Vol. 48 JANUARY 1972 No. 1

THE

Pan-Pacific Entomologist

HALSTEAD — A review of the genus Arhaphe Herrich-Schaffer (Hemiptera:

Largidae) 1

FISHER AND ORTH — Resurrection of Sepedon pacifica Cresson and Re-
description of Sepedon praemiosa Giglio-Tos with biological notes
(Diptera: Sciomyzidae) 8

RASKE — Immature forms, genitalia, and notes on the biology of Anelaphus

albofasciatus Linnell (Coleoptera: Cerambycidae) 21

SCHROETER AND HEWITT — Cytology of some California grasshoppers.

1. Taxonomic considerations (Orthoptera: Acridoidea) 27

HALL — New North American Heterotropinae (Diptera: Bombyliidae) 37

WILCOX — The genus Sintoria Hull (Diptera: Asilidae) 51

OBITUARY NOTICE 59

BOOK REVIEW 61

SCIENTIFIC NOTES 63

ZOOLOGICAL NOMENCLATURE 65

PROCEEDINGS 66

SAN FRANCISCO, CALIFORNIA • 1972

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

THE PAN-PACIFIC ENTOMOLOGIST

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The Pan-Pacific Entomologist

Vol. 48 January 1972 No. 1

A Review of the Genus Arhaphe Herrich-Schaffer

(Hemiptera: Largidae)

Thomas F. Halstead

Arizona Commission of Agriculture and Horticulture, Phoenix 85005

Herrich-Schaffer originally described the genus from “Carolina” in
the form of Arhaphe Carolina. In 1873 Walker described A. cicindeloides
from Mexico; and in 1911 Barber united Japetus Distant with Araphe
(apparently following Uhler’s 1872 misspelling of the generic name)
and described A. mimetica , thus including /. sphaerodes in Araphe. In
1924 Barber described A. breviata from Kansas and again separated
Japetus from Arhaphe , including J. sphaerodes Distant and J . mimeticus.
In 1956 Bliven added Jarhaphetus argutus. This paper again unites the
two genera, places Jarhaphetus Bliven as a synonym of Arhaphe , and
describes two new species.

The set of characteristics which sets this genus apart from other
Largidae is the combination of the absence of any pronotal or thoracic
spination and the possession of a stridulatory apparatus consisting of
ridges or corrugations along the edge of the corium with accompanying
rasp on hind femora.

Arhaphe Herrich-Schaffer

Arhaphe Herrich-Schaffer, 1850: IX: 175 and 183.

Araphe, Uhler, in Hayden, 1872 : 471.

Japetus Distant, 1883: 227; Barber, 1924: 227.

Arrhaphe, Bergroth, 1913: 166.

Jarhaphetus Bliven, 1956: 10. (New Synonymy) .

Key to Species of Arhaphe

1. Head and prothorax coarsely punctate 2

Head impunctate (prothorax impunctate except in A. capitata) 4

2. Membrane white with central black spot mexicana Halstead, n. sp.

membrane with base white and tip apparently black 3

3. Entire corium brick-red sphaerodes Distant

Corium yellow to white with medial black fascia Carolina Herrich-Schaffer

4. Membrane very much reduced, if present at all represented only by narrow

band 5

Membrane entire 6

The Pan-Pacific Entomologist 48: 1-7. January 1972

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

5. Row of puncta along claval suture entirely immersed in black band; pos-

terior femora almost reaching abdominal apex cicindeloides Walker

Row of puncta along claval suture black, each surrounded by white of
corium; posterior femora not extending beyond fourth abdominal seg-
ment breviata Barber

6. Head level between and behind eyes mimetica Barber

Head rising to an elevated knob between and behind eyes

capitata Halstead, n. sp.

Arhaphe CAROLINA Herrich-Schaffer

Arhaphe Carolina Herrich-Schaffer, 1850: IX: 183, tab. 315, fig. 968.

Araphe Carolina Barber, 1911: 28.

Diagnosis. — Known from North Carolina south to Florida, thence
west through Tennessee to Arizona and Baja California. Cited from
“Mexico” by Stal (1870). Easily distinguished from other punctate
forms by coloration of corium and by characters in key. Length
7-10 mm.

Arhaphe cicindeloides Walker

Arhaphe cicindeloides Walker, 1873: VI: 36.

Jarhaphetus argutus Bliven, 1956: 10. (New Synonymy) .

Diagnosis. — Known from Arizona and New Mexico, thence south
into Mexico at least as far as Sinaloa. Large and black, elongate oval,
head and prothorax impunctate, variously pubescent. Corium white
with central broad black fascia from side to side. Anterior femora armed
with single more or less obvious tooth near apex. Head as broad as or
broader than pronotum. Length 9-12 mm. Bliven’s description and
photograph of Jarhaphetus argutus clearly match A. cicindeloides.
Barber is the only one to mention or use the femoral tooth as a char-
acter in separation of the genera, and he must have had individuals of
A. cicindeloides in which the spines were very small to have missed
them. Bliven based his new genus and species on presence of the spine
and width of the head, both very variable in this species.

Arhaphe sphaerodes (Distant)

Japetus sphaerodes Distant, 1883: 227; Barber, 1924: 227.

Araphe sphaerodes Barber, 1911: 28.

Diagnosis. — The following description of the unique type deposited
in the British Museum (Natural History) by Distant was kindly fur-
nished by Mr. W. R. Dolling of that institution: Entirely black except
hemelytra. Thorax, abdomen and legs fine silvery recumbent pubescent.

JANUARY 1972] HALSTEAD REVIEW OF ARHAPHE

3

Head, thorax and femora fine sparse erect pubescent. Head shining,
coarsely reticulate sculptured. Many interstices of sculpturing bearing
single large punctures, each separated from its neighbors by two-thirds
to one and one-half times its own width. Hemelytra reaching about %e
the distance from pronotal posterior to abdominal apex. Corium en-
tirely and uniformly clear brick-red. Angle of clavus between pronotum
and scutellum infuscate, sharply marked off from rest of clavus, clavus
concolorous with corium except at its inner edge, where fuscus color
runs along innermost line of punctures and fades out just beyond
scutellar apex. Basal third of each membrane ivory white; apical two-
thirds dark brown. A narrow ivory band borders apical margin; but
resulting diamond-shaped spot appears as dark apex, not dark mark in
middle, as apical marginal band very narrow. Ratio of antennal seg-
ments 45:35:22:50 (basal first, arbitrary units). Basal segment of
posterior tarsus accounting for three-fifths of its length. Ratio of head
width ( across eyes) to anterior thoracic lobe to posterior thoracic lobe
33:23:26% (different units than above). Rostrum just surpassing an-
terior coxae. Length 9 mm.

Type.— S. Geronimo, Guatemala, Champion.

Arhaphe mimetica (Barber)

Araphe mimetica Barber, 1911: 28.

Japetus mimeticus Barber, 1924: 227. New Synonymy.

Diagnosis. — Known from the Huachuca, Santa Rita and Atascosa
Mountains of southern Arizona, but very probably more widespread
into south and east. Encountered in oak leaf litter in numbers at Madera
Canyon, Arizona, in August. Well differentiated from other species by
characters in key. Length 6-8 mm.

Arhaphe breviata Barber
Arhaphe breviata Barber, 1924: 227.

Diagnosis.— Known only from Kansas. Well differentiated from A.
Carolina by absence of head and anterior pronotal punctation and from
A. cicindeloides by characters in key. Length 7.3-8.25 mm.

Arhaphe mexicana Halstead, new species

(Fig. 1)

Diagnosis. — Known from two male specimens taken near Cuernavaca,
Morelos, Mexico. Very distinct from A. sphaerodes , the most similar
species in genus.

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THE PAN-PACIFIC ENTOMOLOGIST

[VOL. 48, NO. 1

JANUARY 1972] HALSTEAD REVIEW OF ARHAPHE

5

Length of holotype male 8.3 mm (paratype 8.3 mm), width across head (in-
cluding eyes) 1.9 mm (paratype 1.9 mm), ratio of head to anterior thoracic lobe
to posterior thoracic lobe 24:17:21. Width across corium 1.7 mm (paratype
1.7 mm) .

Male. — Linear elongate, black. Head, entire thorax and scutellum coarsely
punctate. Corial margins with line of punctures; scattered punctation on re-
mainder. Clavus with marginal lines of puncta; diagonal line of puncta from
mid-anterior margin to apex. Anterior femora armed with minute spine near
apex. Entire body except corium covered with fine closely appressed silver
tomentum, this absent on large circular patches of sides of third through sixth
ventral segments and anterior margins of abdominal dorsal segments, appearing
as black spots on silver background. Head and thorax sparsely covered with long
black hairs. Corium pale orange with black line along apex and circular black
spot in middle. Clavus pale orange with black triangle, wider at base and fading-
out to apex, along its inner margin. Rostrum reaching iust beyond anterior coxae.
Antennal ratio 27:23:15:31 (basal first, different units than above), reaching
somewhat beyond pronotal posterior, first segment somewhat curved.

Female. — Unknown.

Holotype male , 3-6 mi. S. Cuernavaca, Morelos, Mexico. 1 April
1959. 4,000 ft. H. E. Evans (Cornell type number 4609).

Paratype male. — Cuernavaca, Morelos, Mexico. 17 March 1959.
5,000 ft. H. E. Evans and D. M. Anderson collectors. Deposited in the
author’s collection.

Arhaphe capitata Halstead, new species

Diagnosis. — Known from two male specimens taken in Morelos,
Mexico. Similar in appearance to A. mimelica but differing by scattered
punctation on pronotal anterior, elevated knob between and behind eyes,
and hemelytral membrane surpassing abdominal apex.

Length of holotype male 6.67 mm (paratype 6.9 mm) . Width across head
(including eyes) 1.9 mm (paratype 1.9 mm). Ratio of head to anterior thoracic
lobe to posterior thoracic lobe 22:17:23. Width across corium 1.9 mm.

Male. — Elongate, black. Head shining black, impunctate, with rather sparse
short fine white pubescence on sides and front, with rounded knob between and
behind eyes. Collar and prothoracic anterior black, sides sparingly punctate, silvery
pubescent. Prothoracic posterior velvety black, coarsely punctate, without pubes-
cence. Scutellum black, silvery pubescent, coarsely punctate. Ventrum black
except white posterior margin of metathorax, silvery pubescent, shining. Clavus
black, with three lines of punctation running from base to apex along margins and
in middle. Femora armed with minute spine. Corium white, with transverse
broad black fascia from side to side, row of black puncta along calaval suture,

<-

Fig. 1. Arhaphe mexicana Halstead, n. sp.

6

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

otherwise heavily punctate. Membrane white with very large central black or
dark brown circular spot; surpassing abdominal apex. Rostrum reaching halfway
to mesocoxae. Antennae slender, fourth segment brown ; lighter in color than
others. Antennal ratio 27:25:17:30 (basal first, different units than above), first
segment strongly curved.

F em ale . — U nknown.

Holotype male , Huajintlan, Morelos, Mexico. 14 May 1959.
2,500 ft. H. E. Evans (Cornell type number 4610).

Paratype male. — Same data, deposited in the author’s collection.

Discussion. — In 1911 Barber described A. mimetica and placed
Distant’s Japetus as a synonym of Arhaphe. In 1924 he re-erected
Japetus , including /. sphaerodes and /. mimetica. He separated Japetus
as follows, “Head more globose, as long as but considerably wider than
pronotum; membrane less abbreviated; body and legs without profuse
coating of long setae, almost nude; posterior lobe of pronotum not at
all or very sparsely tomentose; anterior femora armed with a single
small tooth near apex; rostrum short.” If an attempt is made to separate
specimens on this basis, one immediately runs into many incongruities.
Arhaphe mexicana , closest relative to A. sphaerodes , has a very heavily
tomentose posterior pronotal lobe and long setae on the legs and body.
Arhaphe cicindeloides does have anterior femoral spines, and the head
width is variable. The rostrum of the species apparently falling into
Japetus is shorter, usually only attaining the anterior coxae; but that of
A. capitata almost reaches the mesocoxae. The head of A. mexicana
is very flattened, that of A. mimetica less so, and that of A. capitata
even less — approaching A. cicindeloides in globosity. In addition, I
have a single as yet undescribed Mexican specimen of Arhaphe closely
resembling A. cicindeloides , but possessing hemelytra complete with a
fully developed membrane which surpasses the abdominal apex. At-
tempts to separate specimens by use of a length-to-width ratio result in
some males of A. Carolina being placed with Japetus. I have examined
slide mounts of the parameres of A. Carolina, A. cicindeloides, A.
mimetica, A. mexicana , and A. capitata and can find little of use in
specific, let alone generic, differentiation. These very similar parameres
do show a uniform variation from those of Largus and Stenomacra
species examined. From the above data it becomes clear that separation
cannot reasonably be continued, and members of this homogeneous
group must be placed in a single genus.

Acknowledgments

I wish to thank Dr. R. C. Froeschner, Mr. H. B. Leech, Dr. L. L.
Pechuman, and Dr. F. G. Werner for the loan of specimens; Mr. W. R.

JANUARY 1972 ] HALSTEAD — REVIEW OF ARHAPHE

7

Dolling of the British Museum (Natural History) for examining the
type of A. sphaerodes and sending me a description of that specimen;
Mr. J. A. Childers for the fine illustration; and my wife for typing the
manuscript.

Literature Cited

Barber, H. G. 1911. Descriptions of some new Hemiptera-Heteroptera. J. N. Y.
Entomol. Soc., 19: 23-31.

1924. The genus Arhaphe in the United States (Hemiptera-Pyrrhocoridae) .
Can. Entomol., 56: 227-228.

Bergroth, E. 1913. Supplementum catalogi Heteropterorum Bruxellensis, II:
Mem. Soc. Entomol. Belg., 22: 123-183.

Bliven, B. P. 1956. New Hemiptera from the western states with illustrations
of previously described species and new synonymy in the Psyllidae.
Occidental Entomol., Nov. 1956: 1-27.

Distant, W. L. 1880-1893. Biologia Centrali-Americana. Heteroptera I. London,
462 p.

Herrich-Schaffer, G. A. W. 1850. Die Wanzenartigen Insecten. IX. Nurnberg.
Stal, C. 1870. Enumeratio Hemiptorum. I. Kongliga Svenska Yetenskaps
Akademiens Handlingar. 9 (1).

Uhler, in Hayden. 1872. Preliminary report of the U. S. Geological Survey of
Wyoming.

Walker, F. 1873. Catalog of the specimens of Hemiptera Heteroptera in the
collection of the British Museum. VI. London.

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8 THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Resurrection of Sepedon pacifica Cresson and Redescription
of Sepedon praemiosa Giglio-Tos with Biological Notes

(Diptera: Sciomyzidae)

T. W. Fisher and R. E. Orth 1

Division of Biological Control, University of California, Riverside, 92502

Giglio-Tos (1893) based the original description of Sepedon
praemiosa on a single male from Morelia, Mexico, and later (1896)
elaborated on the description of the type. Neither verbal treatment is
adequate to distinguish S. praemiosa from similar species. The type
specimen is believed to be in the museum of the University, Turin, Italy.
However, unanswered correspondence makes confirmation impossible.

Cresson (1914) described S. pacifica from a male collected in Marin
Co., coastal central California. We have examined this specimen. Its
color is somewhat faded, but the terminalia agrees with our concept
of this taxon. Interestingly, Melander (1920) accepted Cresson’s
designation of S. pacifica. Apparently both workers were aware of
S. praemiosa only from the catalogue by Aldrich (1905), and assumed
from the type locality that it was a different species. It is doubtful
that either worker had seen the Giglio-Tos type, and they, therefore,
considered S. pacifica a distinct species. It is fortuitous that all of
the localities listed by Cresson and Melander fell within the area of
distribution we now ascribe only to S. pacifica.

Steyskal (1950) placed Sepedon relictus van der Wulp (v. d. Wulp,
1897) and S. pacifica Cresson (Cresson, 1914) in synonymy with
S. praemiosa. Through the courtesy of K. G. V. Smith, British Museum
of Natural History, we have examined the van der Wulp type specimen
which is a female (not a male as recorded in Biologia Centrali-
Americana) and have compared it with a number of females of S.
praemiosa taken at various localities from Oaxaca, Mexico, to Colorado,
U. S. A. and cannot assign it to S. praemiosa. We recommend that
S. relictus van der Wulp be removed from synonymy. The S. relictus
type fits very well the description of S. haplobasis Steyskal (Steyskal,
1960) , and we suspect that it is conspecific with that species. However,
at this writing (6 October 1971, revision of the original manuscript)
specimens of S. haplobasis are not at hand for comparison.

Although neither Steyskal nor we have seen the S. praemiosa type

1 Specialist and Staff Research Associate, respectively, Department of Entomology, Division of
Biological Control, University of California, Riverside, California 92502.

The Pan-Pacific Entomologist 48: 8-20. January 1972

JANUARY 1972] FISHER & ORTH SEPEDON SYSTEMATICS

9

4 5 6

Figs. 1-6. Sepedon praemiosa Giglio-Tos, U.S.A., Colorado, Fremont Co., 20
Aug. 1959 (N. Marston). Wing length of this specimen, 7.0 mm. Fig. 1. Termi-
nalia, sinistral view, inverted. Fic. 2. Aedeagus, sinistral view. Fig. 3. Aedeagus,
oblique anterior view from direction of arrow in Fig. 2. Fig. 4. Hypandrium,
sinistral view, inverted. Figs. 5-6. Hypandrium viewed from directions of arrows
in Fig. 4.

specimen, examination of a rather large amount of material from
western North America, including specimens from Morelia, which we
designate as topotypes of S. praemiosa, persuade us that the appelation
is correct.

We became aware of two forms of S. praemiosa in 1966, and since
have had at hand over 4,000 specimens which we assign to the taxon
S. pacifica and 267 specimens to S. praemiosa. Both are clearly sep-
arated by differences in distribution and in the morphology of primary

10

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

10 11 12

Figs. 7-12. Sepedon pacifica Cresson, U.S.A., Colorado, Ft. Collins, 16 April
1958 (N. Marston). Wing length of this specimen 8.3 mm. Fig. 7. Terminalia,
sinistral view, inverted. Fig. 8. Aedeagus, sinistral view. Fig. 9. Aedeagus,
oblique anterior view from direction of arrow in Fig. 8. Fig. 10. Hypandrium,
sinistral view, inverted. Figs. 11 and 12. Hypandrium viewed from directions of
arrows in Fig. 10.

and secondary sexual structures of the males. Although the specimen
drawn by Steyskal (1950, fig. 13) and labelled S. praemiosa cannot
now be located, that illustration agrees well with our concept of S.
pacifica.

Other than the material collected by us during the survey phase of
California Experiment Station Project No. 2037, “Biological Control
of Non-marine Mollusks,” material was kindly provided by C. 0. Berg

JANUARY 1972] FISHER & ORTH SEPEDON SYSTEMATICS

11

(Cornell University), N. Marston (Kansas State University), L. V.
Knutson and G. C. Steyskal (Systematic Entomology Laboratory,
USDA), P. H. Arnaud (California Academy of Sciences), W. F. Barr
(University of Idaho), and entomological collections of Illinois Natural
History Survey, Oregon State Department of Agriculture, South Dakota
State University, Texas A. & M., University of Minnesota, University
of Wisconsin, and the University of California at Berkeley, Davis, and
Riverside. Material from S. S. Roback (Academy of Natural Sciences,
Philadelphia) included the type of S. pacijica Cresson.

Comparison of S. pacijica and S. praemiosa reveals consistent differ-
ences in primary and secondary genitalic structures in the males (Figs.
1-12) . With the genitalia extruded — as with fresh material, or with
dried material after light boiling in 10% KOH — the lateral and oblique
anterior aspects of the aedeagus are diagnostic (Figs. 2, 3, 8, 9).

In general S. pacijica appears to be a larger and darker species
than S. praemiosa. The relative sizes of the sexes are also different.
In S. praemiosa the males are smaller than the females, but the reverse
occurs in S. pacijica. However, in proportion to their size, the male
genitalic structures of S. praemiosa are larger than those of S. pacijica.
This size relationship is illustrated by comparing the terminalia and
especially the hypandria of the two species (Figs. 4, 5, 6, 10, 11, 12 —
all drawn to the same scale) .

