
Berkeley for the excellent illustrations of the pupa and adult.

Literature Cited

Byzova, Y. B., and M. S. Gilyarov. 1956. Soil-dwelling tenebrionid larvae of the tribe Helopini
(Coleoptera, Tenebrionidae). Zool. Zhurn., 35:1493-1508.

Doyen, J. T., and J. F. Lawrence. 1979. Relationships and classification of some Tenebrion¬
idae and Zopheridae (Coleoptera). Syst. Entomol., 4:333-377.

Eschscholtz, J. F. 1831. Zoologischer atlas, enthaltend Abbildungen und Beschreibungen
neuer Thierarten, wahrend des Flottcapitains von Kotzebue zweiter Reise um die Welt,
auf der Russisch-Kaiserlichen Kriegsschlupp Predpriaetie in den Jahren 1823-1826.
Viertes Heft, 19 pp.

Gebien, H. 1938-42. Katalog der Tenebrioniden, Teil II. Mitteilungen Munchner Entomol.
Gesell., 28-32:370-744 [repaged].

Hayashi, N. 1966. A contribution to the knowledge of the larvae of Tenebrionidae occurring
in Japan (Coleoptera: Cucujoidea). Ins. Matsumurana, Suppl., 1:1-41, 32 pis.

Horn, G. H. 1870. Revision of the Tenebrionidae of America, north of Mexico. Trans. Amer.
Philos. Soc., ser. 2, 14:253-404.

Kaszab, Z. 1969. The scientific results of the Hungarian soil zoological expedition to South
America. 17. Tenebrioniden aus Chile. Opusc. Zool. (Budapest), 9:291-337.

Keleinikova, S. I. 1963. Larval types of Palearctic darkling beetles (Coleoptera, Tenebrioni¬
dae). Entomol. Rev., 42:292-296.

Keleinikova, S. I. 1966. Descriptions of larvae of some Palearctic genera of darkling beetles
of the Tribe Pedinini (Coleoptera, Tenebrionidae). Entomol. Obozrenie, 45:589-598.

136

PAN-PACIFIC ENTOMOLOGIST

Keleinikova, S. I. 1968. Soil-dwelling darkling beetle larvae (Col., Tenebrionidae) of the
fauna of the USSR. I. Coll. Works Biol. St. Univ. Moscow, 11:205-239.

Kulzer, H. 1962. Neue Tenebrioniden aus Sudamerika (Col.). Entomol. Arbeit. Mus. Frey,
13:79-100.

Lacordaire, T. 1859. Histoire naturelle des Insects. Genera des Coleopteres. Tome Cinquieme.
Premier Partie. (Heteromera, in Part), pp. 1-750.

LeConte, J. L. 1851. New species of California Coleoptera. Ann. Lyceum Nat. Hist. New
York, 5:125-216.

LeConte, J. L. 1862. Classification of the Coleoptera of North America. Prepared for the
Smithsonian Institution. Smithson. Misc. Coll., 136:209-286.

LeConte, J. L., and G. H. Horn. 1883. Classification of the Coleoptera of North America.
Smithson. Misc. Coll., 507:38-567.

Mayr, E. 1963. Animal Species and Evolution. Harvard University Press. Cambridge, Mass.,
xiv + 797 pp.

Skopin, N. G. 1960. Data on the morphology and ecology of larvae of the tribe Blaptini
(Coleoptera: Tenebrionidae). Trudy Inst. Zool. Akad. Nauk Kaz. S.S.R., 11 (entomol-
ogy):37-71, 13 pis.

Skopin, N. G. 1964. Die Larven der Tenebrioniden des Tribus Pycnocerini (Coleoptera, Het¬
eromera). Ann. Mus. r. Afr. cent. (Ser. 8), No. 127:1-35.

Solier, A. J. 1851. Orden III. Coleopteros. In Gay, Historia fisica y politica de Chile, Vol. 5,
pp. 1-285.

Smith, D. S. 1964. The structure and development of flightless Coleoptera: a light and electron
microscopic study of the wings, thoracic exoskeleton and rudimentary flight muscula¬
ture. Jour. Morphol., 114:107-183.

Spilman, T. J. 1979. Larvae and pupae of Centronopus calcaratus and suppressus (Coleop¬
tera, Tenebrionidae), with an essay on wing development in pupae. Proc. Entomol. Soc.
Wash., 81:513-521.

Tschinkel, W. F. and J. T. Doyen. 1980. Comparative anatomy of the defensive glands,
ovipositors and female genital tubes of tenebrionid beetles (Coleoptera). Intern. Jour.
Insect. Morph. Embryol. (in press).

Watt, T. C. 1974. A revised subfamily classification of Tenebrionidae (Coleoptera). New
Zealand Jour. Zool., 1:381-452.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 137-143

TWO NEW SPECIES OF MACRUROHELEA FROM CHILE
WITH A KEY TO THE NEOTROPICAL SPECIES
(DIPTERA: CERATOPOGONIDAE)

William L. Grogan, Jr.

Department of Biological Sciences, Salisbury State College,

Salisbury, Maryland 21801

AND

Willis W. Wirth

Systematic Entomology Laboratory, IIBIII, Agric. Res., Sci. and Educ.
Admin., USDA, % U.S. National Museum Washington, D.C. 20560

Ingram and Macfie (1931) described Macrurohelea for two species from
southern South America, M. caudata I. & M., the type species, and M.
thoracica I. & M. Wirth (1965) presented a revised diagnosis of the genus,
described two new species from Chile, and offered a key to separate the
known species. He also discussed the relationship of the genus with Stilo-
bezzia Kieffer and other genera considered to be related to that genus. Wirth
(1974) assigned Macrurohelea to the tribe Stilobezziini and Wirth et al.
(1974) included the genus in their key to the genera of Ceratopogonidae.
Grogan and Wirth (1977), in their revision of the Nearctic Parabezzia Mal-
loch stated that the male genitalia of Macrurohelea are similar to those of
Parabezzia, and to the closely related Fittkauhelea Wirth and Blanton.

Subsequently, Grogan and Wirth (1979), in describing the new genus No-
tiohelea from Chile, remarked on the similarity of that genus and Macru¬
rohelea to Ceratopogon Meigen. They stressed that all three of these genera
possess sensilla coeloconica on the first flagellomere of the antenna, a char¬
acter absent in Stilobezzia and its relatives. At least one species of Macru¬
rohelea, M. thoracica Ingram and Macfie, has sensilla coeloconica present
on flagellomeres 5-8 as well as on one. We have seen only one other species
of Ceratopogonini possessing these sensilla beyond the first flagellomere,
Brachypogon fuscivenosus (Lutz), which also has them on flagellomeres 2-
4. We have not found any other ceratopogonids except for the Culicoidini
with these sensilla on the distal flagellomeres.

On the basis of its sensillar characters, we assign Macrurohelea to the
tribe Ceratopogonini. This placement of Macrurohelea is supported by its
possession of pubescence between the eye facets. The genus is perhaps
most closely related to Ceratopogon and is probably a sister group of that

138

PAN-PACIFIC ENTOMOLOGIST

genus that is restricted to the southern hemisphere. Ceratopogon is a Hol-
arctic genus, ranging north to at least 82°N latitude and probably as far
south as 35°N. Macrurohelea exhibits a similar latitudinal distribution in
South America.