Neallotype, S. PACIFICA. — female, % mi. w. Valley Ford, Sonoma
County, California, 100 ft., 13 June 1966, AS-485 (T. W. Fisher &

R. E. Orth) , plus 20 California specimens were deposited in the Academy
of Natural Sciences, Philadelphia. In addition 20 California specimens
were deposited at the United States National Museum.

Topotypes, S. praemiosa. — 1 2,2 $, 8 mi. e. Morelia, Michoacan,
Mexico, 1 May 1953 (R. C. Bechtel & E. I. Schlinger). Specimens were
collected in sweep nets while sampling the edge of a bean and squash
field which was adjoined by a swampy river and marsh. 1 9,1 A
plus 10 additional specimens deposited in United States National
Museum. 1 $ topotype plus 10 specimens to the University of California
at Berkeley.

Variation. — Unlike most specimens seen, certain specimens from
several localized populations of S. pacijica in northern California,
Oregon, Washington, and Idaho possess prominent black parafrontal
spots. [We have also seen this variation in occasional specimens of

S. fuscipennis from Oregon, Washington, Idaho, and New Mexico.]

The ability to separate S. praemiosa from S. pacijica on genitalic

characteristics has made it possible to reevaluate certain superficial

12

THE PAN-PACIFIC ENTOMOLOGIST | VOL. 48, NO. 1

characters to the degree that such characters now assume a diagnostic
role of some importance. One such character pertains to the facies.
In S. praemiosa the color of the facies is much lighter, almost yellowish,
when compared to the darker, brownish face of S. pacifica. All the
S. pacifica we have seen have from few to many fine hairs on the
medifacies, but approximately 25% of the specimens of S. praemiosa
we have seen from several localities have no hairs on the medifacies.

Localities where barefaced S. praemiosa were collected include :
Mexico. — Durango (Durango); Michoacan (Morelia). U.S.A . — Ari-
zona (Lake Mary nr. Flagstaff) ; Colorado (Las Animas and Walsen-
burg) ; Nebraska (North Platte) ; New Mexico (Las Cruces, Bob
Crosby Draw) .

Because of the occurrence of a rather large percentage of barefaced
S. praemiosa , it now seems advisable to utilize a key character other
than “medifacies with fine black (scattered) hairs” to lead to this
species, or if key characters currently in use are perpetuated, mention
should be made of the rather common occurrence of glabrous medi-
facies in S. praemiosa in the same couplet wherein hairy medifacies
are referred to. In similar context, parafrontal spots become of question-
able value as primary criteria to separate species of Sepedon.

The following key is adapted from Steyskal 1950, with particular
emphasis on robust species. The wing length is measured from the
basal fracture line. The stated measurements represent the extremes
as based on the specimens at hand. The means fall well within the
extremes which constitute a very small percentage of total numbers
examined. In most species females are larger than males. S. pacifica
occasionally overlaps the size range of S. praemiosa , but since the two
do not overlap in distribution to our knowledge, size alone will not
be a critical determinative factor. Determination of material from
areas of known or potential overlap should be based primarily on
proper examination of male genitalia.

In the area covered by our key taxa currently included in the armipes
group are S. anchista Steyskal, S. armipes Loew, S. bifida Steyskal,
S. capellei Fisher and Orth, S. haplobasis Steyskal, S. melanderi Stey-
skal, and S. pseudarmipes Fisher and Orth. Sepedon haplobasis Steyskal
is reported only from Mexico, D. F. ; male hind femur notched, wing
length 4.6 to 5.5 mm, females 5.1 to 5.9 mm. The pusilla group consists
of S. borealis Steyskal, S. lignator Steyskal, S. neili Steyskal, and S.
pusilla Loew. Sepedon guatemalana Steyskal was reported by Neff
and Berg (1966, p. 41) from Las Cruces, Chiapas, Mexico, approxi-
mately 17° latitude, and thus is the most northern taxon known of the

JANUARY 1972] FISHER & ORTH SEPEDON SYSTEMATICS

13

lindneri group. A presumed range extension of S. macropus Walker
is provided by a single female collected by the senior author on the
west coast of Mexico, Sonora, 0.5 mi. s. of sign to Potam, 300 feet,
5 October 1967.

Publications which contain illustrations of adult Sepedon are those
by Steyskal (1950, 1956, 1960), Foote (1961), Yano (1968), and
Fisher and Orth (1969). Biologies and distribution maps for 14 taxa
of North and Central America are reported by Neff and Berg (1966).

Key to the Nearctic and Mexican Species
of Sepedon Latreille

1 Supraspiracular convexity of metathorax with black hairs 2

Supraspiracular convexity of metathorax without black hairs 10

2 Robust species; wing length 5.8 to 8.9 mm; hind femur of males without

midventral notch 3

Smaller species; wing length 3.6 to 5.5 mm; hind femur of males with or
without midventral notch. North of Tropic of Cancer 8

3 Medifacies with or without fine black hairs; femora orange-yellow to

reddish brown with or without bicoloration of black 4

Medifacies, females, without fine black hairs; femora yellowish, not bi-
colored; wing length 5.5 to 6.4 mm, (smaller females and all males
covered by 2') . U.S.A. north of 38° latitude and Canada. [Steyskal,
1950] S. spinipes americana Steyskal

4 Hind femur slender, much longer than abdomen, length — 69-78% wing

length, black apically with or without distinct black annulus. Subtropical

to tropical 5

Hind femur thickened, not or but little longer than abdomen, length
= 56-59% wing length. Subtropical to temperate 7

5 Middle femur with strong midanterior spinule 6

Middle femur without strong midanterior spinule. Trinidad. [Steyskal,

1950] S. trinidadensis Steyskal

6 Hind femur with blackish pre-apical annulus. Tropical lowlands of eastern

Mexico north to extreme southern Texas, Central America, Columbia.

[Walker, 1849, p. 1078] S. macropus Walker

Hind femur black on apical third. Puerto Rico, Dominica, Haiti. [Melander,
1920] S. caerulea Melander

7 Face yellowish to amber, with or without scattered fine black hairs on

medifacies — may be totally bare; wing length males 5.8-7.2 mm, females
6.3-7.3 mm; hind femur usually less than 4 mm; genitalia as Figs. 1-6.
Southwestern Nebraska, western Kansas, Colorado, Arizona, New Mexico,
Mexico highlands to Oaxaca; 1,100-8,000 feet. [Giglio-Tos, 1893, 1896]

S. praemiosa Giglio-Tos

Face amber to brownish, medifacies with fine black hairs scattered to
moderately dense; wing length males 7.2-8.9 mm, females 7. 0-8.0 mm;
hind femur usually greater than 4 mm; genitalia as Figs. 7-12. All
states west of Rocky Mts. (no records from Arizona) ; east of Rocky Mts.

14

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

north of 40° latitude and east to 96° 28' longitude; extreme southern
portions (50° latitude) of western Canadian Provinces; sea level to 10,150
feet. [Cresson, 1914] S. pacifica Cresson

8 Male hind femur emarginate ventrally; female hind femur simple; hind

tibia distinctly more curved in distal third; abdomen brown with little
more than a trace of bluish reflection; oral margin usually raised,
rectangular in profile ; tp almost straight. Canada, U.S.A., Mexico.

[Steyskal, 1950, 1956, 1960; Fisher and Orth, 1969] S. armipes group

Hind femur of both sexes simple 9

9 Hind tibia more or less evenly arcuate; abdomen frequently almost black

with bluish reflections; oral margin usually low, the angle with face
acute; wing length males 3. 6-4. 5 mm, females 3. 9-4.9 mm; femora orange-
yellow to dark brown; medifacies with fine black hairs. North of 37°

latitude. [Steyskal, 1950] S. pusilla group

Wing length males 5.0 to 5.5 mm, females 5.5 to 6.4 mm; femora yellowish;
medifacies without fine black hairs. U.S.A. north of 38° latitude, Canada.
[Steyskal, 1950] S. spinipes americana Steyskal

10 Robust species; wing length 6.3 to 7.6 mm. U.S.A., Canada 11

Smaller species. South America, Central America, Mexico (Chiapas, 2,200

feet) . [Steyskal, 1950] S. lindneri group

11 Second antennal segment compressed, about three times as long as wide

(viewed laterally), near base much wider than first segment. Widespread
in U.S.A. and Canada, rare in California (Plumas, Sierra counties) and
New Mexico (Rio Arriba, Sandoval counties), no Arizona or Nevada

records. [Loew, 1859; subspecies Steyskal, 1950] S. fuscipennis Loew

Second antennal segment almost cylindrical, nearly five times as long as
wide, basally not thicker than first segment; oral margin in profile acute.

U.S.A. and Canada east of 101° longitude. [Cresson, 1920]

S. tenuicornis Cresson

Biology, laboratory. — In late July 1970 we collected 50 S. praemi-
osa at Lake Mary, near Flagstaff, Arizona, and brought them to River-
side for propagation in the laboratory. Upon our return to Riverside
we collected S. pacifica locally. The object of these collections was to
initiate reciprocal cross mating studies. Many progeny of S.
praemiosa were produced, and on 24 August 1970, 60 F x puparia were
sent to Honolulu, Hawaii, c/o C. J. Davis, Chief Entomologist, for prop-
agation and release against Lymnaea ollula Gould. There were enough
F x adults of S. pacifica produced to enable us to set up reciprocal
crosses of groups of four females and four males as well as control
groups of both species. The results indicated interspecific mating
incompatibility, but were inconclusive. The Fi controls of S. praemiosa
mated and produced viable eggs but all the larvae died before or during
the third instar. The Fj controls of S. pacifica mated but no eggs were
laid. In the reciprocally cross mated groups no mating of Fi adults
was observed and no eggs were laid.

JANUARY 1972] FISHER & ORTH — SEPEDON SYSTEMATICS

15

Although the reasons for failure of these attempted crosses are
obscure, certain comments are considered appropriate. The Fj larvae
were reared on a diet of virtually 100% Physa virgata Gould, and a
high percentage of both species attained the third instar and then
died. Many successfuly pupated, but emerged deformed and soon died.
Curiously, our insect pathologist colleagues at Riverside could identify
no pathogens in moribund or recently dead material. Possibly, the
diet solely of P. virgata did not provide all the nutrients required for
normal development. Another factor we have recognized in these and
other species of sciomyzid flies — as also reported by Neff and Berg
(1966) — is that late season (September, October) attempts at laboratory
propagation often fail, even with fresh appearing adults. Perhaps a
form of ovarian diapause is responsible — a possible survival factor in
species suspected of overwintering as adults, and whose immatures
could not survive.

We concur with the generalized observations reported by Neff and
Berg (1966) for their laboratory rearings, i.e., 3 to 5 days egg incuba-
tion period, 12 to 20 days in the three larval stadia, and 5 to 9 days
in the pupal stage. In their paper, the biology of S. praemiosa , includ-
ing descriptions of the immature stages, was presented in considerable
detail. However, we call attention to the facts that their cultures
consisted of material from three localities — (1) Oxaca, Mexico, col-
lected August 1958; (2) Cypress Lake, Saskatchewan, Canada,

collected July 1957; and (3) Riverside, California, collected January-
February 1961. [Larvae from the Riverside material were taken to
Hawaii and Australia by Berg the following August for testing against
the freshwater snails Lymnaea ollula Gould and L. tomentosa Pfeiffer,
respectively. Ref. Neff and Berg (1966, p. 48) ]. Although no biological
or morphological differences between immatures from the three localities
were reported, we consider the first to be S. praemiosa , and the second
and third to be S. pacifica.

Biology, field. — The general statement by Neff and Berg (1966,
p. 48) that “the species ( praemiosa , = pacifica ) breeds throughout
the year in southern California . . .” may hold for coastal or low eleva-
tions, but it does not pertain to the higher elevations, such as Big
Bear Lake at 6,750 feet in the San Bernardino Mts. or to Lake Hemet,
at 4,500 feet in the San Jacinto Mts. We have sampled both localities
extensively over a period of years. Our contention is particularly true
at Big Bear Lake which sometimes is frozen over, and nearly every year
has substantial snow cover during December, January, and February.
Water temperatures during these months are in the low 40’s °F or

16

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

upper 30’s °F. At comparable and higher elevations in the northern
parts of the state where S. pacifica occurs winter conditions are more
severe. Because of these observations we are reluctant to comment on
the exact mode of overwintering in cold areas.

It seems reasonable that the onset of winter would find eggs, larval
stadia, and pupae present as well as adults. To begin elucidation of
the mechanisms involved, temperature tolerances of all stages require
critical study as do other ecological parameters such as day length,
light intensity, etc., and the interactions of such stimuli. Perhaps all
stages are merely arrested developmental^, perhaps developing a little
on warmer winter days. What the larvae would feed on is another
question, since at many sites migrating water fowl consume most of
the exposed mollusks during the fall and early winter. Surviving
aquatic mollusks are buried in the mud, or are on rocks below the mud
line, or in lakes they may be in vegetation well below the surface.
In such situations they are beyond the reach of bird or fly-maggot
predators.

Distribution and Relative Abundance of S. praemiosa, S.
pacifica, and S. fuscipennis. — These three large and superficially
similar species are usually encountered in open, unshaded marshes.
Most such habitats contain aquatic pulmonate snails of the families
Planorbidae, Lymnaeidae, and/or Physidae. In the laboratory the
larvae of the three species of marsh flies consume a wide variety of
aquatic snail species (Neff and Berg, 1966). Although precise host
associations in nature or nutritional requirements remain to be eluci-
dated, the assumption presently is that since the larvae of these species
feed as overt predators, their nutritional requirements are not tied
to any particular species of mollusk.

We know of no locality (microhabitat) where S. pacifica and S.
praemiosa occur together. A waterway common to both appears to be
the South Platte River. Sepedon pacifica occurs along its western reaches
in northeastern Colorado, and S. praemiosa occurs at North Platte in
southwestern Nebraska. The intermittent nature of the river during
the summer may serve to keep the species separated, but it is possible
that seasonally the two species may coexist along this watercourse as
well as the North Platte River possibly in the vicinity of the western
boundary of Nebraska.

Sepedon fuscipennis is widespread in the U.S.A. and Canada. It is
found in association with S. pacifica in several localities west of 100°
longitude. In the Pacific states the two species occur together in Cali-
fornia (south of Crescent Mills, Plumas Co.; west of Sierraville, Sierra

JANUARY 1972] FISHER & ORTH — SEPEDON SYSTEMATICS

17

Fig. 13. Distribution of S. pacijica and S. praemiosa. We have seen specimens
from all localities indicated by the blackened circles and stars. Clear circles
indicate material not seen but referred to in the literature as S. praemiosa or
S. pacifica and which because of distribution, we assign to pacifica.

Co.), Oregon (Eugene, Lane Co.; Klamath Game Refuge, Klamath Co.),
and in Washington (south of Kalama, Cowlitz Co.). Foote (1961)
reported S. praemiosa (= S. pacifica ) and S. fuscipennis from Idaho
(Coeur d’Alene and Robinson Lake) and Alberta, Canada (Lethbridge) .
The only sites where we are certain S. praemiosa and S. fuscipennis

18

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

occur together is a roadside marsh just east of North Platte, Nebraska,
and in the Jemez Mts., Sandoval Co., New Mexico. In spite of rather
intensive collecting efforts by us and others in the western states and
Canada it is curious that S. fuscipennis has been reported together with
S. pacifica or S. praemiosa at relatively few sites.

Localities where the three species have been reported singly hut
where overlap may be anticipated are as follows. We have taken only
S. fuscipennis at Amargosa and Pike’s Stockade in southern Colorado;
at Chehalis, Centralia, and Silver Lake in Washington; and at Cave
Junction, in Oregon. Foote (1961) reported only S. fuscipennis from
six localities in Idaho, one in Montana, four in Washington, two in
Oregon, two in Alberta, and three in British Columbia. We have taken
only S. praemiosa at Walsenburg, Colorado, and at Lake Mary (8 miles
southeast of Flagstaff), Arizona. Only S. pacifica (= S. praemiosa,
Foote, 1961) was reported by Foote (1961) from 14 sites in Idaho,
28 in Utah, five in Washington, two in Oregon, and one in Alberta.

Material we have seen, and assumptions from literature records
(Steyskal, 1950; Neff and Berg, 1966; Foote, 1961) indicate that in
the U.S.A. S. praemiosa occurs east of the Rocky Mts. at North Platte,
Nebraska, (approximately 41° latitude, 120° longitude), in eastern
Colorado, southwestern Kansas, and along the Rio Grande watershed
in New Mexico. West of the Rocky Mts., S. praemiosa occurs in Arizona
near Phoenix and Flagstaff and in the southeastern portion of the state.
The gorges of the Colorado River and Little Colorado River appear
to be the northern limit of its distribution in Arizona, but a male was
seen from Moab, Utah. South of the International Border, S. praemiosa
occurs in the central highlands of Mexico to Oaxaca, 17° latitude. The
present occurrence of isolated populations of S. praemiosa in well sep-
arated drainages suggests that there may have been wet periods in
the geological past that would have effected a continuous suitable habitat
from Oaxaca to Nebraska, or possibly farther north. Although we
have seen no material from Texas or Mexico from the Rio Grande or its
tributaries, this watershed probably enabled the species to penetrate
as far north as it has.

Sepedon pacifica occurs west of the Rocky Mts. from northern Baja
California del Norte, Mexico, 32° latitude, to 50° latitude, becoming
rare in northwestern Washington and western British Columbia, Canada.
East of the Rocky Mts. it occurs between 40° and 50° latitude. Its
easternmost confirmed occurrence is Sioux City, Iowa, 96° 28' longitude.
We saw the material on which the Neff and Berg (1966) record from

JANUARY 1972] FISHER & ORTH — SEPEDON SYSTEMATICS

19

North Dakota and records in South Dakota and northeast Colorado
were based and assign it all to S. pacifica.

The arid southwestern U.S. — Sonoran Desert creates the zoogeo-
graphic barrier which effectively separates S. pacifica from S. praemiosa.
A curious fact is that we have never collected either species, nor do we
know of any confirmed records, along the Colorado River which bisects
this desert, nor in the high or low deserts of southeastern California
and the contiguous areas of Nevada and Arizona.

The paucity of S. praemiosa and S. pacifica in the Colorado River
drainage system may be caused by the intermittent turbulence and
velocity with the heavy load of silt carried by that river which is a
deterrent to snail development. The dams which have been constructed
over the past forty years are holding much of the silt that otherwise
would flow the length of the river. The result is that relatively calm
and clear water habitats are becoming more common. Perhaps in time
this alteration of the habitat will be taken advantage of by species of
pulmonate gastropods which are suitable, if not necessary, host of
S. pacifica and S. praemiosa, namely planorbid snails, and these sciomy-
zid flies will thus be able to extend their range.

Literature Cited

Aldrich, J. M. 1905. A catalogue of North American Diptera. Smithson. Misc.

Collect., 46 (2— publ. 1444) : 1-680. Sciomyzidae, pp. 577-581.
Cresson, E. T., Jr. 1914. Descriptions of new North American Acalyptrate
Diptera — I. Entomol. News, 25(10) : 457.

1920. A revision of the Nearctic Sciomyzidae (Diptera: Acalyptratae) . Trans.
Amer. Entomol. Soc., 46: 27-89.

Fisher, T. W., and R. E. Orth. 1969. Two new species of Sepedon separated
from S. armipes Loew in western North America. Pan-Pac. Entomol.,
45(2) : 152-164.

Foote, B. A. 1961. The marsh flies of Idaho and adjoining areas (Diptera:

Sciomyzidae). Amer. Midland Natur., 65(1): 144^167.

Giglio-Tos, E. 1893. Diagnosi di nuovi generi e di nuove specie di Ditteri.
Boll. Mus. Zool. Anat. Comp. Univ. Torino, 8(158) : 1-14.

1896. Ditteri del Messico. Parte IV. Muscidae, Calypteratae. Muscidae
Acalypteratae. Mem. Reale Acad. Sci. Torino, Serie seconda., 45: 1-74.
Loew, H. 1859. Die nordamerikanischen Arten der Gattungen Tetanocera und
Sepedon. Wien. Entomol. Monatsschrift, 3(10) : 289-300.

Melander, A. L. 1920. Review of the Nearctic Tetanoceridae. Ann. Entomol.
Soc. Amer., 13(3) : 305-332.

Neff, S. E., and C. 0. Berg. 1966. Biology and immature stages of malaco-
phagous Diptera of the Genus Sepedon (Sciomyzidae). Va. Agr. Exp.
Sta. Bull., 566, 113 pp.

Steyskal, G. C. 1950. The genus Sepedon Latreille in the Americas (Diptera:
Sciomyzidae) . Wasmann J. Biol., 8(3) : 271-297.

20

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

1956. New species and taxonomic notes in the family Sciomyzidae (Diptera:

Acalyptratae) . Pap. Mich. Acad. Sci. Arts Lett., (1955), 41: 73-87.
1960. New North and Central American species of Sciomyzidae (Diptera:
Acalyptratae). Proc. Entomol. Soc. Wash., 62(1): 33^43.

Walker, F. 1849. List of the specimens of dipterous insects in the collection
of the British Museum. Part IV: pp. 689-1172.

Wulp, F. M. van der. 1897. Sciomyzinae, pp. 354-360. In Biologia Centrali-
Americana. Diptera. II. Godman, F. D. and 0. Salvin, [editors].
489 pp. 1888-1903. London.

Yano, K. 1968. Notes on Sciomyzidae collected in paddy field (Diptera) , I.
Mushi, 41(15) : 189-200.

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JANUARY 1972] RASKE BIOLOGY OF ANELAPHUS

21

Immature Forms, Genitalia, and Notes
on the Biology of Anelaphus albofasciatus Linnell 1

(Coleoptera: Cerambycidae)

A. G. Raske

Canadian Forestry Service, Department of Fisheries and Forestry,

Edmonton, Alberta 2

Long-horned beetles of the genus Anelaphus generally feed on various
hardwood trees (Linsley, 1963). Anelaphus albofasciatus (Linnell)
is unique within this genus for its habit of feeding on cactus. Adults
have previously been collected on Opuntia bigelovii Engelmann (Davis,
1928) and on Echinocactus sp. (Linsley, 1963). This paper reports
my observations of A. albofasciatus on Opuntia whipplei Engelmann
and Bigalow (3 to 6 miles west of Seligman, Yavapai County, Arizona,
during mid July and early September 1964 and mid June 1966) and
describes characters that separate Anelaphus from Moneilema, another
cerambycid genus which may occur on the same host.

Notes on the Biology

The adult beetles are active in June and July, the larvae are present
in July and the pupae appear from late July and throughout the fall.
These observations suggest that there is one generation per year on
Opuntia whipplei.

The adults become active after dusk, and are usually found near
the top of the cactus where they feed on the newest growth. They are
most often found crawling slowly over the host plants but are also
often motionless on the top of a branch with their legs extended
and their bodies held high off the substratum; the antennae are held
high in the air at a 45° angle. The beetles may remain montionless
in this position, which resembles preflight posture, for up to five hours.

Several mating pairs were observed, all at night and on the top of
the cacti.

The size and shape of larval mines at early larval instars indicate
that eggs are deposited near or on the surface of the host plant. The
young larvae apparently mine directly into the pith of the host through
an opening of the lace in the cactus skeleton, and proceed either up or

1 The work was started at the University of California, Berkeley, and completed at the Canadian
Forestry Service, Department of Fisheries and Forestry, Calgary, Alberta.

2 Present address: Canadian Forestry Service, Department of Fisheries and Forestry, P. O. Box
6028, St. John’s, Newfoundland, Canada.

The Pan-Pacific Entomologist 48: 21-26. January 1972

22

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Fig. 1. Anelaphus alhofasciatus Linnell, a) adult, b) pupa, c) larva (X4)
(Photos by P. Debnam, Canadian Forestry Service).

down from this point of entry. In contrast, the larvae of the cerambycid
Moneilema sp. first mine the exudate produced by the plant and then
proceed into the pith (Raske, 1966). The larvae of A. albofasciatus
constructs a pupal cell from pith and frass, usually at the bottom of
the larval mine (Fig. 2).

The cerambycid Moneilema semipunctatum forte LeC. (Raske, 1966)
attacks O. whipplei in the same area. The larvae of the two species
can be distinguished by the presence ( Anelaphus ) or absence ( Monei-
lema ) of prolegs. Generally, A. albofasciatus larvae are found in the
upper, smaller stems and branches of the plant, while M. semipunctatum
forte larvae tend to be in the main stem near the ground or in the roots.

Larvae of A. albofasciatus were reared successfully in the laboratory
on the artificial diet used by Lyon and Flake (1966). From one of
the larvae, 41 adults of the internal parasite, Tetrastichus holbeini
Girault (Hymenoptera: Eulophidae), emerged in early August.

JANUARY 1972]

RASKE — BIOLOGY OF ANELAPHUS

23

Fig. 2. Larval mine of A. albofascicitus in Opantia whipplei with pupal cell (a)
and adult exit hole (b) . (X 1.3) (Photo by A. A. Blaker, Univ. of California).

24

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

d.

ventral

Fig. 3. Antenna of larva of A. albofasciatus. Fig. 4. Male genitalia of A.
albofasciatus, a) aedeagus, b) intromittent structures, c) endophallus (dorsal
and lateral), d) tegmen (ventral and lateral), aed — aedeagus, ejd — ejaculatory
duct, endph — endophallus, ints — internal sac, tg — tegmen.

Description of Life Stages

Adult. — Linsley (1963) described the external characters of adult
A. albofasciatus (Fig. la), but the internal reproductive structures of
taxonomic value were not included. These are illustrated in Figs. 4
and 5.

Mature larva. — Length 18-22 mm. (Fig. lc). Head transverse, moderately
depressed. Genae testaceous behind ocelli for about one-fourth distance between
ocelli and base of mandible; sclerotization of mouth frame narrow, setae scattered,
most not in testaceous area; antennae three-segmented (Fig. 3) ; mandible short,
stout, cutting edge “gouge-like”; labrum setose, evenly rounded anteriorly, sud-
denly constricted posteriorly ; clypeus transverse, four times as wide as long,
anterior margin sclerotized; ligula obtusely conical, tip setose; labial palp two-

JANUARY 1972] RASKE — BIOLOGY OF ANELAPHUS

25

Fig. 5. Anelaphus albofasciatus, female genitalia, a) ovipositor (dorsal),
b) ovipositor (ventral), c) stylus, d) internal genitalia, accgl — accessory
gland, ovd — oviduct, sp = spermatheca, spd = spermathecal duct, spgl —
spermathecal gland.

segmented, lightly sclerotized; maxillary palp three-segmented, lightly sclerotized,
process of palpifer prominent, maxillary lobe, cardo, and stipes sclerotized
basally, with scattered setae; mentum and submentum membranous; gula with
sides widely diverging posteriorly; ocelli two, pigmented, subcontiguous. Pronotum
transverse, moderately setose, distinctly raised from pleural regions, dorsal plate
divided, with two distinct light testaceous areas, longitudinally striate; eusterna

26

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

rugose. Abdomen rugose, with ampullae shining, bilobed with transverse and
lateral deep grooves, subtuberculate; epipleura inconspicuous; pleural disc a
deep pore with radial striations; spiracles broadly oval to suborbicular ; peritreme
not raised above general level of cuticle.

Pupa (female ?) — Length 13-15 mm. (Fig. lb). Head with labrum subequal
to clypeus in length, clypeus with deep longitudinal groove; antennae reaching
to third abdominal segment. Pro-, meso- and metanotum with few inconspicuous
setae. Abdominal terga with heavily sclerotized incurved spines, fewest and
smallest on first abdominal segment, increasing in size and number on posterior
segments; most spines with subapical setae; posterior margin of eighth tergite
with row of large spines; sterna glabrous, tenth sternum deeply, irregularly
furrowed, with pronounced median blister-like structure, bearing two small setae
posteriorly.

Acknowledgments

Financial assistance from the National Science Foundation Grant
# GB 2326 is gratefully acknowledged. For the identification of the
adults and for helpful advice, I thank Dr. E. G. Linsley and Dr. J. A.
Chemsak, both of the University of California, Berkeley. Thanks to
Dr. L. Gardiner, Canadian Forest Service, Sault Ste. Marie, for check-
ing the description of immature forms. The parasites were identified
by Dr. B. D. Burks of the U. S. National Museum.

Literature Cited

Davis, A. C. 1928. Southern California collecting notes. Pan-Pac. Entomol.,
4(4) : 183.

Linsley, E. G. 1963. The Cerambycidae of North America, Pt. IV. Univ. Calif.
Publ. Entomol., 21 : 1-165.

Lyon, R. L., and H. W. Flake. 1966. Rearing Douglas-fir tussock moth larvae
on synthetic media. J. Econ. Entomol., 59(3) : 696-998.

Raske, A. G. 1966. Bionomics and Taxonomy of the genus Moneilema (Cole-
optera: Cerambycidae). Univ. Calif., Berkeley, Ph.D., 268 pp.

JANUARY 1972] SCHROETER & HEWITT — GRASSHOPPER CYTOLOGY 27

Cytology of Some California Grasshoppers.

1. Taxonomic Considerations

(OrthopterarAcridoidea)

Gilbert L. Schroeter and Godfrey M. Hewitt

Department of Biology, Texas A&M University, College Station 77843, and
School of Biological Sciences, University of East Anglia, Norfolk, England

The purpose of this paper is to call attention to the extent of chromo-
somal variation, both structural and numerical, in several species of
grasshoppers and its bearing on orthopteran taxonomy.

Many grasshoppers have evolved novel genetic systems which are
not only of intrinsic interest to evolutionary biologists but in many
cases can be taxonomically useful (White, 1951, 1954). During the
course of studying the effects of chromosome rearrangements on genetic
recombination in various species (Hewitt, 1967; Hewitt and Schroeter,
1968) , numerous other species were collected in California which had
not been previously studied cytologically. The taxonomic status of
some of these has remained in doubt due to the fact that morphological
features considered alone have not proved reliable enough in establish-
ing phylogenetic relationships. Six species are reported on here which
make clear that a chromosomal analysis could, indeed should, be used
in any serious taxonomic treatment.

Materials and Methods. — The species discussed, sites and dates
of collection, and number of individuals examined cytologically are
given in Table 1.

Testes from adult male grasshoppers were removed following a
mid-dorsal incision and immediately placed in a fixative of 3:1
absolute ethyl alcohol: glacial acetic acid. Ovarioles of females were
removed by vivisection under insect saline and then cultured for 30
minutes in 0.05% colchicine in insect saline to arrest and accumulate
mitotic metaphases before being similarly fixed. A detailed description
of the reproductive anatomy of grasshoppers is to be found on pages
138-150 in Uvarov (1966).

In our experience young male imagines are a good source of meiotic
material but meiotic divisions can usually be found even in the testes
of males collected late in the season. Only young female imagines
have proved to be satisfactory for the study of ovariole wall mitosis.

Following fixation the material was stored in the fixative or 70%
ethanol in a refrigerator and subsequently squashed in acetic-orcein.

The Pan-Pacific Entomologist 48: 27-36. January 1972

[VOL. 48, NO. 1

28

THE PAN-PACIFIC ENTOMOLOGIST

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O O

JANUARY 1972] SCHROETER & HEWITT GRASSHOPPER CYTOLOGY 29

For details concerning stain preparation and the squash technique the
reader is referred to pages 199-205 in Lewis and John (1964).

Cytological Observations and Discussion

Tanaocerus koebelei Bruner. — The family Tanaoceridae contains but
three species, two of which are found in California. The relationship
of these desert longhorns to the rest of the Acridoidea remains obscure.
In examining the proventricular region of the tanaocerids and compar-
ing it with other grasshoppers, Grant and Rentz (1967) concluded
that the tanaocerids are quite different from both the acridids and
eumastacids. On the other hand, Dirsh (1956, 1961) considers the
Tanaoceridae to be more allied to the eumastacids and includes them
in the Eumastacoid section of his taxonomic scheme which is based
upon comparative studies of the phallic complex. It should be pointed
out that only the epiphallus of Tanaocerus koebelei was studied, a
structure considered by Dirsh (1956) to be a taxonomic character
reliable enough for differentiating families or even groups of families
in some cases.

Figure 1 shows an ovariole colchicine-arrested (c-mitotic) metaphase
with 22 acrocentric chromosomes. In the past the term “acrocentric”
has been used to describe chromosomes with visible short arms and
also those in which short arms were not able to be resolved with the
light microscope but presumed to be present. This is discussed by John
and Hewitt (1968) and White (1969). To avoid confusion, the term
acrocentric will be used in this paper in its broad sense, realizing that
rod-shaped chromosomes in many instances may possess truly terminal
centromeres (i.e., are telocentric) . Until a male individual of Tanaocerus
is studied, a decision concerning the sex-determining mechanism cannot
be made. However, if it is the same as in most other grasshoppers,
it is an XO S /XX 9 system, with the X in this case also being acro-
centric.

Hundreds of acridid species have been studied and most have a
karyotype consisting of 2n$ = 23 (2n2 = 24) acrocentric chromo-
somes (White, 1969). Although fusions between chromosomes have
occurred in many species, thus reducing the chromosome number, this
does not explain the peculiar karyotype of T anaocerus koebelei in
which all the chromosomes are acrocentric. A reduction in number from
22 to 20 by “centric fusion” would be expected to result in a pair of
chromosomes with median or sub-median centromeres.

On the other hand, the chromosome complement of Tanaocerus
koebelei graded in size is very similar to what could be argued as the

30

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

JANUARY 1972] SCHROETER & HEWITT— GRASSHOPPER CYTOLOGY 31

basic karyotype in the Eumastacidae. It closely resembles that of the
Pseudoschmidtiinae and Mastacideinae subfamilies, the cytogenetics
of which have been described by White (1970a). As his study makes
clear, however, the eumastacids are chromosomally quite varied and
a morphologically diverse group.

Certainly more information about the cytogenetics of these groups
needs to be obtained before any conclusion concerning their phylogenetic
interrelationships can be made.

Dracotettix monstrosus Bruner. — White (1954) has drawn attention
to the discussion as to whether the Old World subfamily Pamphaginae
is represented in the New World by such genera as Plirynotettix and
Dracotettix , or whether these belong in the subfamily Romaleinae. He
points out that the Old World Pamphaginae have 21 (20 + X) chromo-
somes in males while the Romaleinae possess 23 (22 + X). Since
Phrynotettix males also have 22 + X chromosomes (McClung, 1914)
its inclusion in the latter subfamily seems warranted. Consequently
we have determined the basic karyotype of Dracotettix monstrosus and
found a male to have 22 + X acrocentric chromosomes (Fig. 2). This
clearly indicates that its inclusion in the Romaleinae is warranted
and both cases together argue against the presence of the Pamphaginae
in the New World.

Oedaleonotus borckii (Stal).- — A total of 20 individuals from two
populations on either side of the Central Valley of California has
been examined and both samples appear to be cytologically uniform
with males having 22 + X acrocentric chromosomes (Fig. 3).

Oedaleonotus enigma (Scudder). — Over 1,000 males of this species
have been collected in the lower San Joaquin Valley near Coalinga.
This particular species exhibits considerable chromosome variation
and differs strikingly from other members of the genus so far studied
in its sex-determining mechanism (Hewitt and Schroeter, 1968). In
contrast to the XO & /XX $ sex-chromosome systems common to most

4 -

Fig. 1. Mitotic complement of T anaocerus koebelei. 2n $ =: 22. Note the
terminal or near-terminal position of the centromeres. X 1125. Fig. 2. First
meiotic metaphase of Dracotettix monstrosus. 2n $ — 23. Eleven bivalents and
a single unpaired sex chromosome (arrow). X 925. Fig. 3. First meiotic metaphase
of Oedaleonotus borckii. 2n $ = 23. All chromosomes possess terminal or near
terminal centromeres, including the X chromosome. X 1250. Fig. 4. First meiotic
metaphase of Oedaleonotus enigma with the ‘basic’ karyotype of 2n $ — 22. Note
the neo-XY sex bivalent (X'Y')- All chromosomes are aero- (telo-?) centric except
the neo-X. X 850. (Line scale rr 10/x.)

32

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

grasshoppers, 0. enigma possesses a neo-XY (X'Y') S /Neo-XX (X'X') 2
mechanism which has evolved from the former by the fusion of the
heterochromatic X with an autosome to form the Neo-X, the homologous
autosome becoming the Neo-Y. Instead of possessing an unpaired
X-chromosome, males of this species are characterized by having a
heteromorphic sex bivalent at the first meiotic metaphase composed
of a metacentric Neo-X and an acrocentric Neo-Y (Fig. 4). The few
females which have been examined possessed two Neo-X’s. Thus,
instead of males having 23 (22 autosomes + X), both males and females
of this species have 22, 20 autosomes plus the two sex chromosomes.

While 2n = 22 is considered to be the basic chromosome number for
this species, male individuals have been found in all populations so
far sampled with 20 and 21 chromosomes. This variation in chromo-
some number is due to the fact that 0. enigma is polymorphic for a
centric fusion involving two non-homologous acrocentric chromosomes,
numbers 4 and 5. A fusion between these two chromosomes has oc-
curred at or near their centromeric regions resulting in one large
metacentric chromosome. An individual may (i) possess the basic
karyotype of 20 acrocentric autosomes, (ii) be heterozygous for the
fused and unfused chromosomes or, (iii) be homozygous for the two
fused metacentric chromosomes. Those individuals which are heter-
ozygous for the fusion possess nine chromosome pairs and a V-shaped
chain-of-three at Ml of meiosis (Fig. 5), while those homozygous for
the fusion have 10 pairs, one of which is a large ring bivalent made up
of the two fused metacentrics (Fig. 6) .

The frequencies of the two types of chromosomes involved in this
fusion polymorphism vary from population to population, but no
population so far sampled has become fixed for either of the two types,
i.e., no monomorphic population has been found. There are, however,
differences in frequencies between populations for the fused and
unfused chromosomes (Schroeter and Hewitt, in preparation).

Although the Neo-XY sex-determining mechanism and the fusion
polymorphism are the most conspicuous cytological features of O.
enigma, there are several other chromosomal alterations for which the
species is also polymorphic.

The first of these is a centromere ‘shift’ in the third smallest chromo-
some (no. 8) . Whereas all the chromosomes of the basic karyotype
are acrocentric, a transposition of the centromere has occurred in the
number 8 chromosome presumably by means of a pericentric inversion,
resulting in a chromosome with the centromere in a sub-median position.
Again, in all samples studied, three types of individuals have been

JANUARY 1972] SCHROETER & HEWITT — GRASSHOPPER CYTOLOGY 33

observed: (i) those homozygous for the two acrocentric no. 8 chromo-
somes; (ii) heterozygous for the acrocentric and submetacentric
forming a heteromorphic bivalent (Fig. 5) and, (iii) homozygous for
the two submetacentrics.

In some individuals one member of the smallest chromosome pair,
no. 10, has an additional heterochromatic segment attached to it,
resulting in a conspicuous unequal bivalent at Ml of meiosis (Fig. 6).
While this enlarged chromosome has been found in all but one of the
populations sampled to date, its frequency is usually so low that only
a very few individuals have been found to be homozygous for it.

The last chromosomal variable to be mentioned is that some in-
dividuals in some populations possess heterochromatic supernumerary
or B-chromosomes which do not pair with any member of the standard
chromosome complement. The occurrence of these B-chromosomes is
even lower than that of the segment attached to chromosome 10, its
frequency not exceeding 2.5% in any of the populations so far sampled.
The meiotic behavior of these and the other variable components of
the chromosome complement of O. enigma and their possible role in
the population dynamics of the species have been discussed in a
previous paper (Hewitt and Schroeter, 1968). However, it is worth
emphasizing that these B’s are both metacentric and rare, and conse-
quently, unlikely candidates for the extra sex chromosome arm de-
scribed in three individuals of this species (Hewitt and Schroeter,
1968) as has been suggested by White (1970b) .