In the present paper we describe two new species of Macrurohelea re¬
cently taken by M. E. Irwin in Chile. These descriptions increase the num¬
ber of known species of the genus to seven, and there are probably many
more yet to be discovered. An Australian species, M. commoni Lee (1962),
is not included in the present key but is distinguished from all of the Neo¬
tropical species by having the second radial cell over four times the length
of the first.

For an explanation of general ceratopogonid terminology see Wirth
(1952); terms dealing with male genitalia are those of Snodgrass (1957);
terminology of antennal sensilla follows Wirth and Navai (1978). We grate¬
fully acknowledge the aid of Susan Powell, who prepared the illustrations.

Key to the Neotropical Species of Macrurohelea

1. Females. 2

Males. 6

2. Second radial cell of wing twice as long as the first . 3

Second radial cell at least 3 times as long as the first. 4

3. Flagellum very short, flagellomeres 9-12 each broader than long,

antennal ratio 0.59; very small species, wing length 0.94 mm ...

. kuscheli Wirth

Flagellum longer, flagellomeres 9-12 each twice as long as broad,
antennal ratio 1.06-1.16; small species, wing length 1.27-1.42
mm . irwini new species

4. Small species, wing length 1.3-1.5 mm; intercalary fork of wing pres¬

ent . caudata Ingram and Macfie

Larger species, wing length 2.1 mm or greater; intercalary fork of
wing absent . 5

5. Flagellomeres 5-8 with apical sensilla coeloconica; legs with incon¬

spicuous setae; wing including the veins pale .

. thoracica Ingram and Macfie

Flagellomeres 5-8 lacking sensilla coeloconica; legs with numerous
long bristly setae; wing including veins infuscated dark brown . .

. setosa Wirth

6. Large species, wing length 2.1 mm or greater . 7

Smaller species, wing length less than 1.5 mm. 8

7. Legs with numerous long bristly setae; wing including veins infus¬

cated dark brown . setosa Wirth

VOLUME 56, NUMBER 2

139

Figs. 1-5. Macrurohelea irwinr, 6, M. paracaudata. 1, female antenna; 2, female wing; 3,
spermathecae; 4, female palpus; 5, 6, male genitalia.

Legs with inconspicuous setae; wing including veins pale .

. thoracica Ingram and Macfie

8. Very small species, wing length 0.90 mm ... paracaudata new species

Small species, wing length 1.3 mm or greater.9

9. Intercalary fork of wing present; second radial cell subequal to first

. caudata Ingram and Macfie

Intercalary fork of wing absent; second radial cell twice as long as
first. irwini new species

140

PAN-PACIFIC ENTOMOLOGIST

Macrurohelea irwini, new species
(Figs. 1-5)

Diagnosis .—A small species of Macrurohelea distinguished by the fol¬
lowing combination of characters: Wing length 1.27-1.43 mm, the second
radial cell twice as long as the first; females with long antennal flagellum,
flagellomeres 9-12 twice as long as broad, antennal ratio 1.06-1.16; males
with tip of aedeagus bifid.

Holotype female .—Wing length 1.27 mm; breadth 0.50 mm.

Head. —Brown. Eyes pubescent; barely contiguous; facets broadly sep¬
arated. Antenna with dark brown pedicel; flagellum (Fig. 1) with proximal
eight flagellomeres globose, distal five flagellomeres more elongated, about
twice as long as broad; first flagellomere with 2-3 apical sensilla coelocon-
ica; proximal flagellomeres each with subbasal whorl of six sensilla chaetica
and a central whorl of three long sensilla trichodea; distal five flagellomeres
with scattered sensilla chaetica and sensilla basiconica; flagellomeres with
lengths in proportion of 12-7-7-7-7-7-7-7-11-12-12-12-20; antennal ratio 1.10.
Palpus (Fig. 4) light brown; lengths of segments in proportion of 6-12-11-7-
12; third segment twice as long as broad with a small shallow apical pit
bearing slender capitate sensilla. Mandible with 9-10 teeth, distal teeth long¬
est.

Thorax: Dark brown with sparse scattered setae and fine pubescence.
Legs uniform brown; femora with sparse setae, setae on tibiae moderately
dense; first tarsomere of fore and hind legs with well developed palisade
setae; fourth tarsomeres cordate; fifth tarsomeres about three times longer
than broad with small equal simple claws. Wing (Fig. 2) hyaline, moderately
broad, about 2.5 times longer than broad, veins grayish; second radial cell
twice as long as first; costa extending to 0.70 of wing length; venation as
figured. Flalter stem light brown; knob pale.

Abdomen: Dark golden brown; covered with uniform, fine pubescence
and a few scattered setae on sterna and terga. Tenth segment elongated and
bent forward ventrally as is typical for members of the genus. Spermathecae
(Fig. 3) heavily sclerotized, spheroid, subequal, with stout, moderately long
necks.

Allotype male .—Similar to holotype female with the usual sexual differ¬
ences. Wing length 1.43 mm; breadth 0.49 mm; costa extending to 0.62 of wing
length; flagellum and fifth tarsomeres and claws of fore and mid legs lost.
Genitalia as in Fig. 5. Ninth sternum about three times broader, than long,
with a shallow caudomedian excavation; ninth tergum tapering rather
abruptly distally to a narrow truncate apex bearing two short apicolateral
processes, cerci very short and subapical to apicolateral processes. Basi-
mere nearly straight, about twice as long as broad; telomere about half the
length of basimere, curved gradually distally to pointed tip. Aedeagus tri¬
angular, slightly broader than long, moderately heavily sclerotized with deep

VOLUME 56, NUMBER 2

141

basal arch about Vs of total length; membrane spiculate but ventral surface
of aedeagus bare; basal arms heavily sclerotized and recurved 90°; distal
portion tapering rather abruptly to slightly bifid tip. Claspettes nearly sep¬
arated; basal arm very heavily sclerotized and doubly recurved; distal por¬
tion lightly sclerotized with tips bent at 90°.

Variation. — Females. —Wing length 1.37 (1.27-1.41, n = 4); breadth 0.57
(0.50-0.61, n = 4). Antennal ratio 1.11 (1.06-1.16, n = 3).

Distribution .—Chile; known only from the type locality.

Types. —Holotype female, allotype male, 3 paratype females, Chile, San¬
tiago Prov., Quebrada de la Plata Maipu, 33°30'S, 70°55'W, 10 Aug. 1966,
M. E. Irwin (Deposited in Calif. Acad. Sci., San Francisco; paratypes in
U.S. National Museum and University of California, Riverside).

Discussion .—We take great pleasure in naming this species for its collec¬
tor, Michael E. Irwin of the University of Illinois, in recognition of his
contributions to our knowledge of Chilean Diptera.

M. irwini was taken at the same time and place as the holotype male of
M. paracaudata n.sp. We are associating the single male of M. irwini with
the females of this species because they are nearly identical in size, as is
the case with other species in this genus.