Oedaleonotus orientis Hebard. — Only two individuals of this species
have been collected, but both exhibited a striking chromosomal feature
which may prove to be of considerable taxonomic importance. This
is the ditactic bivalent (Fig. 7). This peculiar type of bivalent has
been thought to be due to chiasma formation between the short arms
of two acrocentric chromosomes, but John and Hewitt (1966, 1968)
have interpreted ditactic bivalents to result from chiasma formation
within truly terminal centromeres. Whatever interpretation may ulti-
mately be applied, in this instance the ditactic bivalent is a feature
that was consistently observed in all Ml cells of both individuals and
has not been observed in any of the allied species so far analyzed. As
in 0. borckii, males of this species possess 23 acrocentric chromosomes.

Oedaleonotus phryneicus Hebard. — Only a single population of this
species has been sampled to date with a total of 96 males having been
examined cytologically. The chromosome complement, typical of most
acridids, consists of 22 autosomes and one unpaired sex-chromosome
in males, all being rod-shaped with terminal centromeres. There is

34

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

JANUARY 1972] SCHROETER & HEWITT — GRASSHOPPER CYTOLOGY 35

one feature which this species shares with O. enigma, i.e., a super-
numerary chromosome segment attached to a member of the smallest
autosomal pair. About 12% of the individuals observed were found
to be heterozygous for this segment, resulting in an unequal bivalent
as seen in Fig. 8. Each of two individuals also possessed an unpaired
heterochromatic supernumerary chromosome.

Four of the seven presently recognized species of Oedaleonotus
having been studied cytologically, the fact emerges that these four
differ in some unique way from each other. 0 . enigma contrasts
strikingly with the others by possessing a Neo-XY sex-determining
mechanism and being polymorphic for an autosomal centric fusion.
O. orientis is the only species so far studied in which a ditactic bivalent
is formed at Ml of meiosis. O. phryneicus differs from 0. enigma in
its sex-determining mechanism (X0$/XX9) but is similar to the
latter in that it also is polymorphic for a supernumerary chromosome
segment and supernumerary chromosomes. 0 . borckii differs from
the other three in its being monomorphic, there being no apparent
chromosome polymorphism nor any type of peculiar bivalent formation
or sex-determining mechanism. Thus the Oedaleonotus complex would
seem to be a most suitable and extremely rewarding area for cytogenetic
and evolutionary investigations.

Based on the studies made, it is clear that cytological techniques
can be used to great advantage in a serious taxonomic treatment of
this genus. Indeed, we would consider any such study to be lacking
if the chromosome complement was not taken into account. Strohecker,
et al. (1968) state that “satisfactory treatment of the forms of this
genus will require comprehensive study not feasible at present. Speci-
mens from widely separated localities appear to represent quite distinct
species but many intermediate forms occur and we have not been able
to find sets of characters consistent enough to be regarded as of specific
worth.” It may well be that the chromosome complement will prove
to be a worthy character.

«-

Fig. 5. Metaphase I of Oedaleonotus enigma which is heterozygous for the
4-5 fusion forming a chain-of-three (large arrow) and a centric shift in chromo-
some no. 8 (small arrow). X 1050. Fig. 6. Metaphase I of Oedaleonotus enigma
which is homozygous for the 4^5 fusion (large arrow) and heterozygous for the
supernumerary segment on chromosome no. 10 (small arrow). X 1000. Fig. 7.
Metaphase I of Oedaleonotus orientis. 2n $ = 23. Note the ditactic bivalent
(arrow). X 1700. Fig. 8. Metaphase I of Oedaleonotus phryneicus. 2n $ = 23.
Note the unequal bivalent (arrow). X 1350. (Line scale = 10,u.)

36

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Literature Cited

Dirsh, V. M. 1956. The phallic complex in Acridoidea (Orthoptera) in relation
to taxonomy. Trans. Roy. Entomol. Soc. London, 108: 223-356.

1961. A preliminary revision of the families and subfamilies of Acridoidea.
Bull. Brit. Mus. Natur. Hist. Entomol., 10: 349^419.

Grant, H. J., and D. C. Rentz. 1967. A biosystematic review of the family
Tanaoceridae including a comparative study of the proventriculus. Pan-
Pac. Entomol., 43: 65-74.

Hewitt, G. M. 1967. An interchange which raises chiasma frequency. Chromo-
soma, 21: 285-295.

Hewitt, G. M., and G. L. Schroeter. 1968. Population cytology of Oedaleonotus.

I. The karyotypic facies of Oedaleonotus enigma (Scudder). Chromo-
soma, 25: 121-149.

John, B., and G. M. Hewitt. 1966. Karyotype stability and DNA variability
in the Acrididae. Chromosoma, 20: 155-172.

1968. Patterns and pathways of chromosome evolution within the Orthoptera.
Chromosoma, 25: 40-74.

Lewis, K. R., and B. John. 1964. The matter of Mendelian heredity. Little,
Brown, and Co., Boston. 269 pp.

McClung, C. E. 1914. A comparative study of the chromosomes in orthopteran
spermatogenesis. J. Morphol., 25: 651-749.

Strohecker, H. F., W. W. Middlekauff, and D. C. Rentz. 1968. The grass-
hoppers of California (Orthoptera: Acridoidea). Bull. Calif. Insect
Surv., 10: 1-177.

Uvarov, B. 1966. Grasshoppers and locusts. Vol. 1. Cambridge Univ. Press,
London. 481 pp.

White, M. J. D. 1951. Cytogenetics of orthopteroid insects. Advan. Genet., 4:
267-330.

1954. Animal cytology and evolution. Cambridge Univ. Press, London. 454 pp.

1969. Chromosomal rearrangements and speciation in animals. Ann. Rev.
Genet., 3: 75-98.

1970a. Karyotypes and meiotic mechanisms of some eumastacid grasshoppers
from East Africa, Madagascar, India and South America. Chromosoma,
30: 62-97.

1970b. Asymmetry of heteropycnosis in tetraploid cells of a grasshopper.
Chromosoma, 30: 51-61.

JANUARY 1972] HALL — NEW HETEROTROPINE BOMBYLIIDAE

37

New North American Heterotropinae

(Diptera: Bombyliidae)

Jack C. Hall

Division of Biological Control, University of California, Riverside, 92502

The subfamily Heterotropinae is for the most part made up of an
anomalous group of microbombyliids, rarely exceeding 5 mm in length.
The anomaly is attested to by the fact that Prorates Melander was
described as an empidid, Caenotus Cole as a therevid and Apystomyia
Melander runs to the Scenopinidae in most keys to families.

Melander (1950) gave an account of the Heterotropinae together
with a rearrangement of Curran’s (1935) key to genera. Since 1950
none of the species belonging to the various genera comprising the
Heterotropinae have been reported upon. The following account not
only describes a new genus and several new species but greatly extends
the range of distribution of the genus Prorates , hitherto known only
from New Mexico and southern California.

The Heterotropinae may be separated from other Bombyliidae by
the following combination of characters: Occiput flattened, not or
only slightly foveate behind ocellar tubercle; eyes bisected into two
sizes of facets, small below, large above, but without dividing line;
inner margin of eyes usually indented opposite antennal bases; eyes
of male usually holoptic, female dichoptic, wing with two submarginal
cells; four posterior cells with first always open; anal cell closed before
the margin and petiolate or closed at wing margin; anal lobe greatly
developed; legs without conspicuous bristles; body not densely hairy.

Key to Heterotropine Genera

1. Second posterior cell petiolate at base 2

Second posterior cell sessile, medial crossvein between fourth and fifth
veins arises from discal cell 3

2. Third vein forks closer to r-m crossvein than to tip of wing Prorates Melander

Fork of third vein closer to tip of wing than to r-m crossvein

Apystomyia Melander

3. Arista terminal, normal, of one or two segments 4

Third antennal segment terminates in microscopic tuft of hair

Caenotoidcs Hall. n. gen.

4. Proboscis projects well beyond oral margin Heterotropus Loew

Proboscis at most only slightly projecting beyond oral margin Caenotus Cole

The Pan-Pacific Entomologist 48: 37-50. January 1972

38

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Apystomyia Melander
(Fig. 1)

Apystomyia Melander, 1950: 146.

Melander based this genus on a number of specimens collected along
streams in the mountains of southern California. Apystomyia was
placed by Melander (1950) in the Heterotropinae “because it does
not conform with any other subfamily . . . .” An examination of part
of the type series confirms its placement but if and when additional
species are discovered the genus should be reevaluated.

Known only from the genotype species- — A. elinguis Melander.

The genus may be recognized not only by the characters given in
the key to genera but also by the delicate neuration in the wing, the
small second basal cell and consequently large discal cell. The head
and thorax are polished black and the abdomen subopaque; the hair
on the head and thorax is black and quite coarse, that on the abdomen
is white. The knob is entirely black and the proboscis is short and
non-projecting.

Heterotropus Loew
(Fig. 2)

Heterotropus Loew, 1873: 182.

This genus contains a large number of Palearctic species. It is
represented in the Nearctic region by a single species, H. senex Melander.
Melander’s specimen was collected in Arizona at flowers of Baileya
multiradiata.

Heterotropus senex has the second posterior cell sessile and the anal
cell closed at the wing margin ; proboscis long ; body black with yellow
markings and white hair. The halters are whitish.

Type locality, Headquarters Organ Pipe National Monument, Arizona.
The fly was collected on 16 April.

Prorates Melander
(Fig. 3)

Prorates Melander, 1906: 372 (as an empidid) , 1927: 377; 1950: 145.
Alloxytropus Bezzi, 1926: 186.

Originally described as an empidid but later transferred to the
Bombyliidae because of the short costal vein. For many years the
genus was known from a single species, P. claripennis Melander.
Efflatoun (1945) under the name Alloxytropus Bezzi, recorded two
species from Egypt. Melander (1950) synonymized this latter genus

JANUARY 1972 J

hall — new

HETEROTROPWE EOMBYLItDAE

1

2

Prorates frommeri

elin

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'guis. Pig o tv

Heterotropus

40

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

with Prorates. Five additional species are described below bringing
the total known species to eight.

Prorates may be diagnosed as follows: Proboscis projecting usually as far as
or farther than the tip of the antennae, except in P. boydi Hall described below.
Antennae with three segments, (not two as originally stated) first two very small,
third elongate conical and covered with minute pubescence. Scutellum with two
apical marginal bristles. Costal vein extends to or only slightly beyond the third
longitudinal vein; wing usually with darkened stigma below apex of first vein;
branch of fourth vein closer to r-m crossvein than to tip of wing; second posterior
cell petiolate at base; discal cell smaller than second basal cell. Vestiture of
short, fine, scattered hair, longer on underside of head and at base of abdomen.

Genotype species — P. claripennis Melander.

Key to Species of Prorates

1. Proboscis projecting well beyond oral margin 2

Proboscis short, at most labellae extend beyond oral margin; body and

legs black with white hair; knob of halter pale boydi Hall, n. sp.

2. Hair on mesonotum white or yellow, may reflect brownish in some

lights 3

Hair or mesonotum and scutellum black; scutellar bristles of male black,
of female testaceous; abdominal hair white nigrescens Hall, n. sp.

3. Proboscis projecting beyond oral margin for a distance equal to or less

than height of eye 4

Proboscis projects twice eye height beyond oral margin; palpi long,
extending as far as first antennal segment; legs fuscous; hair on
abdomen yellow arctos Hall, n. sp.

4. Legs fuscous, testaceous or dark brown, never mostly pale 5

Legs and thoracic pleural area pale, sternopleura and femora may be

dusky; basal antennal segments pale yellow; abdominal hair yellow
frommeri Hall, n. sp.

5. Abdominal hair golden yellow; proboscis usually extending beyond tip of

antennae; legs or at least the tibiae usually yellowish brown; wing veins

light brown melanderi Hall, n. sp.

Hair on abdomen pale yellow to white; proboscis extends as far as tip of
antennae or only slightly beyond; legs brownish yellow, tibiae of female
lighter; wing veins dark brown claripennis Melander

Prorates boydi Hall, new species

Immediately separable from the congeners by the short, non-protruding
proboscis and the entirely black body with pale hair.

Male. — Black, knees and tibiae brownish. Eyes contiguous down front for
distance greater than length of ocellar tubercle. Ocellar tubercle and small
triangular front bare. Antennae black, third segment may appear lighter in some
lights; basal segments short, without apparent vestiture; third segment, in lateral
view conical, broadest at base then tapering to narrow apex, at least twice as long
as wide, microscopically pubescent; arista terminal, as broad as apex of third

JANUARY 1972] HALL — NEW HETEROTROPINE BOMBYLIIDAE

41

segment. Face retreating, bare above, scattered short white hair below. Proboscis
short, not projecting beyond oral margin. Palpi nearly as long as proboscis, one
segmented. Occiput white pilose, hair not dense, nearly bare above.

Mesonotum dull, when viewed from in front brownish pollinose becoming
whitish laterally; hair short, scattered, yellowish, longer and tending towards
white near scutellum; bristles wanting (may be broken off). Pleura white
pollinose, bare except for few thin whitish hairs on mesopleuron. Legs with
short pale yellow hair, that on tibiae setaceous; bristles other than apical tibial
bristles wanting; hind tarsus longer than hind tibia. Pulvilli as long as claws.
Halter stem brownish, knob white. Squama white with fringe of white hair.
Scutellum dull with long, erect, white hair, bristles wanting.

Wing hyaline, stigmal area faintly yellowish; anterior veins brown, rest pale.
Costa terminates at fourth vein; second posterior cell short petiolate, and petiole
shorter than r-m crossvein; r-m crossvein at middle of discal cell; discal and
second basal cells of nearly equal length; petiole on anal cell longer than
posterior crossvein; anal lobe large; alula moderately well developed.

Abdomen with short, erect, scattered white hair; posterior margin of second
segment narrowly pale. Venter with posterior margins of second and third
segments narrowly pale; short white hair overall. Genital capsule brownish,
round, lateral plates moderately truncate at apex.

Female. — Unknown.

Holotype and two paratypes from P. L. Boyd Desert Research
Center, (Deep Canyon), 3 mi. S. Palm Desert, Riverside County,
California, 20 June 1969 (S. Frommer and B. Worley), malaise trap
at marker 57. One paratype male from Fish Springs, Imperial County,
California, 13 September 1965 (R. C. Dickson), from a yellow sticky-
board trap. Type deposited in U. S. National Museum.

Prorates claripennis Melander

Melander, 1906: 373.

Differs from the congeners by the black legs and black halters. The
basal antennal segments are also black. Prorates claripennis is very
similar to P. melanderi Hall. From this latter species claripennis may
be separated by the shorter proboscis, which does not extend beyond
the tip of the antennae and by the pale yellow to white, not golden,
abdominal hair.

Melander’s original description is short and as such incomplete. A
full redescription of both sexes follows :

Male. — Black, knees and fore coxae a little lighter. Halter stem and knob
black. Eyes in contact down front for distance equal to length of ocellar tubercle;
inner margin of eyes at most only slightly indented opposite antennal bases.
Small triangular front bare. Basal antennal segments short, equal in length,
each segment with two or three short yellowish hairs; third segment in lateral
view elongate conical, flattened, one and one-half times longer than two basal

42

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48 , NO. 1

segments combined, microscopically pubescent; arista terminal, minute, not
distinctly separated from third segment. Face receding, bare above, pale yellow
to white hair below. Proboscis projecting, short, not reaching beyond apex of
antennae; hair on labellae pale. Palpi testaceous, approximately one-third length
of proboscis, hair short, pale. Occiput vaguely cinereous pollinose with short,
scattered white hair.

Mesonotum vaguely cinereous pollinose with two median darker vittae; hair
short, yellow; bristles yellowish. Pleura cinereous pollinose, essentially bare.
Hair on legs pale yellow; hind tibia and tarsus of equal length. Pulvilli as long
as claws. Squama pale, whitish with fringe of long pale hair. Scutellum cinereous
pollinose, hair short, fine, scattered, pale; two apical bristles dark brown.

Wing hyaline, stigma dark brown, veins dark, only vaguely reaching posterior
margin. Costa terminates at fourth vein; petiole at base of second posterior
cell shorter than r-m crossvein; discal and second basal cells of nearly equal
length; petiole on anal cell one and one-half times longer than posterior crossvein;
alula moderately well developed.

Hair on abdomen pale yellow to white; posterior margin of third segment
narrowly white in ground color. Venter with white hair. Genital capsule with
y ellowish hair, elongate and narrowly rounded apically.

Female. — Very much like male except legs lighter colored. Knob of halter
brownish yellow. Eyes at narrowest point separated by nearly one and one-half
times width of ocellar tubercle. Eyes definitely emarginate opposite bases of
antennae. Posterior margins of third and fourth segments narrowly pale; lateral
margin from base to sixth segment pale; basal sternites with posterior margins
narrowly pale. Otherwise as described for male.

Other than the holotype and allotype, both from New Mexico, I have
seen no other specimens. Both are in the U. S. National Museum.

Prorates arctos Hall, new species

The long proboscis, dark color, well pronounced wing venation and
the northern distribution will serve to distinguish this species.

Female. — Black, tips of tibiae, knees and knob of halter pale; legs fuscous;
head and thorax somewhat grayish pollinose. Eyes at narrowest point separated
by more than width of ocellar tubercle. Front bare, slightly concave in front of
ocellar tubercle. Eyes emarginate opposite bases of antennae. Basal antennal
segments equal in length, each with a few short pale hairs; third segment slightly
longer than two basal segments combined, elongate oval, microscopically pubescent;
arista small, terminal. Face bare above, scattered pale hair below. Proboscis
projecting nearly twice eye height beyond oral margin. Palpi testaceous, tips
darker, with pale hair. Hair on occiput pale, short, scattered.

Mesonotum with two dark median vittae; hair short, yellowish; bristles brown.
Pleura bare. Coxae and hind femur with short pale hair; hind tibia and tarsus
of nearly equal length; pulvilli as long as claws. Squama white with fringe of
long white hair. Scutellum with yellow hair (bristles apparently broken off) .

Wing veins dark; stigma well pronounced. Wing hyaline; petiole at base of
second posterior cell extremely short; second basal cell slightly longer than

JANUARY 1972] HALL— NEW HETEROTROPINE BOMBYLIIDAE

43

discal cell ; petiole on anal cell much longer than posterior crossvein ; alula
well developed.

Abdomen rather evenly blackish brown; hair golden yellow; venter with lateral
margins of segments one to four pale; hair pale yellow.

Male. — Unknown.

Holotype female from 10 mi. S. Shoshoni, Fremont County,
Wyoming, 3 July 1965 (F. R. Holland) as prey of the robberfly
Heteropogon wilcoxi James. Deposited in the U. S. National Museum.

This fly represents the most nothern record for the genus Prorates
which was hitherto known only from New Mexico and southern Cali-
fornia. It seems probable that specimens should be found throughout
the Great Basin.

Porrates melanderi Hall, new species

This species consists primarily of the series of specimens which
Melander collected in Borrego, California and which he considered to
be the same as P. claripennis from New Mexico. I have three additional
specimens from Borrego.

The resemblance to P. claripennis is remarkably close. Prorates
melanderi may be distinguished from P. claripennis by the golden
yellow hair on the abdomen, the lighter colored legs and the slightly
longer proboscis.

Male. — Brownish black; basal antennal segments light brown; legs testaceous;
knob of halter dark. Eyes in contact in middle of front for a distance shorter
than ocellar tubercle. Inner margin of eyes incised opposite bases of antennae.
Small triangular front bare, cinereous pollinose. First two antennal segments
equal in length, first segment with a few short, fine, pale yellow hairs at apex;
third segment twice as long as two basal segments combined, in lateral view
elongate conical, microscopically pubescent; arista minute, terminal. Face bare,
few scattered pale hairs below oral opening. Proboscis projecting shorter than
height of eye beyond oral margin but reaching beyond tip of antennae. Palpi
yellowish, at least half as long as proboscis, with short pale yellow hair. Occiput
cinereous pollinose with short pale hair, that on ocellar tubercle yellowish.