This species appears to be most similar to M. kuscheli Wirth in having
a small, short, broad wing with the second radial cell twice as long as the
first. However, M. kuscheli differs from M. irwini in having a smaller wing
(wing length 0.94 mm), shorter antenna (antennal ratio 0.59) with flagello-
meres 9-12 broader than long. The male genitalia of M. irwini are distinctive
and easily distinguished from those of other species by the broad triangular
aedeagus with bifid tip.

Macrurohelea paracaudata, new species

(Fig. 6)

Diagnosis .—A small species of Macrurohelea : males distinguished by the
following combination of characters: Small size (wing length 0.90 mm); ninth
sternum with distinct caudomedian notch bearing setose tubercles on each
side; aedeagus with hastate tip.

Holotype male .—Wing length 0.90 mm; breadth 0.34 mm.

Head .—Brown. Eyes pubescent, moderately broadly separated. Pedicel
of antenna dark brown; flagellum lost. Palpus light brown; segments shrunk¬
en and not measured; fifth segment more than twice as long as fourth.

Thorax .—Dark reddish brown with sparse scattered setae and fine pu¬
bescence. Legs uniform lighter brown; femora with sparse setae, tibiae with
more dense setae; first tarsomere of fore and hind legs with well developed
palisade setae; tarsi of mid legs and all but first tarsomere of hind leg lost;

142

PAN-PACIFIC ENTOMOLOGIST

fourth tarsomere of fore leg cordate; fifth tarsomere of fore legs slender with
small, equal, simple claws. Wing similar to that of M. irwini n. sp. (Fig. 2);
second radial cell about 1.4 times the length of first; costa extending to 0.64
of wing length. Halter lost.

Abdomen .—Dark golden brown. Genitalia as in Fig. 6. Ninth sternum
with caudomedian notch bearing setose tubercles on each side; ninth tergum
gradually tapering distally to a broad truncate tip bearing two rather long
apicolateral processes; cerci short, subapical to apicolateral processes. Bas-
imere greatly curved, nearly twice as long as broad with basal mesal lobe;
telomere nearly the length of basimere, greatly curved to pointed tip. Ae-
deagus rather short, 1.3 times broader than long, heavily sclerotized with
very shallow basal arch only about Va of total length, membrane and ventral
surface not spiculate; basal arm rather long and slender with recurved tip;
distal portion very short and not as heavily sclerotized, the tip hastate or
shaped like the point of a crowquill pen. Claspettes nearly fused at base;
basal arm heavily sclerotized with slender, ventrally projecting portion; dis¬
tal portion more lightly sclerotized with broad flat tip that bends at apex.

Female. —Unknown.

Distribution. —Chile; known from a single specimen from the type local¬
ity.

Type .—Holotype male, Chile, Santiago Prov., Quebrada de la Plata Mai-
pu, 33°30'S, 70°55'W, 10 August 1966, M. E. Irwin (Calif. Acad. Sci.).

Discussion .—The affinities of M. paracaudata with other species of the
genus are uncertain, but this species has a low aedeagus similar to that of
M. caudata Ingram and Macfie, hence the specific name. M. caudata differs
by having a straight stout basimere, a narrower ninth sternum, apex of ninth
tergum lacking apicolateral processes, and wing with an intercalary fork.

Literature Cited

Grogan, W. L., Jr., and W. W. Wirth. 1977. A revision of the Nearctic species of Parabezzia
Malloch (Diptera: Ceratopogonidae). J. Kans. Entomol. Soc., 50:49-83.

Grogan, W. L., Jr., and W. W. Wirth. 1979. Notiohelea, a new genus of biting midges of the
tribe Ceretopogonini from Chile (Diptera: Ceratopogonidae). Pan-Pacific Entomol.,
54:283-286.

Ingram, A., and J. W. S. Macfie. 1931. Ceratopogonidae. Diptera of Patagonia and South
Chile, Part II, Fascicle 4, pp. 155-232.

Lee, D. J. 1962. Australasian Ceratopogonidae (Diptera, Nematocera). Part IX. The genus
Macrurohelea. Proc. Linn. Soc, N.S.W., 87:339-340.

Snodgrass, R. E. 1957. A revised interpretation of the external reproductive organs of male
insects. Smithson. Misc. Collect., 135:1-60.

Wirth, W. W. 1952. The Heleidae of California. Univ. Calif. Publ. Entomol., 9:95-266.

Wirth, W. W. 1965. Two new species of Macrurohelea from Chile (Diptera, Ceratopogonidae).
Pan-Pacific Entomol., 41:46-50.

VOLUME 56, NUMBER 2

143

Wirth, W. W. 1974. A catalogue of the Diptera of the Americas south of the United States.
14. Family Ceratopogonidae, pp. 1-89. Museu de Zoologia, Universidade de Sao Paulo,
Brazil.

Wirth, W. W., and S. Navai. 1978. Terminology of some antennal sensory organs of Culicoides
biting midges (Diptera: Ceratopogonidae). J. Med. Entomol., 15:43-49.

Wirth, W. W., N. C. Ratanaworabhan, and F. S. Blanton. 1974. Synopsis of the genera of
Ceratopogonidae (Diptera). Ann. Parasitol. Hum. Comp., 49:595-613.

NOTICE

BELTSVILLE AGRICULTURAL RESEARCH CENTER

SYMPOSIUM V

The Beltsville Agricultural Research Center sponsors an annual research
symposium with a specific theme. The subject of the fifth “BARC Sympo¬
sium” will be “Biological Control in Crop Production.” It is scheduled for
May 19 to May 21, 1980. Subject matter will be presented as invited lectures
and contributed posters with the lectures published in the BARC symposium
series (5th volume). Previous symposia in this series were: (1) Virology in
Agriculture, (2) Biosystematics in Agriculture, (3) Animal Reproduction,
and (4) Human Nutrition Research: Questions and Answers.

Registration and a reception will be held Sunday evening followed by five
technical sessions held Monday morning through Wednesday noon. The
sessions are as follows:

Session 1—Relevance of ecological theories to practical biological control.
Session 2—Concepts, principles and mechanisms of biological control of
pests.

Session 3—Recent advances in mass production of biological control agents.
Session 4 —Strategies of biological control.

Session 5—General considerations: Environmental, regulatory, safety, eco¬
nomic and biocontrol in integrated pest management systems.

Voluntary poster presentations will be held Monday from 5:30 to 7:30 pm.
Registration fee $60.00.

Anyone wishing to receive a registration packet for this symposium
should contact:

Publicity Chairman, Symposium V, Room 214, Bioscience Bldg 011 A,
BARC-West, Beltsville, Md. 20705.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 144-152

PARASITISM OF SWEAT BEES IN THE GENUS AGAPOSTEMON
BY CUCKOO BEES IN THE GENUS NOMADA
(HYMENOPTERA: HALICTIDAE, ANTHOPHORIDAE)

George C. Eickwort and Judith Abrams
Department of Entomology, Cornell University, Ithaca, New York 14853

Nomada is a large Holarctic genus of cleptoparasitic (“cuckoo”) bees
whose species superficially resemble wasps. Most species parasitize soil¬
nesting bees in the genus Andrena (Andrenidae), but a few have been re¬
corded from other soil-nesting Andrenidae ( Panurgus, Nomadopsis ), Hal-
ictidae ( Halictus , Lasioglossum, Evylaeus, Nomia), Melittidae ( Melitta ,
Dasypoda), and Anthophoridae ( Eucera ) (Stoeckhert, 1933, 1954; Stephen,
Bohart, and Torchio, 1969; Bohart, 1970). Despite the abundance of many
species of Nomada, only Linsley and MacSwain (1955) have conducted
detailed studies of their biology. Rozen (1966) described the larvae of five
species and he (Rozen, 1977) also described the pupa of one species and
summarized nomadine behavior.

In this paper we record the first definite association between species of
Nomada and host species in the sweat bee genus Agapostemon (Halicti-
nae). Species of Nomada previously had been suspected of parasitizing
species of Agapostemon (Linsley, MacSwain, and Smith, 1954; Roberts,
1969) but were never recovered from nest cells. We also present observa¬
tions on nest-entering behavior, development of immature stages, and in¬
cidence of parasitism to supplement the meager biological data on Nomada.

Materials and Methods

Nomada ( Nomadula ) articulata Smith was studied in Ithaca, Tompkins
Co., New York, principally on the Cornell University campus. J. A. made
a detailed study of the above-ground behavior of one of its hosts, the uni-
voltine, communally nesting Agapostemon virescens (Fabricius), at an ag¬
gregation of 13 nests in a 2 sq. m area in a weedy, sporadically mowed
campus location in 1975 (Abrams, 1977 1 ). Three female N. articulata were
captured as they emerged from host nests, on 1 and 3 June and 5 July.
She observed 15 female parasites at the nest site in June and July.
Eleven of these were marked for individual recognition; seven reappeared
at the nest site. She also recovered three adult N. articulata in host nests
excavated in October. At the same site, G.C.E. excavated a nest containing
immature N. articulata on 18 July 1972. G.C.E. also recovered N. articu¬
lata from nests of univoltine Agapostemon radiatus (Say) in a level, moist,

VOLUME 56, NUMBER 2

145

grassy lawn on the Cornell campus about 270 m from the nest site of A.
virescens. Five nests, excavated on 19 July 1974, 23 June 1978, and 25 July
1978, contained immature Nomada. Adult Nomada were collected at the
site but we did not conduct detailed behavioral observations.

Nomada ( Micronomada ) formula Viereck was studied near Richmond,
Contra Costa Co., California, at the entrance to San Pablo Reservoir.
G.C.E. observed Nomada flying about an aggregation of active nests of
Agapostemon texanus Cresson located in level, bare, dry sandy soil, on 22
and 29 August 1975. He excavated five nests that contained immature N.
formula on 24 and 25 September.

Techniques of observation, nest excavation, and rearing and preservation
of immature bees are given by Abrams and Eickwort (1980a). In addition,
some prepupae were kept in a refrigerator for several months to break dia¬
pause. Twenty-nine N. articulata immatures and 12 N. formula immatures
were recovered from nest cells, of which six of the former and two of the
latter species were reared to adults.

Voucher specimens of cleptoparasites and hosts and their immature stages
are deposited in the Cornell University insect collection, lot number 995.
Larvae and pupae of Nomada are also deposited in the American Museum
of Natural History.

Nomada (Nomadula) articulata Smith

Marked adult females returned repeatedly to the Agapostemon virescens
nest site. One female, marked on 2 July, visited the site on 15 of the next
18 observation days. Females were active from 0800 to 1230 hr. They flew
in both sun and shade, but not when air temperatures were below 22°C.
Typically a female visited the aggregation once or twice per morning and
remained each time from 15 to 45 minutes, repeatedly attempting to enter
nests. Her behavior was similar to that of Nomada opacella Timberlake
and Nomada sp. at Andrena nest sites in California (Linsley and MacSwain,
1955; Thorp, 1969). Nomada articulata flew slowly through the aggregation
a few centimeters above the ground and alighted on raised perches of earth,
twigs, leaves, or stones, 3 to 12 cm from nest entrances. A female waited
on the perch with her antennae directed towards the nest opening from 5
sec to 10 min before approaching the entrance (Fig. 1). About half of the
attempts to enter nests were made immediately after an Agapostemon had
left on a foraging trip. The other attempts were made just after an Agapos¬
temon had entered her nest or when no activity could be seen at the nest
entrance. Nomada did not approach a nest if a guard was visible at the
entrance.

Each N. articulata female followed the same behavioral sequence when
attempting to enter an A. virescens nest. She walked slowly towards the

146

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-6. Nomada articulata. Fig. 1, adult female perched near host nest. Fig. 2, adult
female attempting entry into nest of Agapostemon virescens. Fig. 3, first instar larva on egg
of A. virescens. Fig. 4, larva feeding on provision mass of A. virescens. Fig. 5, prepupa,
lateral view. Fig. 6, prepupa, ventral view.

VOLUME 56, NUMBER 2

147

nest entrance with her wings vibrating. She stopped at the entrance, still
fanning her wings, and inserted her antennae into the burrow for 1 or 2 sec
(Fig. 2). She then stopped vibrating her wings and walked into the burrow.

If the Nomada encountered an Agapostemon guard when she inserted
her antennae into the nest entrance, she immediately backed away from the
burrow and returned to a watching position or investigated another nest.
The guard either snapped at the intruder with her mandibles or advanced
towards the Nomada until her thorax was outside the entrance. If the guard
was further down the burrow, the Nomada entered the nest but always
reappeared within 10 sec. The guard followed the Nomada to the surface
and blocked the nest entrance with her head or the dorsal surface of her
abdomen. Of 50 attempts to enter nests, 48 were unsuccessful.

Agapostemon virescens did not interact with Nomada outside the nests.
Guards did not chase parasites perched nearby, and on several occasions
a returning forager flew right over a Nomada in order to enter her nest. In
these instances, the Nomada immediately retreated.

Only one nest was entered successfully in 1975, by two N. articulata
females on 1 July. This was the only nest in the aggregation that was oc¬
cupied by a solitary Agapostemon female during the pollen-foraging season.
July 1 was the only day that the host was seen foraging. She brought a load
of pollen into her nest at 0936, emerged at 0941, and flew away. At 0957 an
unmarked N. articulata entered the nest and remained in the burrow 3 min
before emerging. The Agapostemon brought in two more pollen loads by
1050. At 1055, a second, marked Nomada, which had just been driven away
from a communal nest, entered this nest and also remained 3 min before
departing. Observations on 1 July were ended at 1100 hr. Although the nest
entrance was open for the next three days, the Agapostemon had apparently
abandoned her nest because she was never seen again. Two Nomada ap¬
proached the nest entrance during these three days but neither descended
into the burrow. Like other nomadines (Rozen, 1977), N. articulata prob¬
ably oviposits only in host cells that are in the stage of being provisioned.

In two more instances in 1975, Nomada may have successfully entered
nests of A. virescens. Both cleptoparasites were found trapped under plastic
cups that were used to check exiting Agapostemon. Since Nomada walk
to hosts’ nest entrances, they may have approached the burrows through
the gap between the ground and the edge of the cup. In one of these cases,
a guard was blocking the nest entrance with the dorsal surface of her ab¬
domen. The Nomada had apparently been stung, because she was partially
paralyzed and died within 10 min.