Mesonotum cinereous with two darker median vittae; hair short, yellow;
lateral bristles yellow. Pleura cinereous especially below, bare except for few
pale hairs on mesopleura. Coxae and legs with short pale yellow to white hair,
that on femora very short and setiform; hind coxa darker than fore and mid-
coxae; hind tibia longer than hind tarsus. Pulvilli as long as claws. Squama white
with fringe of long pale hair. Scuttellum blackish brown, somewhat cinereous
pollinose ; two marginal bristles testaceous ; hair white.

Wing hyaline, veins pale brownish; petiole of second posterior cell longer
than r-m crossvein; discal cell small, nearly one-half length of first basal cell;
petiole on anal cell longer than posterior crossvein; alula developed, rounded.

Abdomen dark brown with golden-yellow hair, may be lighter towards base

44

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

of abdomen; posterior margins of second and third segments narrowly white;
lateral margin of first three segments white; posterior margins of second and
third sternites narrowly whitish. Genital capsule brown with yellow hair, tri-
angular in lateral view with apex bluntly rounded.

Female. — Grayish-black; basal antennal segments darker than in male; knob
of halter pale yellowish. Eyes separated at narrowest point by at least width
of ocellar tubercle ; lateral margin of abdominal segments one to five white.
Otherwise as described for the male.

Holotype male and allotype from Palm Canyon, Borrego, Imperial
County, California, 3 May 1945 (A. L. Melander) . In U. S. National
Museum.

Nine paratopotypes and three paratypes, Borrego, Imperial County,
California, 3 May 1956 (P. H. Timberlake) in the authors collection.

Prorates nigrescens Hall, new species

This species is readily differentiated from the others by the black
hair on the mesonotum and the black scutellar bristles as well as by
the darker knob of the halter.

Male. — Blackish brown; palpi and legs brownish yellow; posterior margins
of first four abdominal segments narrowly white. Eyes contiguous down most
of front. Ocellar tubercle and small triangular front bare. First two antennal
segments short, small, of nearly equal length, both microscopically white
pollinose; third segment in lateral view conical, a little more than twice as
long as wide, microscopically pubescent; arista apical, minute. Face receding,
cinereous pollinose below. Proboscis projecting, reaching beyond tip of antennae.
Palpi nearly one-third length of proboscis. Underside of head and lower half of
occiput with whitish hair.

Mesonotum when viewed from in front shows three grayish brown median
vittae which do not reach scutellum; mesonotal hair short, scattered, erect, black,
longer towards scutellum; one prealar and one postalar bristle, both pale in color.
Pleura whitish pollinose, bare with few short, fine, pale hairs on mesopleuron.
Legs apilose and asetose, covered with minute hair-like setulae; posterior tarsus
a little longer than posterior tibia; pulvilli nearly as long as claws. Stem of
halter pale, knob black or dusky. Squama dusky with fringe of long pale hair.
Scutellum with short black hair with two median, strong, black, marginal bristles.

Wing hyaline, stigma dark brown, anterior veins brown, rest pale; petiole on
second posterior cell longer than r-m crossvein; vein M 3 +Cui reaches hind
margin of wing; petiole of anal cell longer than posterior crossvein; discal cell
a little shorter than second basal cell; anal lobe large; alula moderately developed.

Abdomen sparsely white haired. Genital capsule with yellowish hair, in lateral
view elongate, narrowed apically.

Female. — Eyes at vertex separated by one and one-half times width of ocellar
tubercle. Head cinereous pollinose. Front bare with a narrow longitudinal median
depression. Lower half of pleura and all of legs yellowish; spines on ovipositor
pale yellow. Otherwise as described for the male.

JANUARY 1972] HALL NEW HETEROTROPINE BOMBYLIIDAE

45

Holotype male and allotype from Riverside, Box Springs Mts.,
Riverside County, California, 4 July 1969 (J. C. Hall, M. E. Irwin)
malaise trap. Both in U. S. National Museum.

One paratopotype and one paratype, topotypic except collected 3
October 1967, in the author’s collection.

Prorates frommeri Hall, new species

This species differs from the congeners by the pale yellow basal
antennal segments, pale hair on the mesonotum and by the pale legs.

Male. — Brownish, legs, basal antennal segments, thorax somewhat, venter of
abdomen and genitalia pale yellow. Eyes contiguous down most of front; ocellar
tubercle and small triangular front bare. Basal antennal segments small, at most
with few short hairs; third segment long conical, nearly four times longer than
wide, microscopically pubescent; arista terminal, minute. Proboscis projecting
beyond oral margin for distance nearly equal to height of eye. Palpi short, pale
with pale hair. Face bare. Underside of head and lower half of occiput with
white hair.

Mesonotum, when viewed from in front, with three lighter colored median,
longitudinal vittae which become coalesced in front of scutellum, lateral margin
brownish yellow; hair short, fine, erect, yellow; bristles yellow. Pleura bare
small patch of fine hair on mesopleuron. Long hair on legs wanting, femora
and tibiae with small setaceous hair; bristles wanting. Pulvilli as long as claws.
Halter stem and knob yellowish white. Squama white with fringe of white hair.
Scutellum brownish, hair short, scattered, yellow, two convergent marginal
bristles testaceous.

Wing hyaline, stigmal area at most only faintly colored; anterior veins brown,
rest pale; second posterior cell long petiolate, petiole as long as posterior cross-
vein; second basal cell much longer than discal cell; petiole on anal cell much
longer than length of posterior crossvein; no veins reach posterior margin of
wing; alula moderately well developed.

Abdominal dorsum with short, scattered yellow hair; posterior margins of
nearly all segments narrowly pale in ground color. Venter with yellow hair.
Genital capsule brownish, in lateral view elongate triangular, lower apical corner
slightly elongated.

Female. — Head cinereous pollinose; eyes widely separated. Body and legs
entirely pale yellowish or straw colored. Otherwise as described for male.

Holotype male and allotype, from P. L. Boyd Desert Research
Center, (Deep Canyon) 3 mi. S. Palm Desert, Riverside County,
California, 1 October 1969 (S. Frommer), malaise trap at marker
#57. In U. S. National Museum.

Two hundred twenty-one paratopotypes collected from 1 June 1969
to 4 October 1969, in a malaise trap.

There is a rather wide range of color expressed from nearly entirely
brown (except the legs) to entirely straw colored. The majority of the

46

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

specimens are of this latter color, although the males generally tend
to be darker. I can find no structural differences between the color
variants of this species.

Caenotus Cole
(Figs. 4, 8)

Caenotus Cole, 1923: 14; Melander, 1950: 148.

Caenotus was originally described as a therevid. Cole based his
decision on the placement of the genus in the Therevidae primarily
because of the five posterior cells found in the wing of the genotype
species, C. inornatus Cole. An examination of part of the type series
of C. inoratus shows that although there are five posterior cells in each
wing the longitudinal veins forming the extra cell arise from the discal
cell. In the Therevidae the posterior vein forming the fifth posterior
cell arises from the small crossvein at the base of the discal cell. The
other three species of Caenotus : C. canus Melander, C. minutus Cole
and C. hospes Melander, all have only four posterior cells.

Melander (1927) transferred Caenotus to the Bombyliidae. In 1950
he recharacterized the genus and gave a key to the species together
with the descriptions of two new species. I can add nothing more to
Melander’s remarks.

Caenotoides Hall, new genus

(Figs. 5, 6, 7)

The new species described below were discovered during the course
of study for this report. The one striking difference noted about the
specimens was that the third antennal segment terminated in a
microscopic tuft of hair. The only other genus of bombyliid sharing
this characteristic is Anthrax, a group to which these specimens ob-
viously do not belong.

Caenotoides is similar in habitus to Caenotus but differs by the
shorter and less abundant pile on the body, the shorter one-segmented
palpi, the tuft of hair at the tip of the third antennal segment which
is not separated by a distinct suture and by the costa terminating at
the third longitudinal vein or only slightly beyond.

Small flies, up to 4 mm in length. Eyes of male contiguous, widely separated
on female; lower half of eye of male more finely faceted than upper half,
facets of female uniform in size. Ocellar tubercle large, ocelli form equilateral
triangle. Antennae approximate at base, first two segments nearly equally short,
both broadened apically; basal half of third segment short, broad, apical half
styliform, about as long as basal portion, terminating in tuft of minute hair.

JANUARY 1972] HALL— NEW HETEROTROPINE BOMBYLIIDAE

47

4

5

Figs. 4 & 5, wing of holotypes. Fig. 4. Caenotus hospes. Fig. 5. Caenotoides
californica. Figs. 6-8, lateral view of antenna. Fig. 6. Caenotoides califormca.
Fig. 7. Caenotoides idahoensis. Fig. 8. Caenotus hospes.

43

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Oral opening reaches base of antennae. Proboscis short, about half as long as
oral opening; palpi very small, one-segmented. Face rounded, not protruding,
receding below. Occiput flat. Head slightly wider than width of thorax. Mesonotum
rounded, without apparent bristles, hair short and sparse. Legs with short
setula-like hair; pulvilli present, much shorter than claws. Scutellum rounded
without marginal bristles. Wing hyaline, at most stigmal area slightly colored;
two submarginal cells; four posterior cells, all open in wing margin; costa
terminates at tip of third vein or only slightly beyond; apical portion of vein R 5
from branch of R* to tip of wing longer than basal portion from branch of R* to
r-m crossvein; medial crossvein at base of third posterior cell arises from discal
cell; discal cell much smaller than either basal cell; anal cell closed before
wing margin and petiolate; anal lobe large; alula small. Abdomen of male with
seven visible segments, female with eight. Genital capsule large, rounded,
medially divided.

Genotype . — Caenotoides calif ornica Hall, described below.

Key to Caenotoides Species

1. Third antennal segment gradually attenuated to styliform apical one-half,

legs not black 2

Base of third antennal segment broad then suddenly constricted to apical
styliform portion; legs black, tibiae testaceous californica Hall, n. sp.

2. Legs flavo-testaceous ; abdomen pale yellowish with basal black stripe

on second segment mexicana Hall, n. sp.

Legs dark brown; abdomen white or whitish, base of second segment
black , idahoensis Hall, n. sp.

Caenotoides californica Hall, new species

(Figs. 5, 6)

Male. — Black, subshining, abdomen other than base of second segment, venter
and genitalia, white. Legs black, tibiae testaceous; halter stem brown, knob
white. Vertex vaguely whitish pollinose. Hair on ocellar tubercle short, erect,
white. Small triangular front bare, whitish pollinose. Basal antennal segments
equal in length, second segment much broader apically than the first, both with
few minute pale hairs; base of third segment, in lateral view, not as broad as
second segment but broadly flattened, suddenly and sharply constricted to apical
styliform half which terminates in a tuft of microscopic hair or setae (Fig. 6).
Face whitish pollinose laterally near antennae, white hair at sides below. Oral
opening wide, reaching base of antennae or nearly so. Proboscis short, black.
Palpi extremely small, black. Lower half of occiput with long pale hair, upper
half nearly bare except for a few short hairs next to eye margin.

Mesonotum, when viewed from in front, with three vague brownish median
vittae which do not reach scutellum; pile pale, short, scattered. Pleura hare
with few short fine hairs on meso- and stenopleurae. Hair on legs setiform,
not long or dense; apical tibial bristles small, fine. Squama white with fringe
of white hair. Scutellum brownish pollinose, hair white, scattered, fine, those
hairs along posterior margin a little stronger but not bristle-like.

Wing hyaline, veins brown ; costa terminates slightly beyond apex of third

JANUARY 1972 ] HALL NEW HETEROTROPINE BOMBYLIIDAE

49

longitudinal vein, ambient vein not evident; third posterior cell of nearly equal
width throughout; petiole on anal cell equal to length of posterior crossvein.

Abdomen elongate, cylindrical with short, fine, scattered pale hair. Venter
with much shorter pale hair. Genital capsule rounded, shiny, lateral pieces with
short white hair, apex rounded.

Female. — Front whitish yellow pollinose, small, narrow, median black stripe
just below median ocellus. Mesonotum brownish yellow pollinose without apparent
vittae; humeral and postalar calli a little lighter in color than notum; hair on
mesonotum pale yellow. Pleura black, upper part of sternopleuron and small
spot on posterior portion of mesopleuron yellowish. Abdominal dorsum with
small blackish spots on sides of second to fifth segments, sixth and following
segments entirely black as is middle of venter. Each side of ovipositor with
five stout, blunt spines. Female otherwise as described for male.

Holotype male and allotype from 3 mi. N. of North Palm Springs,
Riverside County, California, 25 February 1968 (M. E. Irwin).
Both in U. S. National Museum.

These specimens were take in copulo and are mounted on one pin,
the male being uppermost.

These flies were collected in a sand dune association. They were
flying close to the ground among the sparse grass and when captured
they were very reluctant to rise into the net. Only one other male was
collected at the same time. There were probably more specimens but
because of their small size and camouflaged appearance they were
overlooked. Repeated attempts to collect more specimens have brought
negative results.

Caenotoides idahoensis Hall, new species

(Fig. 7)

Very similar to C. calif ornica. Differs primarily in the shape of the
third antennal segment and the blunt apices of the lateral pieces
enclosing the genitalia.

Male. — Basal portion of third antennal segment gradually constricted to apical
styliform one-third. Mesonotum entirely black. Wing veins yellow. Black mark
at base of second abdominal segment not dark, rather obscure. Extreme lateral
margins of abdominal segments four to seven yellowish. Legs dark brown.
Otherwise identical to description of C. calif ornica.

Holotype male from Bruneau, Owyhee County, Idaho, 23 June
1938 (H. M. Harris). In U. S. National Museum.

Caenotoides mexicana Hall, new species

Quite similar to the congeners. Differs mainly by the flavotestaceous
legs and the absence of the black stripe across the base of the second
abdominal segment.

50

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Male. — Palpi pale yellow. Abdomen yellowish not white; mesonotum vaguely
brownish yellow pollinose without vittae. Antennae like that in C. idahoensis
except styliform portion equal in length to broadened basal portion. Body with
white hair except that on scutellum yellow. Dark stripe across base of second
abdominal segment wanting. Venter dark in middle of fourth segment only.
Specimen otherwise as described for C. californica.

Holotype male from 23 mi. E. San Luis, Baja California, Mexico,
24 April 1962 (M. E. Irwin). In U. S. National Museum. Type was
collected in a sand dune association.

Literature Cited

Bezzi, M. 1926. Notes additionelles sur les Bombyliides d’Egypte. Boll. Soc.
Roy. Entomol. Egypt, 18: 244—273 (1925).

Cole, F. R. 1923. A revision of the North American two-winged flies of the
family Therevidae. Proc. U. S. Nat. Mus., 62(4) : 1-140.

Efflatoun, H. C., Bey. 1945. A monograph of Egyptian Diptera. Part VI.

Family Bombyliidae. Sec. 1: Subfamily Bombyliidae Homeopthalmae.
Bull. Soc. Fouad d’Entomol., 29: 1-483.

Loew, H. 1873. Beschreibung europaischer Dipteren. Systematische Beschrei-
bung der bekannten europaischen zweifliigeligen Insecten, von Johann
Wilhelm Meigen. Vol. 3, 320 pp. Halle.

Melander, A. L. 1906. Some new or little known genera of Empididae. Entomol.
News, 17: 370-379.

1927. Diptera. Fam. Empididae. Fasc. 185, 434 pp. In Wytsman, P., ed.,
Genera insectorum. Bruxelles.

1950. Taxonomic notes on some smaller Bombyliidae. Pan-Pac. Entomol.,
26: 134-144, 145-156.

JANUARY 1972]

WILCOX SINTORIA REVISION

51

The Genus Sintoria Hull

(Diptera: Asilidae)

J. Wilcox

Anaheim, California 92806

The genus Sintoria Hull (1962) was proposed for S. emeralda Hull
from Mexico. Material collected in recent years has extended the range
of the genus to Baja California and north to Utah. Six species are
described as new and a key to the species is given.

Sintoria is closely related to Cyrtopogon Loew. In both genera the
mystax is composed of hairs, there is a comb of about five curved
bristles at the apex of the tibiae, and they lack strong dorsocentral and
scutellar bristles. In Sintoria the facial gibbosity arises abruptly below
the antennae to a height subequal to the length of the first antennal seg-
ment from which it recedes gradually to the oral margin ; in Cyrtopogon
the gibbosity is usually evenly rounded with the greatest height at the
middle. There are weak occipital bristles in Sintoria while in Cyrtopogon
they are usually long dense hairs. In Sintoria the scutellum is more or
less flattened and the disc largely bare of pollen and with short discal
and long marginal hairs; in the groups of Cyrtopogon with flattened
scutellum, the disc is pollinose and the hairs long erect on the disc and
posterior margin. In Sintoria the head, thorax and coxae are largely
pollinose; spots on the mesonotum, the abdomen largely and the legs
are bare of pollen, and these areas are shining black with metallic
reflections of blue, green or purple; no Cyrtopogon have been noted with
these metallic colors.

Sintoria is mainly a fall genus, occurring from August to November.
Cyrtopogon occurs mainly in the spring and summer and until recently
had not been collected in Mexico. Its range in North America in the
west, is from New Mexico to Baja California Norte and north to Alaska;
in the east it occurs from Florida to Nova Scotia and occurs in all the
Canadian Provinces; it is not found in Texas and is absent or rare in
the Great Plain States.

I am indebted to the following persons for the loan of specimens:
J. G. Rozen, American Museum of Natural History, AMNH; Mont A.
Cazier, Arizona State University, ASU; Paul H. Arnaud, Jr., California
Academy of Sciences, CAS; J. A. Powell, California Insect Survey, Uni-
versity of California, Berkeley, CIS; E. L. Sleeper, California State
College, Long Beach, CSCLB; Eric Fisher, Long Beach, Calif., EF ;
Charles H. Martin, Tucson, Arizona, CHM; P. H. Timberlake, University

The Pan-Pacific Entomologist 48: 51-58. January 1972

52

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

of California, Riverside, UCR; W. W. Wirth, Entomology Research
Division, Agr. Res. Serv., U. S. D. A., USNM, W. J. Hanson, Utah State
University, USU.

Measurements were made with an ocular micrometer: of the face
and wings at 40 times, and of the antennae at 75 times.

Key to Species

1. Mesonotum behind suture largely bare of pollen; wings lightly infuscated 2

Mesonotum behind suture largely pollinose, small intermediate spot and

postalar calli bare; wings hyaline 3

2. Hairs of face light yellow; hairs of mesonotum yellowish; abdomen bluish

green, posterior corners of male tergites 1-5 and female 1-6, yellowish

pollinose; length 8-9 mm. (Mexico) emeralda Hull

Mystax largely black; anterior mesonotal hairs on central stripe largely
black; abdomen blue black, small posterior corners of tergites 2-5, gray
pollinose; length 7-9 mm. (Baja California Sur) lagunae Wilcox, n. sp.

3. Central mesonotal crest of hairs largely black and nearly as long as antennal

segments 1-3; mystax largely and hairs of frons and ocellar tubercle,

black; length 9 mm. (Mexico) rossi Wilcox, n. sp.

Central mesonotal hairs white or yellowish and subequal in length to antennal
segments 1-2 ; mystax, hairs of frons and ocellar tubercle, white or
yellowish 4

4. Pollen on sides of abdominal tergites 2-6 extending inward on anterior

margins - 5

Pollen on sides of abdominal tergites if extending inward, on posterior
margins 6

5. Anterior pollinose fascia on male abdominal tergites 2-6 broadly separated,

entire on female 2-5; central stripe of mesonotum bare of pollen anterior
to suture and in male confluent with intermediate spots, posterior inter-
mediate spots narrowly separated from postalar calli ; length 8 mm.

(Arizona, Utah) cazieri Wilcox, n. sp.

Anterior pollinose fascia on male abdominal tergites 3-5 entire; central stripe
of mesonotum pollinose, intermediate spots small and widely separated from

each other and from postalar calli; length 7 mm. (Texas)

pappi Wilcox, n. sp.

6. Scutellum all pollinose; broad sides of abdominal tergites 1-6 gray pollinose,

broadly extending inward for short distance on posterior corners of 2-6;

length 10 mm. (California) mojavae Wilcox, n. sp.

Disc of scutellum largely bare of pollen, narrow posterior margin and basal
central spot on disc, gray pollinose; sides of abdominal tergite 1 and pos-
terior corners of 2-6, gray pollinose, the pollen extending narrowly forward

to anterior margins on most specimens; length 7-8 mm. (Arizona)

cyanea Wilcox, n. sp.

Sintoria cazieri Wilcox, new species

Male. — Length 8 mm. Head black, densely white pollinose. Hairs and bristles
white; 8 erect bristles on ocellar tubercle. Face at antennae 21/30 width of one

JANUARY 1972]

WILCOX — SINTORIA REVISION

53

eye. Antennae black; hairs white; two yellowish biistles below on segment 2;
segment 3 of nearly uniform width, narrowed on apical fourth; segments 10:11:
33:16 in length.

Mesonotum black, gray pollinose, central stripe anteriorly brown, area between
humeri and central stripe golden. Anterior intermediate bare spots broadly con-
nected across dorsum and narrowed laterally; posterior spots connected narrowly
with bare postalar calli. Hairs white, short sparse, not much longer than antennal
segment 1, longer in posterior dorsocentral rows and long erect on humeri. Two
presutural bristles yellowish, supraalars and postalars white, hair-like. Pleura and
coxae densely gray pollinose, hairs white. Disc of scutellum bare, posterior margin
and narrow base gray pollinose, 6 long white marginal hairs.

Abdomen black with greenish reflections; sides of tergites 1-7 densely gray
pollinose, pollen extends inwards narrowly on anterior margins of 1 and 3-5,
interrupted at middle by about half width of segments. Hairs short sparse white,
longer laterally; lateral bristles on tergite 1 white. Sternites 1-7 densely gray
pollinose, hairs white. Genitalia black, hairs white.

Legs black, reflections blue, on hind femora greenish. Hairs white; bristles
yellowish white; pulvilli and empodia whitish; claws black, base narrowly reddish.

Halteres yellowish brown, lower stem brown. Wings hyaline, veins brown, an-
terior crossvein at 42/66 length of discal cell, third vein branched beyond end
of discal cell, anal cell open, posterior cell 4 narrowed at wing margin.

Female. — Length 8 mm. Bristles on ocellar tubercle and hairs below on antennae,
yellowish. Anterior intermediate bare spots not connected dorsally, and posterior
spots separated from bare postalar calli. Anterior pollinose bands on abdominal
tergites 2-5 entire, narrowly separated on 6; segments 7-8 bare; apical spines
brown. Anterior crossvein at 40/67 length of discal cell, minute stump vein on
one side, posterior cell 4 not narrowed at margin.

Holotype male , Holden, Utah, 18 September 1959, CIS. Allotype
female, Beryl, Iron County, Utah, 27 September 1953 (M. Cazier)
AMNH.

Paratypes. — 1 female, Five Points, Utah, 5 September 1931 (G. F.
Knowlton, M. J. Jones) beets; antennal segment 3 missing, JW; 2
females, House Rock, Arizona, 14 September 1946 (F. H. Parker) A. L.
Melander Collection, USNM.

Named in honor of Mont A. Cazier, Arizona State University, Tempe,
a most persistent and productive collector who has supplied me with
innumerable Asilidae in the last ten years.

Sintoria cyanea Wilcox, new species

Male. — Length 8 mm. Head black, densely grayish white pollinose, frons with
slight golden tinge. Hairs white, in mystax longer on oral margin; 4^5 yellowish
lateral hairs on frons; about 6 erect yellowish bristles on ocellar tubercle; oc-
cipitals white. Face at antennae 21/33 width of one eye. Antennae black, segment
1 grayish and remainder brownish pollinose; sparse hairs below on segments 1-2
white, above brownish; two brownish bristles below on segment 1 and one on

54

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

segment 2; segment 3 slightly coarctate, tapering on apical fourth; segments
14:13:39:17 in length.

Mesonotum black, densely grayish brown pollinose; central stripe subshining;
small intermediate spots before and behind suture and postalar calli, shining
black. Hairs erect yellowish, as long as antennal segments 1-2. Bristles yellowish,
2 presutural, 2 supraalar, 3 postalar. Pleura and coxae grayish pollinose, hairs
white. Scutellum shining black, narrow posterior margin and basal semicircular
spot, grayish brown pollinose; hairs yellowish white, on disc as long as mesonotal
hairs, posterior margin with fringe of about 20 longer erect hairs.

Abdomen shining blue-black, sides of tergites 1-6 gray pollinose widening
slightly on the posterior corners of 2-5. Hairs white, long laterally, short sparse
dorsally. Sternites 1-7 gray pollinose with long white hairs. Genitalia black, long
hairs white.

Legs blue black; hairs white; bristles white becoming yellowish brown on tarsi;
claws black; pulvilli and empodia light brown.

Halteres yellowish, base brown. Wings hyaline, veins brown, anterior crossvein
at 36/70 length of discal cell, third vein branched opposite end of discal cell.

Female. — Length 7 mm. Apical spines of abdomen black. Third vein branched
beyond end of discal cell.

Holotype male , Sabino Canyon, Santa Catalina Mts., Arizona, 25
October 1962 (J. Wilcox) CAS. Allotype female, same locality, 20
October 1962, CAS.

Paratypes. — Arizona: 9 males, 19 females, same locality, 19 to 25 October 1962,
22 October 1965 (Itol J. and J. Wilcox) JW ; 3 males, 3 females, same data, 11
October 1964 (Eric Fisher) EF ; 1 female, 1.5 mi. N. E. Carrizo, Navajo Co., 28
September 1962 (P. H. Timberlake) UCR; 1 female, Huachuca Mts., 13 October
1931 (E. D. Ball) USNM; 2 males, 2 females, 3 mi. E. Pantano, 20 October 1962,
22 October 1965 (J. Wilcox) JW ; 1 male, 1 female, 7 mi. W. Peach Springs,
Mohave Co., 28 September 1964 (P. H. Timberlake) UCR; 14 males, 21 females,
2 mi. N. E. Portal, 10 October 1964 (Eric Fisher, R. R. Snelling) late afternoon
and evening, some by sweeping Tar Bush, and 3 at light, EF ; 1 male, 2 mi. N. E.
Portal, 22 October 1960 (M. A. Cazier) ASU ; 1 male, 3 females, 2 mi. N. E.
Portal, 23 October 1962 ( J. Wilcox) JW ; 1 male, 2 females, Tucson, September
1939 (R. H. Crandall) AMNH; 1 male, Tucson, 31 September 1958 (R. S. Beal)
ASU ; 1 male, Tucson, 9 July 1920, JW.

The specimens collected at the mouth of Sabino Canyon, elevation
about 3,000 feet, were mostly on the dead stems down in or among the
seed clusters and flowers of Burro Brush. Near Portal, they were found
on the stems and leaves of Tar Bush. On hot days, they were not found
until 3:30 or 3:45 p.m., but on a cloudy day they appeared at 11:30
a.m., but soon disappeared when it became cooler. Another time a few
were collected resting on the green foliage of mesquite about four
feet high at 11:00 a.m. on a warm day. Metapogon punctipennis
Coquillett was collected at the same time in both localities.

The specimens from near Portal have only the posterior corners of

JANUARY 1972]

WILCOX SINTORIA REVISION

55

abdominal tergites 2-5 pollinose and some of the specimens from Sabino
Canyon do not have the pollen on tergites 2—3 extending forward to
the anterior margins on the sides of the tergites.

SlNTORIA EMERALDA Hull

Sintoria emeralda Hull, 1962, U. S. Nat. Mus. Bull. 224: 199.

Type $, Nepantla, Mexico, 28 October 1923 (E. G. Smith) USNM; Allotype $,
La Cascada, Mexico, 28 October 1922 (E. G. Smith) USNM.

Male. — Length 7 mm. Face and frons golden, occiput gray pollinose; hairs all
yellowish: face at antennae 19/28 width of one eye. Antennae black, hairs brown;
segments 14:15:41:10 in length.

Mesonotum golden pollinose; central stripe, broad anterior and posterior inter-
mediate spots and postalar calli, shining purple, anterior spots extend narrowly
to humeri. Hairs white to yellowish and subequal in length to antennae 1-2.
Scutellum shining bluish purple, very narrow posterior margin golden pollinose;
discal and about 12 longer marginal hairs yellowish.

Abdomen bluish green; narrow posterior corners of tergite 1 and posterior
corners of 2-5, golden pollinose. Genitalia blue black, hairs yellowish.

Legs bluish green, hairs and bristles yellowish.

Halteres lemon yellow, base and lower stem brown. Wings light brown; veins
brown, at base anteriorly yellowish; anterior crossvein at 35/59 length of discal
cell; third vein branched before end of discal cell, in one wing with minute stump
vein.

Female. — Length 7 mm. Posterior corners of tergites 1-6 yellowish pollinose.
Anterior crossvein at 39/70 length of discal cell, third vein branched opposite
end of discal cell.

Additional specimens. — Mexico, Aguascalientes, 19 mi. S. Aguascalientes, Hy.
45, Km. 508, 19 August 1962 (Dorothy and C. H. Martin) ; Zacatecas, 10 mi. S.
Jalapa, 17 September 1970 (G. E. and R. M. Bohart) , Rio Grande, 4 September
1966 (G. E. and A. S. Bohart), 5 mi. N. Zacatecas, 19 September 1970 (G. E. and
R. M. Bohart) .

The following records were kindly supplied by Eric Fisher. Mexico: Guanajuato,
41 mi. N. Queretaro, 6,950 ft., 3 September 1963, (Scullen and Bolinger) ; Jalisco,
10 mi. S. W. Lagos de Moreno, 8 September 1970 (E. M. and J. L. Fisher) ; San
Luis Potosi, 19 mi. S. San Luis Potosi, 7,200 ft., 4 September 1963 (Scullen and
Bolinger) .

Sintoria lagunae Wilcox, new species

Male. — Length 8 mm. Head black, grayish white pollinose. Hairs of mystax
black, the lower fourth white; hairs on frons black; about 12 long slender black
bristles on ocellar tubercle; occipitals, beard and hairs on palpi and proboscis
white. Face at antennae 21/38 width of one eye. Antennae black, brownish
pollinose; hairs black, two bristle-like below on 2; segments 15:15:45:16 in
length.

Mesonotum black; yellowish pollinose; central stripe subshining; intermediate
spots with blue reflections, anterior ones twice as broad as long, posterior ones
longer than broad and include postalar calli. Hairs on the anterior central stripe

56

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

erect black and as long as antennae 1-2 with few yellowish anteriorly and mostly
yellowish posteriorly ; lateral hairs yellowish. Bristles yellowish, 2 presutural,
others not separable from the hairs. Pleura and coxae yellowish pollinose; hairs
on coxae white, on hypopleura yellowish. Scutellum black, narrow posterior
margin yellowish pollinose; hairs yellowish, discal hairs sparse semierect, about
20 long erect marginals.

Abdomen blue-black; posterior corners of tergites 2-5 gray pollinose. Hairs
yellowish, long on sides and short on dorsum. Sternites yellowish pollinose; hairs
long erect yellowish. Genitalia black; hairs yellowish.

Legs black with blue reflections; hairs yellowish white; bristles yellowish; claws
black ; pulvilli and empodia light brown.

Halteres yellowish red, lower stem brown. Wings very light brown, veins brown,
anterior crossvein at 37/62 length of discal cell, third vein branched slightly
beyond end of discal cell.

Female. — Length 8 mm. Face and frons with golden tinge; mystax black, hairs
on sides below yellowish. Abdominal hairs white, short on sides of tergites 6-8;
sternites 7-8 bare with short hairs; apical spines black. Anterior crossvein at
41/66 length of discal cell, third vein angulated and with suggestion of a stump
vein.

Holotype male, Mexico, Baja California Sur, Sa. Victoria, La
Laguna, 5,500 ft., 20/23 October 1968 (E. L. Sleeper, F. J. Moore)
CSCLB. Allotype female, same data, CSCLB. Types to be deposited in
Los Angeles County Museum.

Paratypes. — 92 males, 79 females, same data, CSCLB; 7 males, 2
females, La Laguna, Sierra Laguna, L. Calif., 14 October 1941 (Ross
and Bohart) CAS.

The mystax of both sexes varies from all black to the lower half
white in the males and some of the oral hairs as well as the lateral hairs
yellowish in the females. Some of the specimens show a distinct brown
tinge in the wings while others are nearly hyaline.

Sintoria mojavae Wilcox, new species

Female. — Length 10 mm. Head black, densely white pollinose. Hairs and
bristles white; 6 bristles on ocellar tubercle. Face at antennae 24/40 width of
one eye. Antennae black, brownish pollinose; hairs white; two yellowish white
bristles below on segment 2; segment 3 tapers from base to apex; segments 18:18:
45:20 in length.

Mesonotum black, gray pollinose; central stripe indistinctly brown; small inter-
mediate spots and postalar calli, shining black. Hairs white; bristles white, 2
presutural and 2 fine supraalar and postalar. Pleura and coxae gray pollinose,
hairs white. Scutellum gray pollinose, hairs white, about 12 long crinkly marginal
hairs.

Abdomen purple-black; broad sides of tergites 1-6 expanded inward on posterior
margins of 2-6, gray pollinose, fascia separated by about half width of abdomen on
2, widening gradually to 6; 7-8 bare; apical spines black. Hairs white, long on

JANUARY 1972]

WILCOX — SINTORIA REVISION

57

sides of basal segments becoming short apically ; lateral bristles on 1 white.
Sternites 1-7 gray pollinose, hairs long erect white on 1-5, dense on 1.

Femora greenish black, tibiae and tarsi blue black; hairs white; bristles yellowish
white; claws black; pulvilli and empodia light brown.

Halteres light brown, lower stem brown. Wings hyaline, veins brown, anterior
crossvein at 48/84 length of discal cell, third vein branched opposite end of discal
cell.

Holotype female , Grapevine Springs (Cushenbury Springs), San
Bernardino County, California, 10 September 1948 (J. Wilcox) CAS.

Sintoria pappi Wilcox, new species

Male.— Length 7 mm. Head black, densely white pollinose, frons with slight
golden tinge. Hairs and bristles white, on frons and 6 bristles on ocellar tubercle
yellowish. Face at antennae 19/29 width of one eye. Antennae black, gray pol-
linose; hairs and one bristle on segment 2 yellowish white; segments 13:13:36:14
in length.

Mesonotum black, densely gray pollinose; divided central stripe and broad area
behind humeri brown; small intermediate spots and postalar calli shining black.
Hairs white, anteriorly in central stripe as long as antennae 1, longer posteriorly.
Bristles yellowish, 2-3 presutural, 1 supraalar, 2-3 postalar and 3-4 anterior
dorsocentral. Pleura and coxae grayish golden pollinose, hairs white. Scutellum
black; narrow posterior margin and base of disc extending narrowly to posterior
margin at middle, gray pollinose; hairs white, about 8 longer bristle-like marginals.

Abdomen blue-black; sides of tergite 1, broad sides of 2-7 and anterior margins
of 2-6, gray pollinose, anterior fascia narrow at middle and interrupted on 2 and
6-7. Hairs long white laterally becoming shorter apically ; 4-5 white lateral
bristles on 1. Sternites gray pollinose, hairs white. Short genitalia blue black,
hairs white.

Legs green black, hairs and bristles white; claws black; pulvilli and empodia
light brown.

Halteres yellowish, base brown. Wings hyaline, veins brown except basally;
anterior crossvein at 40/66 length of discal cell; third vein branched slightly
beyond end of discal cell; anal cell broadly open.

Female. — Length 6 mm. Face 16/24 width of one eye; antennal segments
11:11:25:15 in length. Central bisecting pollen on scutellum broad. Abdomen
with blue-purple reflections; broad sides of tergites 1-6 gray pollinose, extending
inward anteriorly on 2-6, narrowly entire on 3-5; sternites 1-6 gray pollinose;
apical spines black. Halteres yellow, lower stem brown. Anterior crossvein at
28/46 length of discal cell.

Holotype male, 8.6 mi. S. W. of New Mexico State Line, Hwy. 180,
Culberson County, Texas, 2 October 1962 (Chas. S. Papp) UCR.
Allotype female, 2 mi. N. W. Mesa, Chaves County, New Mexico, 10
September 1961 (P. D. Hurd) Gutierrezia lucida, CIS.

The facial gibbosity of this species is very weak.

Named in honor of Charles S. Papp, scientific illustrator, University
of California, Riverside and noted Coleopterist.

58

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Sintoria rossi Wilcox, new species

Female. — Length 10 mm. Head black (greased), pollen on face and frons
yellowish, on occiput gray. Mystax long dense black with some yellowish white
hairs intermixed on oral margin; hairs on frons and about 10 bristle-like on ocellar
tubercle, black; occipitals yellowish white; beard and hairs on palpi and proboscis
white. Face at antennae 23/34 width of one eye. Antennae black (greased) ; hairs
black, three or four long ones below on segment 1 and two or three on 2 ; segment
3 narrowed on apical fourth; segments 15:17:45:18 in length.

Mesonotum black (largely greased), golden brown pollinose; bare intermediate
spots before and behind suture confluent and postalar calli in part bare. Central
line with dense crest of black hairs as long as antennal segments 1-3; long sparse
hairs otherwise yellowish. Two yellowish presutural bristles, supraalars and
postalars hair-like. Pleura and coxae yellowish gray pollinose, hairs yellowish
white. Scutellum black with some gray pollen laterally; dense short discal hairs
and about 20 long crinkly hairs on posterior margin, white.

Abdomen black with green-blue reflections; narrow sides of tergites 2-5 ex-
panded a little on posterior corners, grayish pollinose; 1 and 6-8 bare; apical
spines black. Hairs long yellowish white becoming shorter apically, bristles on
1 hair-like. Sternites 1-6 grayish pollinose, long sparse hairs yellowish white.

Legs black with blue reflections; hairs white; bristles yellowish; claws black;
pulvilli and empodia light brown.

Halteres light brown, lower stem brown. Wings hyaline, veins brown; anterior
crossvein at 46/79 length of discal cell; third vein branched a little beyond end of
discal cell; anal cell broadly open.

Holotype female, N. Slope, 11,000 ft., Mt. Popocatepetl, Mexico,
20 November 1946 (E. S. Ross) CAS.

Named in honor of Edward S. Ross, California Academy of Sciences,
who frequently collects rare specimens of insects other than his speciality
on his many trips to numerous parts of the world.

Literature Cited

Hull, F. M. 1962. Robberflies of the World. The genera of the family Asilidae.

Part 1; Part 2. U. S. Nat. Mus. Bull., 224: 1-430, figs. 1-29, 1 pi.;

431-907, figs. 30-35 and 1-2536.

JANUARY 1972]

ARNAUD— G DALLAS HANNA

59

(D SaUas Harnta,

lssr-iam

Paul H. Arnaud, Jr.

California Academy of Sciences, Golden Gate Park, San Francisco 94118

Doctor G Dallas Hanna, a “renaissance man,” biologist, geologist,
micropaleontologist, diatomist, malacologist, ornithologist, instrument
maker and inventor, expert in optics, printer, photographer, was born
on 24 April 1887 in Carlisle, Arkansas, and died in San Francisco, Cali-
fornia, on 20 November 1970, during his 83rd year. He joined the
Academy staff in 1919, and continued this association for 51 years. The
diversity of his interests also included the planning and direction of the
construction of the star projector for the Morrison Planetarium at the
Academy’s Instrument Shop. He adapted an Eastman Kodak Company
three color printing process for illustrating the Academy’s scientific
publications (he and his wife Margaret produced approximately 165,000
color prints). Dr. Hanna received his A.B. (in 1909) and A.M. (in
1913) from the University of Kansas and the degree of Doctor of Philos-
ophy (in 1918) from George Washington University. In May, 1970, Dr.
Hanna was presented with the honorary degree of Doctor of Sciences
by the University of Alaska.