Nomada opacella makes orientation flights after examining host burrows
(Linsley and MacSwain, 1955). Most TV. articulata had probably learned
nest locations before we began constant observations at the Cornell nest

148

PAN-PACIFIC ENTOMOLOGIST

site, but one female was observed making an orientation flight. On 5 July,
the female Nomada emerged from an Agapostemon nest at 0855 hr. She
remained at the nest site, walking and taking short flights, until 1028 and
then flew off. At 1121 she returned and attempted to enter a second nest,
but was stopped by a guard. She then made a short orientation flight of
three or four loops and left the nest site. On 6 July, she returned and at¬
tempted to enter both the nest from which she had first emerged and the
second nest that she had located on 5 July.

In the nest of A. virescens excavated in 1972 that contained Nomada
immatures, at least four of 13 cells (31%) were infested by the cleptoparasite.
This nest also contained only one adult host. In contrast to A. virescens,
Agapostemon radiatus made solitary nests that were scattered and well
hidden in the grassy lawn. Five of the six excavated nests of A. radiatus
yielded Nomada immatures, with percentages of infested cells for each nest
of 100% of 8 cells, 93% of 14 cells, 50% of 2 cells, 33% of 6 cells, and 25%
of 4 cells. The percentage of infested cells from all nests in this site was
63% of 40 cells; for infested nests only, 74% of 34 cells. Another nest of A.
radiatus, well separated from nests of other Agapostemon in another lawn,
yielded no Nomada.

We did not recover eggs of N. articulata, but they probably were placed
in the cell walls, as occurs in all other Nomadinae (Rozen, 1977). Nomada
edwardsii Cresson and N. opacella usually lay two eggs in each cell and
the larva that first hatches kills the other egg (Linsley and MacSwain, 1955).
The presumed oviposition period of N. articulata was half as long as the 4
to 6 min recorded for N. opacella, so double oviposition may not occur in
N. articulata. We recovered first-instar N. articulata larvae on two provi¬
sion masses. One larva apparently was in the act of consuming the host egg
when we broke into its cell. The larva was situated parallel to and partially
on top of the egg (Fig. 3). We brought the provision mass into the laboratory
and provided another host egg. The mobile first-instar Nomada also killed
this egg. All other N. articulata larvae were on provision masses without
host eggs or larvae, so killing host eggs is typical of the first instar of this
species, as it is of most Nomadinae (Rozen, 1977). Later instars (Fig. 4)
perched on top of and consumed provision masses in the manner illustrated
for other Nomada by Linsley and MacSwain (1955).

Mature N. articulata larvae from three nests defecated and then became
quite turgid, resting on their dorsa in the cells with their elevated heads
towards the cell mouths (Figs. 5-6), as illustrated for N. suavis Cresson by
Bohart (1970). They presumably would have spent the winter as postdefe¬
cating larvae (prepupae), pupated in the spring, and emerged as adults along
with their hosts in late May or early June. This developmental pattern is
typical for Nomadinae (Rozen, 1977). In the laboratory, these prepupae

VOLUME 56, NUMBER 2

149

entered diapause and did not further develop unless they were refrigerated
for several months. This cold treatment broke diapause, and development
resumed when the bees were brought to room temperature.

However, N. articulata immatures from an A. radiatus nest excavated
on 25 July 1978 did not follow this pattern. Ten were already pupae when
the nest was excavated, and development of those that survived proceeded
immediately to the adult stage without refrigeration. Three adult N. arti¬
culata were found in nests of A. virescens in October. Overwintering of
adults was hypothesized for Nomada parasitizing vernal Andrena by Lins-
ley and MacSwain (1955) and by Thorp (1969). Our data show that N.
articulata from the same population and indeed the same nest site may
overwinter both as prepupae and as adults.

Nomada (Micronomada) formula Viereck

Agapostemon texanus nests were aggregated, although each was believed
to be occupied by a single female. The population was probably multi vol-
tine, with provisioning for the last generation occurring in late August when
adult N. formula were also present at the nest site. Flight activity of parent
hosts and of cleptoparasites had ceased by late September, when nest bur¬
rows were partially filled with soil and no live parent females were in the
nests. Most of the host cells were deserted at this time; a few contained
older host pupae. Some newly emerged adult female hosts were in burrows
or in hibernacula similar to those of A. virescens (Abrams and Eickwort,
1980a).

All Nomada recovered during these excavations were turgid postdefe¬
cating larvae. As with N. articulata, they were on their dorsa with their
elevated heads towards the cell mouths. The feces appeared to be plastered
on all the cell walls, although denser posteriorly. These feces often were
separated from the cell walls and appeared to form a loose, amorphous,
cocoon-like structure around the larva, the cylindrical fecal pellets being
held together by silk-like strands (these may have been fungal mycelia).
This loose configuration may have been due to the low soil moisture. Ste¬
phen, Bohart, and Torchio (1969) noted that Nomada larvae speckle cell
walls with fecal pellets oriented in various directions.

No pupal or adult N. formula were recovered during the excavations, so
that population was overwintering as diapausing prepupae. Several prepu¬
pae were refrigerated in the spring of 1976 and these moulted into pupae
and then adults when removed to room temperature.

It was impossible to estimate parasitism rates in the largely deserted nests
of A. texanus, although all five burrows traced to cell level appeared to lead
to some cells that contained Nomada prepupae. Twelve Nomada prepupae

150

PAN-PACIFIC ENTOMOLOGIST

were recovered in total. In contrast, a large isolated nest of A. texanus
located several hundred meters from the aggregation contained no Nomada.

Discussion

The Nomada that attack Agapostemon behave very much like those that
attack Andrena, despite the differences in host biology and phylogenetic
relationship, and despite the fact that Agapostemon is a basically Neotrop¬
ical genus whose species have been in contact with species of the Holarctic
genus Nomada for presumably a relatively short period of evolutionary
time. Indeed, Andrena nests were abundant in the nesting areas of A. ra-
diatus and A. virescens, and quite possibly N. articulata is primarily an
Andrena parasite that enters Agapostemon nests when it encounters them.
Several European species of Nomada (i.e., N.fucata Panzer, rufipes Fa-
bricius, and zonata Panzer, see Stoeckhert, 1933) have been recorded as
parasites of both halictines and Andrena.

Tengo and Bergstrom (1977) have analyzed the cephalic secretions of
several species of Nomada and found an interesting pairwise correlation
between the secretions of the male (but not the female) cleptoparasites and
the Dufour’s gland secretions of the females of their host species of An¬
drena. The male Nomada coats his mate with this secretion during copu¬
lation so she has the same odor as the host nest. Presumably the female is
then able to enter a nest without being attacked by the host female. How¬
ever, if some Nomada do indeed attack hosts belonging to more than one
family, then either chemical mimicry does not occur in these species of
Nomada or host species belonging to different genera produce similar chem¬
icals in their Dufour’s glands.

The behavior of N. articulata at nests of A. virescens and the heavy
parasitism of some nests of A. radiatus and A. texanus by Nomada while
others were untouched indicates that the Nomada learned the locations of
nests and repeatedly returned to them. Such learning is apparently wide¬
spread in cleptoparasitic bees, as reviewed by Rozen, Eickwort, and Eick-
wort (1978). We hypothesize that N. articulata were “traplining,” visiting
each remembered nest in turn and, when a host female was not present,
pausing long enough to check for cells being provisioned.