For most of the period between 1911 and 1919, Dr. Hanna was em-
ployed by the United States Bureau of Fisheries in Alaska (first in the
Bristol Bay area and from 1913 on in the Pribilof Islands). It was on
the Pribilof Islands and later on Guadalupe Island and the adjacent
Mexican West Coast Islands that he made his most significant contribu-
tions to the field of Entomology. During 1913-1914, he made important
collections of insects and other arthropods from St. George Island,
Alaska, and these were eventually published on in 1923 by various
specialists in volume 46 of North American Fauna. Many species were
recorded from these islands for the first time, including 17 species new
to science; these collections are deposited in the United States National
Museum. A second collection, made in the Pribilofs in 1920 was pre-
sented to the California Academy of Sciences. In 1921, 86 species (16
of them new) were recorded in the Proceedings of the California Acad-
emy of Sciences, ser. 4, vol. 11, no. 14, pp. 153-195. Dr. Hanna contrib-
uted an introduction to this paper on pages 153-155. From these
combined Pribilof collections the following new species were dedicated
to him: family Tipulidae — Pedicia hannai (Alexander), family Helco-
myzidae — Heterocheila hannai (Cole), and family Muscidae — Mydaea
hannai (Malloch).

1

60

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Fig. 1. G Dallas Hanna. Photograph taken in 1966.

JANUARY 1972]

BOOK REVIEWS

61

In 1922, Dr. Hanna was a key member of an expedition to Guadalupe
Island and the islands of the west coast of Mexico aboard the motorship
Tecate, from July 9th— August 16th. An article by Hanna and A. W.
Anthony entitled, “A Cruise Among Desert Islands” was published in
the National Geographic Magazine (vol. 44, no. 1, pp. 71—99, July 1923) .
On this expedition 1138 insects and arachnids were taken by Hanna and
J. R. Slevin. The General Report of this expedition also was published
by him in the Proceedings of the California Academy of Sciences (ser. 4,
vol. 14, no. 12, pp. 217-275, text figs. 1—2, pis. 15—19, 1925). Dr. Frank
E. Blaisdell, Sr. published an article in the same volume (pp. 321-343),
recording 57 species of Coleoptera, of which 14 were new to science,
including the patronym S tibia hannai Blaisdell (family Tenebrionidae)
collected on Angulo Rock, Asuncion Island. In subsequent years, Dr.
Hanna continued to collect small lots of insects which he presented to the
Department of Entomology, the last received in 1968.

On the occasion of Dr. Hanna’s seventy-fifth birthday, volume 32 of
the Proceedings of the California Academy of Sciences was dedicated as
a “G Dallas Hanna Anniversary Volume.” The first number, published
on April 24, 1962, pages 1 through 40, contains a portrait, biography
(written by Dr. Robert C. Miller) and bibliography of 438 writings and
other scientific contributions through the year 1961. An additional 40
writings have been published, making a total of 478 contributions in a
61 year period, 1909 through 1970.

Dr. Hanna is survived by his wife, Margaret (nee Moore) Hanna,
three sisters, a brother, Dr. Marcus A. Hanna, two granddaughters, and
four great grandchildren.

BOOK REVIEW

CURCULIONIDAE TRIBE OpHRYASTINI OF NORTH AMERICA (COLEOPTERA). David G.

Kissinger, Taxonomic Publications, South Lancaster, Mass. 1970. 8 %" X 11",

238 pp., 128 figures and groups of figures. $12.00 (paper).

This is the latest of Dr. David Kissinger’s several substantial contributions to the
knowledge of the Curculionidae and, at the same time, it constitutes the first part
of a proposed revision of the weevil subfamily Leptopiinae in North and Central
America. The major portion of this book is devoted to a revision of the 48 species
of Ophryastes. These are large, broadnose weevils occurring in the arid regions
of the western United States and northern Mexico. In addition to the thorough
taxonomic treatment of Ophryastes, information is included on the generic char-
acters and relationships of Deracanthus, a Central Asian genus which shares the
tribe with Ophryastes.

62

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48 , NO. 1

In a section entitled “Ecology” Kissinger summarizes what little is known about
the biology of Ophryastes. The paucity of information on this subject reflects the
general lack of attention which entomologists have given to the study of weevil
biology. The weevils under consideration here would seem to lend themselves well
to such studies as they are large, sometimes relatively abundant, tend to form
well-defined local populations, and above all, they should be interesting. Perhaps
the present revision will provide the necessary impetus for investigation of their
biology.

Four species of Ophryastes are described as new. Nomenclatural changes involve
new synonymy for 3 generic and 20 specific names and new combinations for 29
names. Most of the species are placed in 3 species-groups on the basis of characters
of the female genital tube; each of the 3 groups is further divided into species-
subgroups. Eight species are not assigned to groups. The key works quite well
for the determination of species of Ophryastes. One should have little or no diffi-
culty correctly determining the large majority of members of the genus, especially
if the key, descriptions and the many illustrations are carefully used together. The
male and female genitalia are described more thoroughly than is usual in papers on
weevils; otherwise the species descriptions and other taxonomic data are of the
usual kind for such studies. Twelve pages are devoted to a summary of measure-
ments of external characters of both sexes (where available) of the 48 species.

The list of references includes only those not cited by Kissinger in his 1964
generic key to the Curculionidae in America north of Mexico. It appears to me that
the desirability of having this book complete within itself would have overshadowed
the cost of adding the few additional pages required for a complete reference list.

The present book is profusely illustrated with photographs and line drawings in a
manner similar to Dr. Kissinger’s earlier work on weevils of the subfamily Api-
oninae. These illustrations and their accompanying explanations cover 142 pages.
The photographs (of which there are several hundred) show dorsal, lateral and
sometimes ventral habitus views, as well as views of specific external structures of
taxonomic importance, and male and female genitalia. These photographs are
mostly of excellent quality and add greatly to the taxonomic value of the publica-
tion. Because of the exceptionally large number of illustrations included, those
referring to a given species are often arranged on widely separated pages. This,
however, is only a minor inconvenience and does not detract from the general
usefulness of the work.

Distribution maps, and tables and figures of statistical data involving principal
component and multiple discriminant analyses cover 22 pages. An index to the
new species described and new nomenclatural changes is also included.

Dr. Kissinger has once again demonstrated his ability to do high quality work
on a taxonomically difficult group of weevils. The present revisionary study
places the taxonomy of Ophryastes on firm ground and provides the information on
which subsequent workers may base much-needed biological and population studies
of the genus. — Horace R. Burke, Texas A&M University, College Station, 77843.

JANUARY 1972]

SCIENTIFIC NOTES

63

SCIENTIFIC NOTE

Thomas Wrentmore Cook Entomology Library. — The entomological library
of the late Dr. Thomas Wrentmore Cook (188T-1962) was donated to the California
Academy of Sciences by his widow, Mrs. Mary Manning Cook (now Mrs. George
Wale) of Oakland, California. Arrangements for this important donation were
made by Dr. and Mrs. Edward L. Kessel.

The Cook library on Formicidae, Hymenoptera, and General Entomology con-
sists of 289 bound volumes (including 51 books and bound volumes of the papers
of the renowned ant authority William Morton Wheeler), 25 parts of serials,
11 unbound volumes, and 1312 “reprints.” The 51 volumes of the books and
papers published by Dr. Wheeler are a virtually complete collection of the 470
items by this world ant authority, and there are probably few such sets in
existence.

The library also includes numerous papers on world ants by many authors,
including bound volumes of the works of Arnold, Clark, Creighton, Donisthorpe,
Emery (7 volumes), Escherich, Fielde, Forel, Huxley, Janet (2 volumes), Mayr,
McCook, M. R. Smith, and Wasmann; titles on Hymenoptera such as “Catalogus
Hymenopterorum,” vols. 1-10, by Dalla Torre, 1892-1902, “Nouvelle Methode de
Classer les Hymenopteres et les Dip teres,” vol. 1, “Hymenopteres,” by L. Jurine,
1807, and “Additions et Corrections au Volume II de la Faune Entomologique du
Canada traitant des Hymenopteres” by L’Abbe L. Provancher, 1889; and general
works as “The Class Insecta” by Baron Cuvier (with supplementary additions by
Griffith, Pidgeon, and Gray), 1832, “Uber entomologische Sammlungen,
Entomologen & Entomo-Museologie” by Walther Horn and Ilse Kahle, 1935-1937,
and “Histoire Naturelle des Fourmis, et recueil de Memoires et d’Observations sur
les Abeilles, les Araignees, les Faucheurs, et autres insectes” by P. A. Latreille,
1802.

Dr. Cook, who wrote the book “The Ants of California” (Pacific Books, Palo
Alto, pp. i-xiii, 1-462, illustrated, 1953), had a life-time interest in the Formicidae,
having first undertaken graduate studies on this group of insects with Wheeler
in 1932. It was mainly during the last fifteen years of his life, from 1948 on, that
he assembled this extraordinary library on the ants of the world and on the
order Hymenoptera. — Paul H. Arnaud, Jr., California Academy of Sciences, San
Francisco, 94118.

SCIENTIFIC NOTE

The Lost Type of Brachinus sallei Chaudoir (Coleoptera: Carabidae). —

In my 1970 revision of North and Middle American Bombardier Beetles (Quaest.
Entomol., 6: 4-215) I stated that the type(s) of Brachinus sallei Chaudoir was
presumed lost. The basis for this statement was, first, the indication by Chaudoir
that he saw the specimens he described in Salle’s collection, and second, the sub-
sequent search in parts of this collection by George E. Ball in the Museum National
d’Histoire Naturelle, Paris and by E. Taylor in the Hope Museum, Oxford, at my
request. The specimen was not found in either museum where the type(s) should
have been. Then, quite unexpectedly, during a recent visit to the British Museum
(Natural History), supported in part by the American Philosophical Society, I

64

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

discovered a specimen of Brachinus sallei labelled “Orizaba, Mexico, Salle Coll.,
type,” “108” on green paper, and further labelled “Brachynus Sallei Chd. after
Salle, Sallei Chaud.”

Consequently, this specimen, a male, is herewith designated LECTOTYPE. The
type locality was originally stated by Chaudoir to be Mexico, and later restricted
by me to Tabasco, Mexico on the basis of material seen; but Salle’s label “Orizaba,
Mexico . . indicates the real TYPE LOCALITY. This locality record also
indicates a wider distribution than I previously reported; the northern most
locality then known was Lake Catemaco, Veracruz, Mexico. — T. L. Erwin,
Smithsonian Institution, Washington, D. C., 20560.

SCIENTIFIC NOTE

Observations on the Life History of Bombardier Beetles: Mechanism
of Egg-Hatching (Coleoptera: Carabidae). — In 1967 Dr. George Ball and I
collected live specimens of various species of Brachinus throughout Mexico. These
were kept alive in plastic bags until we returned to the University of Alberta,
Canada. Subsequently, the specimens were sorted to species and placed in
refrigerator trays containing moist peat, and maintained at room temperature. The
specimens were fed injured insects and various cuts of beef. The colony of
Brachinus mexicanus Dejean subsequently began mating and ovipositing, in a
manner I have described previously [Erwin, 1967. Coleopt. Bull., 21(2): 41-55].
The eggs in this case were not rolled in mud as they normally would be, because
the peat substrate was unnatural. This allowed me to make continuous observations
on the developing embryo through the very thin-walled egg. The development of
the embryos required 7-10 days.

Most carabid larvae have one or two egg-bursters or spines on the frons with
which the egg membranes are rasped and broken. The frons of known Brachinus
larvae are smooth with no evidence of egg-bursters.

The young embryo of B. mexicanus develops in a C-shape within the ellipsoid
egg; the frons, clypeolabral area, and dorsum of the mandibles are against the
side wall, rather than one end. As the young larva grows, the point of one mandible
presses against the egg membrane. At the proper growth stage, the wall is pierced
by this mandible, and is probably helped by some muscle contraction. In dry,
mud-encased eggs (the normal condition) the amniotic fluid moistens the dry mud
covering the egg and allows the first instar to struggle free. It’s subsequent be-
havior and development has been described before (Erwin, ibid.). — T. L. Erwin,
Smithsonian Institution, Washington, D. C., 20560.

JANUARY 1972]

ZOOLOGICAL NOMENCLATURE

65

ZOOLOGICAL NOMENCLATURE: Announcement A. (n.s.)88

Required six-month’s notice is given of the possible use of plenary powers by
the International Commission on Zoological Nomenclature in connection with the
following names listed by case number:

(see Bull. Zool. Nomencl. 28, pts. 1/2, 10 August 1971) :

1937. Suppression of Papilio actaeon Fabricius, 1775 (Insecta, Lepidoptera)

(see Bull. Zool. Nomencl. 28, pts. 3/4, 8 December 1971)

1954. Type-species for Sminthurinus Borner, 1901 (Insecta, Collembola)

1962. Validation of Polyzonium germanicum Brandt, 1837 (Diplopoda)

(see Bull. Zool. Nomencl. 28, pts. 5/6, 31 December 1971)

1966. Type-species for Heniola Uvarov, 1940 (Insecta, Orthoptera)

Comments should be sent in duplicate, citing case number, to the Secretary,
International Commission on Zoological Nomenclature, c/o British Museum (Natural
History), Cromwell Road, London S.W. #7, England. Those received early enough
will be published in the Bulletin of Zoological Nomenclature. — W. E. China,
Assistant Secretary.

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66

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

PACIFIC COAST ENTOMOLOGICAL SOCIETY

F. L. Blanc D. G. Denning M. S. Wasbauer P. H. Arnaud, Jr.

President President-elect Secretary Treasurer

Proceedings

Three Hundred and Thirty-Eighth Meeting

The 338th meeting was held Friday, 19 February 1971, at 7:45 p.m. in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park,
San Francisco. President Blanc presided.

Members present (34) : R. P. Allen, P. H. Arnaud, Jr., W. E. Azevedo, D.
Bacon, J. Benedict, G. S. Benham, Jr., F. L. Blanc, I. Boussy, G. Brady, R.
Bush'nell, S. L. Clement, Karen S. Corwin, A. R. Dutton, J. F. Emmel, F. Ennik,
K. E. Frick, E. Grissell, J. Guggolz, K. S. Hagen, J. T. Hjelle, E. A. Kane, R. L.
Langston, H. B. Leech, R. Lem, D. P. Levin, A. E. Michelbacher, A. R. Moldenke,
C. B. Philip, H. I. Scudder, R. E. Stecker, J. W. Tilden, M. S. Wasbauer, S. C.
Williams, D. L. Wilson.

Visitors present (10) : V. Cheney, Alexandria Ennik, T. M. Glenine, K. Heston,
Netta Leong, Martha Michelbacher, Alison Moldenke, Hazel Tilden, Constance
Wilson, R. Wong.

The minutes of the meeting held 18 December, 1970, were summarized.

The following names were proposed for membership: C. D. Langston, Jeannette
N. Wheeler, Harriet B. Reinhardt.

President Blanc asked for introductions from the floor. Mr. Steve Clement
introduced John Benedict, a graduate student at U.C. Davis. Dr. Ron Stecker
introduced Don Henley, a student at San Jose State College, recently returned from
Viet Nam.

President Blanc read an announcement of the First International Congress of
Systematic and Evolutionary Biology which will be held at the University of
Colorado, Boulder, from 4-11 August 1973. The Society of Systematic Zoology
and the International Association for Plant Taxonomy are co-sponsors of this
midterm congress of botanical and zoological interaction at the international level.

President Blanc announced that the first thirteen volumes (1924-1937) of the
Pan-Pacific Entomologist are still on sale to members at the reduced price of $1.50
per volume or $19.50 for the entire thirteen. At today’s prices, these would run
$10.00 per volume.

President Blanc announced his appointment of Dr. J. W. Tilden to the Program
Committee, and Dr. Fred G. Andrews to head up an ex-officio refreshment com-
mittee.

He also pointed out that the date given on the meeting notice cards for the
March meeting was in error. There will be no March meeting. The next scheduled
meeting will be 16 April. Mr. Ian Boussy asked why we are having so few meetings
and was answered by Dr. Stecker, who stated that the number of meetings follows
the tradition of past years with three meetings in the Spring and three in the
Fall, but that the policy is flexible, and if the membership wishes to have more
the Program Committee will do its best to find additional speakers. Dr. Stecker

JANUARY 1972 ]

PROCEEDINGS

67

then asked that anyone with suggestions on a site for the annual picnic and field
day contact him as soon as possible, so plans can be made for this year.

The following notes were presented:

The red dragonflies of Tokyo. — There was an apparent mistaken reference
to “red horseflies” (Diptera: Tabanidae) in a recent World Press broadcast, rather
than to red dragonflies (Odonata) which was actually intended. The reference was
in a broadcast on Channel 9 Television in early November, and a letter of inquiry
to the participant, Professor Chalmers Johnson at the Center for Chinese Studies,
University of California, Berkeley, brought a letter and a copy of the Japanese
paper Asahi used. The insect pictured was the characteristic and well-known red
dragonfly of Tokyo, which had made the news because it has largely disappeared
as a result of air pollution. Dr. Johnson commented: “The government is making
an effort both to control smog and to bring back red dragonflies.” The newspaper,
showing a photograph of a man holding a red dragonfly on the Ginza in Tokyo,
was exhibited at the meeting. — C. B. Philip, California Academy of Sciences.

F. C. Hottes, 1899—1970. — Frederick Charles Hottes, born October 20, 1899,
died at Grand Junction, Colorado, 27 October 1970. His brother, Howard H. Hottes,
has sent the following information to the Pacific Coast Entomological Society.

Fred Hottes was born at Mascoutah, Illinois. He graduated from Mt. Lincoln
High School in Palisade, Colorado, in 1919, obtained his B.S. from Colorado
Agricultural College (now Colorado State University), Fort Collins, in 1923,
his M.S. from Iowa State College in 1925, and his Ph.D. in entomology from the
University of Minnesota in 1927. He was Professor of Biology at James Milikin
University, Decatur, Illinois, 1928-1947, and head of the department from 1929
until his retirement in 1947. At that time he returned to Grand Junction, Colorado,
to care for his widowed father, H. G. Hottes, who passed away in 1957. Fred
Hottes began publishing descriptions of new species of aphids before receiving
his doctorate, and one of his early major papers was ‘The plant lice, or Aphididae
of Illinois’ with T. H. Frison, 1931. He continued to publish after his retirement,
and had a lifelong special interest in the genus Cinara. He published on aphids
in co-authorship with other workers, notably E. 0. Essig, and on surface and
shore bugs (veliids, halobatids, saldids, etc.) with C. J. Drake.— Hugh B. Leech,
California Academy of Sciences, San Francisco.

The Sonora Blue — 1971 the earliest season for the north. — Philotes
sonorensis (Felder & Felder) adults were collected in late January in Santa Clara
County, California, and in abundance in early February in El Dorado and Placer
counties. It appears that the “peak” flight for 1971 has already occurred, and
adults will be scarce in March or April at the lower elevations. Almost half of the
males in these northern colonies have a few orange scales on the hindwings cor-
responding to the prominent orange splotch on the females. This has not been
observed on numerous Southern California specimens. There is also variation in the
number and location of the black spots on the forewings within the colonies through-
out the species range from Northern to Baja California. Although the percentages
may vary from coast to desert, or from north to south, no subspecific status seems
to be warranted. These populations are referred to as “colonies,” as P. sonorensis
is a very weak flier and is seldom found far from its Dudleya host. The acceptable
stonecrop species (Crassulaceae) have such a spotty distribution, that one wonders
why the insect shows so little variation, or how some of the more isolated colonies
could have become established. — Robert L. Langston, Kensington, California.

68

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

The principal speaker of the evening was Dr. J. W. Tilden, San Jose State
College, emeritus. His beautifully illustrated talk was entitled “Insects and their
Western Habitats.”

Coffee and other refreshments were served during a social hour in the entomology
rooms following the meeting. — M. S. Wasbauer, Secretary.

Three Hundred and Thirty-Ninth Meeting

The 339th meeting was held Friday, 16 April 1971, at 7:45 p.m. in the Morrison
Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Blanc presiding.