Solitary Agapostemon radiatus and A. texanus appear to be defenseless
against such a strategy once their nests are discovered. Their only effective
counter-strategies are to conceal their nest entrances more effectively and,
especially, to build nests distant from those of other potential hosts. In
contrast, communal Agapostemon virescens escape parasitism by Nomada
through effective nest guarding, and only the occasional solitary nest of this
species falls prey to the cleptoparasite (Abrams and Eickwort, 1980b).

VOLUME 56, NUMBER 2

151

A final point is the overwintering of N. articulata from the same popu¬
lation in two different stages, as prepupa and as adult. Similar develop¬
mental flexibility has been observed in several species of Osmia (Megach-
ilidas) (Levin, 1966; Rust, 1974).

Acknowledgments

We thank F. Robert Wesley and Patricia Lockwood of Cornell University
for assistance in excavating Agapostemon radiatus nests and for lovingly
rearing immature stages of Nomada articulata. Mr. Roy Snelling of the Los
Angeles County Museum graciously identified the Nomada. Dr. Jerome
Rozen, Jr., of the American Museum of Natural History, Dr. Robbin Thorp
of the University of California at Davis, and Ms. Barbara Taraday of Cornell
critically read the manuscript. We thank the East Bay Municipal Utility
District for permitting excavations of nests of A. texanus at San Pablo
Reservoir. G.C.E. acknowledges the hospitality of the Division of Ento¬
mology and Parasitology of the University of California at Berkeley, where
he was a research associate while conducting field studies on A. texanus,
and the Department of Entomology of the University of California at Davis,
where he was a visiting professor during the preparation of the manuscript.
Observations on adult behavior of N. articulata were adapted from the M.S.
thesis of J.A. This research was supported by National Science Foundation
grant nos. BMS-72-02386 and DEB-78-03151.

Literature Cited

Abrams, J. 1977. The biology of the communal sweat bee Agapostemon virescens (Hyme-
noptera: Halictidae) in New York State. M.S. thesis, Cornell University, vii & 74 pp.
Abrams, J., and G. C. Eickwort. 1980a. Biology of the communal sweat bee Agapostemon
virescens (Hymenoptera: Halictidae) in New York State. Search (Cornell Univ. Agr.
Exp. Sta.), in press.

Abrams, J., and G. C. Eickwort. 1980b. Nest switching and guarding by the communal sweat
bee Agapostemon virescens (Hymenoptera: Halictidae). Insectes Soc., in press.
Bohart, G. E. 1970. The evolution of parasitism among bees. Utah State Univ., Logan,
30 pp.

Levin, M. D. 1966. Biological notes on Osmia lignaria and Osmia californica (Hymenoptera:

Apoidea, Megachilidae). J. Kans. Entomol. Soc., 39:524-535.

Linsley, E. G., and J. W. MacSwain. 1955. The habits of Nomada opacella Timberlake with
notes on other species (Hymenoptera: Anthophoridae). Wasmann J. Biol., 13:253-276.
Linsley, E. G., J. W. MacSwain, and R. F. Smith. 1954. A note on the nesting habits of
Exomalopsis solani Cockerell (Hymenoptera, Anthophoridae). Pan-Pacific Entomol.,
30:263-264.

Roberts, R. B. 1969. Biology of the bee genus Agapostemon (Hymenoptera: Halictidae).
Univ. Kans. Sci. Bull., 48:689-719.

Rozen, J. G., Jr. 1966. The larvae of the Anthophoridae (Hymenoptera, Apoidea). Part 2. The
Nomadinae. Amer. Mus. Novitates, 2244:1-38.

152

PAN-PACIFIC ENTOMOLOGIST

Rozen, J. G., Jr. 1977. Immature stages of and ethological observations on the cleptoparasitic
bee tribe Nomadini (Apoidea, Anthophoridae). Amer. Mus. Novitates, 2638:1-16.

Rozen, J. G., Jr., K. R. Eickwort, and G. C. Eickwort. 1978. The bionomics and immature
stages of the cleptoparasitic bee genus Protepeolus (Anthophoridae, Nomadinae). Amer.
Mus. Novitates, 2640:1-24.

Rust, R. W. 1974. The systematics and biology of the genus Osmia, subgenera Osmia, Chal-
cosmia, and Cephalosmia (Hymenoptera: Megachilidae). Wasmann J. Biol., 32:1-93.

Stephen, W. P., G. E. Bohart, and P. F. Torchio. 1969. The biology and external morphology
of bees. Agr. Exp. Sta. Oregon State Univ., Corvallis, 140 pp.

Stoeckhert, F. K. 1933. Die Bienen Frankens (Hym. Apid.). Deut. Entomol. Z. 1932. Beiheft,
vii & 294 pp.

Stoeckhert, F. K. 1954. Fauna Apoideorum Germaniae. Abhandl. Bayer. Akad. Wiss. n. f.,
65:1-87.

Tengo, J., and G. Bergstrom. 1977. Cleptoparasitism and odor mimetism in bees: do Nomada
males imitate the odor of Andrena females? Science, 196:1117-1119.

Thorp, R. W. 1969. Systematics and ecology of bees of the subgenus Diandrena (Hymenop¬
tera: Andrenidae). Univ. Calif. Publ. Entomol., 52:1-146.

Footnotes

1 Nomada articulata was erroneously identified as Nomada australis Mitchell in Abrams
(1977).

NOTICE

AWARDS FOR STUDY AT
The Academy of Natural Sciences,

Philadelphia, Pa.

The Academy of Natural Sciences of Philadelphia, through its Jessup
and McHenry funds, makes available each year a limited number of awards
to support students pursuing natural history studies at the Academy.
Awards usually include a stipend to defray living expenses, and support for
travel to and from the Academy. Current application deadlines are 1 April
and 1 October 1980. Further information may be obtained by writing to:
Chairman, Jessup-McHenry Award Committee, Academy of Natural Sci¬
ences of Philadelphia, 19th and the Parkway, Philadelphia, Pennsylvania
19103.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 153-154

DASYMUTILLA PHOENIX (FOX), A NEW
SYNONYM OF D. FOXI (COCKERELL)
(HYMENOPTERA: MUTILLIDAE)

Donald G. Manley

Department of Entomology & Economic Zoology, Clemson University,

Clemson, South Carolina 29631

The species Dasymutilla foxi was first described by Cockerell (1894) in
the genus Sphaerophthalma (sic) later transferred to the genus Dasymutilla
by Mickel (1928). This was followed by a first description of the species
Dasymutilla phoenix by Fox (1899). In reviewing the genus, Mickel (1928)
stated that the two species are not different structurally from one another
but that D. phoenix is larger in size than D. foxi and that the pubescence
of the dorsum of D. phoenix is whitish as opposed to scarlet pubescence of
the latter. Inclusion of the following biological evidence and a good series
of specimens from the same location has led to the conclusion that D.
phoenix should no longer be considered as a valid species but is, instead,
a synonym of D. foxi.

Materials and Methods

A series of 129 specimens, including 85 females and 44 males (of which
8 pairs were taken in copula ), were collected from a mating aggregation on
Willow Springs Ranch, north of Tucson, Arizona (described separately by
Manley and Taber, 1978). All were collected between September 18, 1976
and September 29, 1976. Of these, 31 specimens (16 females, 7 males, and
four pairs taken in copula ) have been deposited in the Entomology Museum
of the University of Arizona and 98 (61 females, 28 females, plus 4 females
and 5 males taken in copula ) have been retained in the personal collection
of the author.

All of the specimens collected were determined as belonging to the D.
foxi-phoenix complex. All were active in the same area, at the same times.
It seemed impossible to distinguish two species on the basis of behavior.

Results and Discussion

Since the major criteria for distinguishing these two as valid species have
been coloration and size, these criteria were examined more closely in the
laboratory. The following were found:

154

PAN-PACIFIC ENTOMOLOGIST

Males: Of 44 males that were collected, pubescence of the dorsum of the
head and thorax of 43 was a smokey yellow (“dirty white”), with the ab¬
domen being scarlet. On the remaining male, head, thorax and abdomen
were all scarlet. The latter male was one of those taken in copula, and was
coupled with a female having the smokey yellow pubescence on head and
thorax, and scarlet abdomen. Males ranged in length from 8 mm to 13 mm.

Females: There was considerably more variation in color among the fe¬
males. The continuum ranged from smokey yellow to scarlet. The color
variation has, however, been categorized for descriptive purposes. The pu¬
bescence above for the majority (61) of the 85 females was smokey yellow
(head, thorax, and abdomen). Of the remaining 24, 23 had head and thorax
colored smokey yellow. Of those, the abdomens of 7 were of an orange-
yellow color, 6 were orange, 4 were a red-orange, and 6 were scarlet. Pu¬
bescence of the dorsum of the remaining female was entirely scarlet (head,
thorax, and abdomen). Length in the females ranged from 7 mm to 12 mm
and seemed to be entirely independent of color.

If these were separate species, one would expect to find more individuals
with the colors of the two extremes, with only a few intermediates. The
actual situation was one of many more individuals with the smokey yellow
pubescence of D. phoenix, with a few variants ranging all the way to the
scarlet extreme. It is thus concluded that the two should be considered as
one species, D.foxi, with the updated synynomy being as follows:

Dasymutilla foxi (Cockerell)

Sphaerophthalma (sic) foxi Cockerell 1894:199. 6
Sphaerophthalma (sic) heterochroa Cockerell and Casad 1894:298. 9
Mutilla phoenix Fox 1899:247. NEW SYNONYM. 9
Sphaerophthalma (sic) foxi var. arizonica Cockerell 1900:65. 9
Dasymutilla foxi Mickel 1928:84. Sex not stated.

Dasymutilla phoenix Mickel 1928:85. Sex not stated.

Literature Cited

Cockerell, T. D. A. 1894. Descriptions of new Hymenoptera. Entomol. News, 5:199.
Cockerell, T. D. A. 1900. New insects from Arizona, etc. Entomologist, 33:65.

Cockerell, T. D. A., and J. B. Casad. 1894. New species of Mutillidae. Entomol. News,
5:293-98.

Fox, W. J. 1899. American Mutillidae. Trans. Am. Entomol. Soc., 25:247.

Manley, D. G., and S. Taber, III. 1978. A mating aggregation of Dasymutilla foxi in Southern
Arizona. Pan-Pac. Entomol., 54:231-235.

Mickel, C. E. 1928. Biological and taxonomic investigations on the mutillid wasps. U.S. Nat.
Mus. Bull., 143:84-87.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 155-156

SCIENTIFIC NOTE

ENTOMOBRYA KAN ABA (WRAY) (COLLEMBOLA:

ENTOMOBRYIDAE) AN INDOOR HOUSEHOLD
PEST IN CENTRAL CALIFORNIA

Entomobrya kanaba (Wray) is here reported as an indoor household pest
in central California. Maynard (1951, A monograph of the Collembola or
springtail insects of New York State, p. 14) states “It is only rarely that
Collembola are reported as household pests, and then usually the damage
is in the aesthetic sense.” Curran (1947, Nat. Hist., 56(10):476, 1 fig.) has
pointed out that the presence of springtails can be a sign that there is excess
moisture in the building that should be eliminated.

During the fall months of 1977 Mr. Earl Enos, a Custodian at the Wilson
Elementary School, in San Leandro, Alameda County, California, repeat¬
edly observed small insects in the classroom sink of the kindergarten room.
Often several dozen specimens would be seen in the early morning around
the inside of the white enamel sink (with only a few on the drainboard).
They were not noticed in other classrooms. Finally, as the identity of these
unknown jumping insects was wanted, Mr. Enos collected a sample of 17
specimens and they were submitted to Mrs. Mary A. Davies to have them
identified. The writers forwarded the specimens to Dr. David L. Wray,
Collembolist and Entomologist Emeritus of the North Carolina Department
of Agriculture, who identified them as Entomobrya kanaba (Wray) (depos¬
ited in the CAS and Wray collections).

Interestingly, in addition to being observed at the Wilson Elementary
School, in San Leandro, Mrs. Leslie Sweeny, the kindergarten teacher,
stated that she had noted Collembola in her home in Fremont, California
over a period of three or four years. Also, Mrs. Cyndi Rose, a next door
neighbor to Mrs. Sweeny, had a very troublesome infestation of Collembola
throughout her home. Samples of the Collembola in Mrs. Rose’s home were
made (2 specimens on 29-IX-1977 and 60 specimens on 5-X-1977) and these
were submitted to Dr. Wray, and he also identified them as Entomobrya
kanaba (Wray). Samples were not collected in Mrs. Sweeny’s home, but it
is possible that they were also this species.

The infestation in Mrs. Rose’s home was throughout the entire building.
The collection made on September 29th was from the back of a bath¬
room shower curtain and this curtain was described as being “black” with
Collembola. These Collembola were also stated to be in every sink, basin,
tub, in the pots and pans in the kitchen, and in the clothes closets. An
attempt to control the infestation with an insecticide was made (in this
instance RAID was used) but it was not successful. A pest control operator

156

PAN-PACIFIC ENTOMOLOGIST

was hired to control the infestations in both the homes of Mrs. Rose and
Mrs. Sweeny at a cost of $40.00 per home. The infestation of Collembola
at the home of Mrs. Sweeny was minor, however, but there was a problem
with silverfish that she wished controlled.

Entomobrya kanaba was described by D. L. Wray (1953, Nature Notes,
Occ. Pap. No. 1:4, fig. 2, A-F, in the genus Drepanura ) on the basis of a
large series taken by Dr. G. F. Knowlton at Kanab, Utah on June 15-16,
1952. They were collected “on a tile floor of auto camp bath room, on the
sidewalk, and under plank.” Knowlton states that “in the early morning of
June 16 they were graying the ground by the thousands.” About one thou¬
sand specimens were obtained. It would appear that their large numbers
about dwellings may lend to their dispersal by artificial means. Wray and
Knowlton (1956, Great Basin Nat., 16(l-4):4) also record Entomobrya kan¬
aba from the state of Idaho from two collections—from shade tree litter and
boxelder litter. Salmon, in his world catalog of Collembola (1964, Roy. Soc.
New Zealand, Bull. 7 (vol. 2):440), does not list further references or dis¬
tribution for this species. Christiansen (1956, Ent. News, 67(5): 129-130;
1958, Bull. Mus. Comp. Zool., 118(7):469) considers kanaba to a synonym
of Entomobrya unostrigata Stach (1930, Abhand. Senckenberg. Naturf.
Ges., 42(1):63), and provides records under the latter name from San Diego,
California and Fort Collins, Colorado. Stach (1963, Polska Akad. Nauk Inst.
Zool., Cracow, p. 70) discusses unostrigata and kanaba and indicates some
differences.

Collembola are probably present in small numbers in homes or offices,
particularly if indoor plants are kept, and they may often go unnoticed. For
example, Lepidocyrtus cinereus Folsom was recently collected on 29-IX-
1977 in a pot planter in an office (PHA) at the California Academy of Sci¬
ences, San Francisco, where it is an inconspicuous species. In contrast,
Entomobrya kanaba with other habits, probably associated with drains, has
been shown to be a household pest.

Our thanks are extended to Dr. D. L. Wray for his identification of the
Collembolan collections and to Mrs. Mary A. Davies, Mr. Earl Enos, Mrs.
Cyndi Rose, and Mrs. Leslie Sweeny for specimens and information con¬
cerning the infestations, and to the reviewers of this note.

Paul H. Arnaud, Jr. and Thomas W. Davies, California Academy of Sci¬
ences, Golden Gate Park, San Francisco 94118.

PAN-PACIFIC ENTOMOLOGIST
April 1980, Vol. 56, No. 2, pp. 157-160

SCIENTIFIC NOTE

NOTES ON THE LIFE HISTORY OF THREE CONODERUS SPECIES
OF WIREWORMS IN CALIFORNIA: (COLEOPTERA: ELATERIDAE)

Information on the duration of the larval period of three introduced Con-
oderas species was obtained by confining newly hatched larvae individually
in two ounce salve tins containing screened moist soil and a surplus of
wheat, both of which were replenished at two week intervals. The larvae
were confined indoors where temperatures ranged from 15° to 25°C (60° to
78° F).

Conoderus exsul (Sharp)

Two separate rearings of Conoderus exsul were started in 1978. The first,
or early group, included 31 larvae which hatched March 19. Of these, 20,
or 65%, pupated between July 6 and November 11 of the same year, for an
average larval period of 149 days. The remaining 11 larvae pupated the
following year between January 18 and March 19, for an average larval
period of 331 days. Stone (1976, Pan-Pac. Entomol., 52:304) showed that
16% of June hatched larvae had matured the same year and 84% the second
year. The pupal period in this early matured group ranged from 10 to 19
days, averaging 11.7 days. Lower temperatures the second year prolonged
the pupal period from 18 to 22 days (Tables 1 and 2).

Larvae which hatched on July 25, 1978, or approximately 4 months later
than the above, were used in the second rearing experiment. Of the 53 larvae
involved, 13 pupated the same year between October 3 and 25 for an average
larval period of 80 days. All remaining larvae pupated the following year
between February 28 and May 29, the larval period averaging 242 days. The
pupal period for the early and late maturing groups averaged 13 and 17.3
days, respectively.

Conoderus amplicollis (Gyll.) (Gulf Wireworm)

This species was reared in the manner described previously. Of 16 larvae
that had hatched on July 20, 1978, none matured the first year. All pupated
the following year over a period of 3.5 months from March 23 to July 1.
Their larval period averaged 305 days and their pupal period, 14.3 days.

In a 1979 paper, (Stone & Wilcox, Pan-Pac. Entomol., 55:235-238) “The
Gulf Wireworm in California,” ten of 19 larvae pupated the same year,
which was due, perhaps, to their May 5 hatching date.

L/t

Table 1. Duration of the larval stage of three introduced Conoderus species, Riverside, CA. 1978-79.

Date hatched

1978

Records

(no.)

Larvae completing development

1978 Larval period

1979 Larval period

Pupations

(%)

Range

(days)

Average

(days)

Pupations

(%)

Range

(days)

Average

(days)

Conoderus exsul

Mar. 19

109-237

149

305-365

331

July 25

70-92

199-295

242

Conoderus amplicollis

July 20

100

247-347

305

Conoderus falli

Aug. 16

80-106

229-270

259

Oct. 9

62-70

183-291

258

PAN-PACIFIC ENTOMOLOGIST

Table 2. Duration of pupal stage of Conoderus species, Riverside, CA. 1978-79.

Larvae completing development in

1978 Pupal period

1979 Pupal period

Date hatched

1978

Records

(no.)

Range

(days)

Average

(days)

Records

(no.)

Range

(days)

Average

(days)

Conoderus exsul

Mar. 19

10-19

11.7

18-22

19.5

July 25

11-17

14-20

17.3

Conoderus amplicollis

July 20

11-19

14.3

Conoderus falli

Aug. 16

12-19

16.3

16-17

16.3

Oct. 9

21-22

21.5

10-17

11.4

VOLUME 56, NUMBER 2

160

PAN-PACIFIC ENTOMOLOGIST

Conoderus falli Lane (Southern Potato Wireworm)

Of 19 C. falli larvae that hatched August 16, 1978 reared on wheat, 8
larvae died in the larval stage. Seven of the remaining 11 larvae pupated
between November 4 and 30 the same year, for an average larval period of
90 days. The 4 remaining larvae pupated in 1979 between April 1 and May
12. Their larval period averaged 259 days. The duration of the pupal period
for both groups was 16.3 days.

Additional data on the life span of C. falli was obtained in 1978 by rearing
a group of 20 larvae that had hatched later than usual, on October 9. It is
interesting to note that even at this delayed hatching date two larvae pupated
on December 10 and 18, for an average larval period of 66 days. The re¬
maining 18 larvae pupated the following year over a period of 4 months from
April 9 to July 26. Their larval period ranged from 183 to 291 days, averaging
258 days.

The duration of the pupal period varied with the temperature, from an
average of 21.5 days in December to 11.4 days for those maturing the second
year during warmer months.

Discussion

Rearing these species of Conoderus wireworms indoors, possibly under
the most favorable of conditions, one might expect the larvae to develop
more rapidly and mature earlier than would be the case for larvae inhabiting
dry, weedy fields or soils lacking in suitable food. Conditions closer to the
indoor rearing conditions would prevail in irrigated vegetable growing areas
planted to green or dry beans or to other crops more palatable for their
survival.

As judged by the thousands of beetles attracted to a black light at Riv¬
erside, both C. exsul and C. falli have found climatic conditions and food
very favorable. Rearings of larvae of these species showed that, depending
upon the time of beetle emergence and egg deposition, from 10 to 65 percent
of the larvae may complete development the same year, the remainder the
second year. C. exsul larvae also appear to have found conditions favorable
for survival at Olive, in an avocado grove, where the annual collection of
beetles exceeded that obtained at Riverside. However, of interest, was the
fact that no C. falli adults were collected at this location in the 5 year period
1975-79.

Adults of C. amplicollis are not highly attracted to light, nevertheless
their presence in 15 counties is an indication that they too have become well
established in California soils.

M. W. Stone ( Collaborator—USDA SEA AR. Boyden Entomological
Laboratory), 131 Sir Damas Dr., Riverside, California 92507.

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