Members present (28) : R. P. Allen, F. G. Andrews, P. H. Arnaud, Jr., W.
Azevedo, F. L. Blanc, G. Brady, T. Briggs, S. L. Clement, J. E. Cronin, D. G.
Denning, A. R. Dutton, J. F. Emmel, F. Ennik, W. E. Ferguson, J. R. Gabel, L.
Gormley, T. E. Hewton, Jr., R. L. Langston, H. B. Leech, G. A. H. McClelland,
A. E. Michelbacher, C. B. Philip, E. S. Ross, 0. Shields, R. Stecker, R. W. Thorp,
M. S. and J. S. Wasbauer, R. H. Whitsel, S. C. Williams.

Visitors present (21) : Madelaine Arnaud, J. Bass, Grace Bass, D. Carroll,
C. Crawley, T. W. Davies, Sue Dolcini, Alexandria Ennik, Stephenie Ferguson,
Toni Gabel, D. Jolly, W. Rausden, Pam Ringhoff, Ann Sibole, Karen Stecker,
R. Stecker, R. Taylor, Ruth Taylor, Joyce Thorp, W. Vaundell, R. L. Wong.

The minutes of the meeting held 19 February 1971 were summarized.

The following names were proposed for membership: David W. Berggren,
Jacques Chassain, J. Eric Cronin, Carol D. Crosswhite, Timothy L. Delgman,
Dennis R. Engel, Carl Goodpasture, Donald J. Jolly, James D. Marshall, Albert
E. Rackett, Keith B. Taylor, Richard J. Taylor, Norman E. Woodley.

Dr. Ferguson announced that Dr. J. W. Tilden, recently emeritus from San
Jose State College, had undergone open heart surgery at the Stanford Medical
Center last week. Dr. Stecker added that blood donations in Dr. Tilden’s name
would be gratefully accepted.

Mr. Leech exhibited some cardboard boxes with pinning bottoms suitable for
temporary storage of Lepidoptera, etc. Twenty-five of these are available from the
Academy at $0.20 each.

The program chairman, Dr. Stecker, announced the forthcoming picnic and
field day is to be held at Del Puerto Canyon, Stanislaus County on Saturday,
15 May. President Blanc had been there earlier today and described the area
as one of an abundance of various plants, notably digger pine, Senecio and
manzanita, with a picnic area and paved roads. Dr. Stecker announced that the
speaker for the next regular meeting on 15 October will be Rick Main, who will
speak on insect collecting in the Grand Canyon of Mexico.

President Blanc called for introductions. Dr. Wasbauer introduced Miss Pam
Ringhoff and Miss Sue Dolcini, both of the California Department of Agriculture.
These girls serve the Society well in that they do all of the typing and record
keeping for the Secretary and so leave him free to concentrate on other duties.

The following notes were presented :

Oviposition of the California Damsel fly, Archilestes californica Mac-
Lachlan. — On 25 September 1968, while collecting insects in the stream bed
of Alameda Creek in Sunol Valley Regional Park, Alameda County, California, I

JANUARY 1972]

PROCEEDINGS

69

observed a pair of Archilestes calif ornica MacLachlan in the process of oviposition
on the small stems of White Alder, Alnus rhombifolia.

The White Alder found along inland foothill and mountain streams in California
is a large upright growing tree. The Alder in this observation was a small (about
V -2 meter in height) immature plant or the leafed sucker stems of a larger tree.
The plant grew out from beneath a large rock and hung over a shallow pool in
the stream.

Closer observation of this plant showed three and sometimes four pairs of
A. californica, the males riding tandem with females in oviposition. Almost all
the small stems of this miniature Alder were girdled by egg punctures.

The male California Damsel fly seizes the female by the prothorax with the
clasping terminal appendages of the abdomen and maintains this posture all
through oviposition. The female arches her long abdomen so that the ovipositor
rests on the bark of the plant stem. She lays six eggs in each puncture that is
made with an incising tool, the terebra of the ovipositor. The punctures are
easily recognizable by the reddish puckered scars on the twigs. The flat elongated
ovate eggs are inserted beneath the outer bark and into the cambium layer in fan-
like fashion three on each side from the puncture. As many as seventy five to one
hundred seventy five eggs may be inserted by one female in numerous incisions
on the twigs. The eggs lie dormant beneath the bark through the winter.
Hatching in spring the young naiads then drop to the water below.

A return trip on 5 October 1968 revealed only one pair of Damsel flies in oviposi-
tion on the same plant. The terminal twigs of the branches of the larger alders
hanging over Alameda Creek were examined at this time but no indications of
scarring by Archilestes could be found. The supposition proposed herein is
that Archilestes californica prefers only the small tender shoots of the immature
alder trees for ovipositing in this area. — T. W. Davies, California Academy of
Sciences , San Francisco.

Sound production in Agrilus pulchellus Bland (Coleoptera: Bupresti-
dae). — At the American Museum’s Southwestern Research Station, 5 miles west
of Portal, in Cochise County, Arizona, on 11 September 1966, while inspecting
an insect flight trap situated in a small meadow, partly surrounded by Salix sp.,
I sighted a specimen of a beautifully colored Agrilus resting on one of the inside
panels of the flight trap. The specimen was collected by hand and held between
my thumb and index finger. To my surprise I could discern stridulatory sounds.
The specimen was held so that the elytra and abdomen were tightly held, with
the head of the beetle free. The stridulatory sounds were produced at the time the
Agrilus rotated its head from side to side in its prothoracic socket. The specimen
was subsequently determined as a female of Agrilus pulchellus Bland.

Carlson and Knight (1969, Contrib. Amer. Entomol. Inst., 4(3) : 69-71) discuss
stridulation in four sympatric species of Agrilus in Michigan. The method of
stridulation is apparently unique to Agrilus. The sounds produced are considered
stress sounds, are identical in both sexes, and are not species specific. With their
Agrilus species the stridulations could not be heard without amplification or by
placing the insect next to the ear. In the case of Agrilus pulchellus the stridulatory
sounds were audible at a distance of at least 15 inches. — Paul H. Arnaud, Jr.,
California Academy of Sciences, San Francisco.

The principal speaker of the evening was to have been Mr. Michael R. Gardner,
but Mr. Gardner was taken ill the day of the meeting and Dr. G. A. H. McClelland

70

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

very kindly offered to fill in by narrating a film he had made on his mosquito
research in Africa.

A social hour was held in the entomology rooms following the meeting.- — M. S.
Wasbauer, Secretary.

Three Hundred and Fortieth Meeting

The 340th meeting was the annual field day and picnic. It was held on Saturday,
15 May 1971 at Frank Raines County Park in Del Puerto Canyon, Stanislaus
County, California.

Members present (19) : F. L. Blanc, R. J. Bushnell, J. E. Cronin, A. R. Dutton,
J. G. Edwards, F. Ennik, W. E. Ferguson, J. R. Gabel, A. Gillogly, Anita Gillogly,
H. B. Leech, C. B. Philip, Harriet Reinhard, R. E. Stecker, R. Taylor, K. Taylor,
M. Wasbauer, Joanne Wasbauer, R. F. Wilkey.

Visitors present (17) : Fern Blanc, Marcia Cronin, Pam Dutton, Alice Edwards,
Sasha Ennik, J. Ennik, A. Ennik, Stephenie Ferguson, Toni Gabel, Gladys Philip,
Karen, Phyllis, Susan and R. Stecker, Jean Walker, Carol and D. Wasbauer.

The day was pleasant, clear and warm. A west wind came up in the afternoon
which kept temperatures down to a pleasant level. Insects were in evidence
throughout the area. Craneflies were particularly abundant in the grass sur-
rounding the irrigated picnic area. Insect collecting and photography were major
activities noted during the day. Frisbee games and sunbathing were also en-
joyed by some. Franklin Ennik reported collecting a number of scorpions in
the canyon and Dr. Edwards collected some interesting aquatic Coleoptera. Dr.
Wasbauer reported that, despite the wind, he took nearly a thousand specimens in
two Malaise traps operated in the Canyon for three hours.- — M. S. Wasbauer,
Secretary.

Three Hundred and Forty-First Meeting

The 341st meeting was held Friday, 15 October 1971, at 7 :45 p.m. in the Mor-
rison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Blanc presiding.

Members present (29) : F. G. Andrews, C. Armin, F. L. Blanc, R. M. Bohart,

I. Boussy, T. Briggs, R. M. Brown, K. S. Corwin, J. Eric Cronin, A. R. Dutton,

W. E. Ferguson, J. R. Gabel, M. R. Gardner, L. Green, J. Guggolz, J. T. Hjelle,
E. A. Kane, H. B. Leech, R. Main, A. E. Michelbacher, C. B. Philip, H. G. Real,
E. S. Ross, R. B. Stecker, K. Taylor, R. Taylor, R. W. Thorp, S. C. Williams.

Visitors present (31) : H. P. Allemndrizen, Fern Blanc, Mr. and Mrs. G.
Cabral, Kathy Cheap, M. P. Cronin, T. Delaman, Thomas Dimock, Cristine Duncan,
Janie Edwards, Beverly Ehreth, Don Emlay, Stephenie Ferguson, P. C. Galam,

T. M. Glimme, Roberto H. Gonzalez, Sue Goodwin, Martha Hjelle, Bob Hishop,

Ron Limandde, Dennis Merrill, Martha Michelbacher, Wayne Nelson, Carl Pinto,
Phyllis Stecker, Don Streke, Joyce Thorp, Marilyn Trochman, Darrell Ubick,
Gregory H. Walker, Wendy Hof man, Robert Wong.

President Blanc announced that Secretary Wasbauer is recovering from a back
operation and is not able as yet to resume his duties as Secretary. President Blanc,
therefore, appointed Michael Gardner acting interim Secretary for the duration of
Dr. Wasbauer’s recuperation.

The minutes of the meeting held 16 April 1971 were summarized. The annual
Field Day and Picnic (340th meeting) was also reviewed.

JANUARY 1972]

PROCEEDINGS

71

The following names were proposed for membership: Student members: John
MacDonald, Carl Goodpasture, Steven Haskett. Regular members: George
Tamaki, C. E. Langston, John D. Glaser, Wayne L. Vaundell, Donald V. Jolly,
Beverly Ehreth, David E. Foster, Fred Roberts, Jim Batts.

Program Chairman Ron Sleeker announced that the next meeting may he held
on the 12 November 1971 and that the principal speaker would be Dr. Henry
Robinson. Dr. Robinson is expected to speak on his work in medical entomology
done for the World Health Organization in Southeast Asia. Dr. Stecker also
announced that the following meeting’s main speaker will be the outgoing President,
Louis Blanc, who will deliver the traditional presidential address.

Dr. Stecker announced that the San Jose Entomology Club had torn 30,000
reprints from excess issues of the Pan-Pacific Entomologist. These were to go on
sale, and the proceeds to benefit both the Pacific Coast Entomological Society
and the San Jose State Entomology Club.

President Blanc expressed his appreciation on behalf of the Society to the
San Jose State Entomology Club for their fine job in making these papers available.

President Blanc announced that the Treasurer, Dr. Arnaud, had received cor-
respondence from the Internal Revenue Service concerning the status of the
Pacific Coast Entomological Society as a non-profit organization. President Blanc
proposed an addition to the Society’s Articles of Incorporation to meet the federal
requirements for a tax-exempt corporation. The addition to the Articles was
thereupon put to the vote of the members and unanimously approved.

President Blanc called for notes and exhibits. The following notes were sub-
mitted :

Some Observations on Tarantula Behavior. — On 27 July 1971, a large,
brown tarantula (Araneae: Theraphosidae) was given to the Department of
Entomology of the California Academy of Sciences. Unfortunately, the origin of
the spider is unknown because it was originally purchased from a biological supply
company. Also, due to the incomplete knowledge of the systematics of this group,
the generic and specific assignment is not known.

On 7 September 1971, the tarantula constructed a whitish egg sac which was
approximately 1.75 in. long, 1 in. wide and % in. thick. The sac was flaccid,
assuming, to some degree, the shape of whatever substrate it was placed on. The
egg sacs of many other spiders, such as the Black Widow ( Latrodectus mactans ) ,
Argiope spp., and the wolf spiders, are strong enough to maintain a circular
shape. It was not tightly packed with eggs, since the contents would shift from
one end to the other when handled. The sac was carried about at times, held
tightly by the chelicerae, and at other times it was left on the ground, seemingly
ignored.

On 1 October 1971, the tarantula destroyed and ate the egg sac and much
of its contents. At this time, I salvaged a portion of the sac and was able to
determine that it consisted of five separate layers of silk, the outermost layer
being thicker than the others. The space inside would have had an approximate
diameter of IV 2 in. if circular.

The eggs were a shimmery yellowish-green with irregular white spots which
moved, as the egg was moved, as if suspended between the “shell” and an inner
central mass. Of the several eggs examined, none showed any trace of embryonic
development.

This spider had been kept as a pet for two years prior to coming to the California

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Academy of Sciences, and during that time never came in contact with a male.
Thus, even though copulation had not occurred, an egg sac was constructed to
accommodate infertile eggs. — John T. Hjelle, California Academy of Sciences,
San Francisco.

High Flight of Butterflies in San Francisco, California. — The purpose of
this note is to report observations made by Mr. Russell W. LaBelle, a stock broker,
who has an office in the Bank of America building in the Business district of
San Francisco. The Bank of America building is San Francisco’s highest; 52
stories and 779 feet in height. It is located in the block surrounded by California,
Kearny, Pine, Montgomery Streets. From his 43rd story window, which is at a
elevation of over 600 feet, on 4 and 5 October 1971, Mr. LaBelle noted butterflies
flying in sufficient numbers to warrant telephoning the Academy to report this
occurrence. During mid-day a butterfly was seen every few minutes. He observed,
from his windows facing on California Street, that the butterflies were flying mostly
from west to east. This direction of flight would direct them inland rather than
towards the coast.

The species involved in this flight is not definitely known. It was not possible
to collect voucher specimens, and an entomologist did not view the flight. It was
at first thought that the Monarch, Danaus plexippus (Lin'naeus) was responsible
for this flight. However, from information on extensive flights of the California
Tortoise Shell, Nymphalis californica (Boisduval) reported in the bay area by
Dr. J. Powell at this same time, this latter species may have been involved in this
flight. — Paul H. Arnaud, Jr., California Academy of Sciences, San Francisco.

Mass movements of Nymphalis californica . — Nymphalis californica under-
goes periodic or sporadic massive population increases and mass movements which
have been called migrations in the literature. The last time this occurred in the
San Francisco Bay area was in 1960, when several aggregations of the butterflies
were observed in spring, followed by tremendous populations developed in June.
During intervening years the species is scarce and may not be a breeding resident
here. Data which will be published elsewhere were given concerning movements
of the butterflies in the Berkeley Hills during 5-14 October 1971. A request was
made for recording of any observations of Nymphalis, particularly mass move-
ments, during this fall and next year. Circumstances indicate that the note given
earlier in this meeting, on high-flying butterflies over San Francisco, involved
observations on N. californica. — J. A. Powell, University of California, Berkeley.

Further Observations of Theraphosid Tarantula Burrows. — Two groups of
colonies of tarantulas are being observed; one is located at Frank Raines County
Park in Stanislaus County, 17 miles west of Patterson. The other is at Los
Gatos Creek County Park, Fresno County, 21 miles west of Coalinga. These
colony sites have been visited once a month since their discovery over a year ago.
Usually the observations have been made during the first weekend of each month.

The Frank Raines Park colonies are on grassy slopes in oak woodlands with
a mixture of digger pines, juniper and chamise and are on the north facing slopes.
At Los Gatos Park, the area is an almost level grassland with oak- juniper wood-
lands having a sparse distribution of digger pine and chamise. Both areas have
been disturbed by man in the making of the parks, but this apparently has had
little effect on the existence of the tarantula colonies.

In late summer and early fall, the tarantula burrow is easy to distinguish because
its occupant has woven a collar of webbing around the edge of the opening and

JANUARY 1972]

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73

usually has a thin veil of webbing across the entrance. If the veil web is destroyed
by inquisitive poking with a twig, the tarantula will usually emerge, examine its
doorway and then descend head-first and spin a new veil web. The constant
respinning of the veil web builds up the collar around the opening. On several
occasions it has been observed that rosettes of dried leaves have been created
around the burrow opening, particularly if the hole emerges on a grassless, hard,
earth-surface of a campground or picnic area. Late in the summer, some of the
tarantulas at Los Gatos Park pushed empty egg cases out of the burrows and
left them beside the entrances. Mature females and males usually have a burrow
entrance about the size of a 250 piece (2.4 cm) . However, when the webbing
collar is removed, the burrow may be as much as 6 to 7.5 cm in diameter.

With the onset of winter, several tarantulas closed their burrows. It has not
been observed how they do this, and there is variation in their techniques. Some
plug their burrows at the surface, while others plug the entrance from 2 cm
below the surface to 20 to 25 cm below the surface. The veil web and collar
are destroyed in the plugging. Sometimes the burrows will develop a growth of
grass around the edges or across the plug. Relocation of the burrows is difficult
unless they are marked.

What the stimulus for reopening the holes may be has not been determined,
nor has the stimulus for plugging the holes been determined. So far there is
little indication of correlation between hole-plugging activities and temperature
drop. Some tarantulas remained active all winter, even when temperatures at
Frank Raines Park dropped to below freezing at night. Others plugged their
holes early in the Fall and remained closed through May. Rain doesn’t seem to be
a direct cause of closure. At Frank Raines Park there was much more rain, yet all
occupied burrows observed at Los Gatos Park, were closed this winter and the
rainfall there has been extremely sparse this year. In the laboratory “colony”
of 13 female and 5 male tarantulas, six females were stimulated to dig holes
after an artificial “rainy season” was created by pouring water on them every
other day for two weeks. In the field, spiders sometimes plug their burrow
entrances after water was poured into the burrow in attempts to wash them out
of the tunnel. One female in the laboratory closed her burrow entrance after a
“rainstorm” where both the ground and the tarantula were well dampened by
water.

The “watering” technique has proved to be a good mea'ns of “sexing” colony
members. Theraphosid tarantulas usually emerge from the burrow when water is
poured into the opening; sometimes half a liter is all that is necessary to bring
emergence and at other times it has been necessary to pour over three to five
liters into the burrow. This technique saves the need to dig out the tarantula
and thus destroy its home. Sometimes, these spiders will plug the entrance with
their bodies when water is introduced — the hairy body forming an impenetrable
plug. This may be a “defensive” mechanism against sudden flash flooding.

It is unwise to poke fingers into open burrows because the tarantula is often
quite agressive when in the burrow. When teasing spiders out of the entrance
with a twig, pencil or similar tool, the tarantula frequently attacks the object with
its chelicerae. It has been observed that the spider can sink its fangs into the wood
of a pencil. However, once the spider has emerged from its burrow, it then usually
assumes the docile attitude of the Theraphosidae. — J. Russel Gabel, San Francisco
State College, San Francisco.

74

THE PAN-PACIFIC ENTOMOLOGIST

[VOL. 48, NO. 1

Death of E. R. Leach. — Mr. Edwin R. Leach of 217 Hillside Avenue, Piedmont,
California, died the 22nd of July 1971, age 93. He was a member of the Pacific
Coast Entomological Society for 55 years, having joined in 1916. He later became
a Life Member, and in 1948 was elected an Honored Member. His interest in the
society was an active one: he helped it with gifts of money, and served as
Treasurer from 1931 to 1942.

Mr. Leach joined the California Academy of Sciences in 1920, became a Life
Member and Patron in 1946, and a Fellow in 1955. Through the kindness of
his wife and daughter, his wish that his collection of beetles be given to the
Academy has been complied with, and the material not yet accessioned but esti-
mated to be between 40,000 and 50,000 specimens, has just been received. His
interest centered in the Cetoninae of the Scarabaeidae, and the Lucanidae, in each
of which he had outstanding world wide collections of identified species. In
addition, there are strong collections of the black tiger beetles, Omus spp., and
the rain beetles, Pleocoma spp., as well as ruteline Scarabaeidae, Australian
Cerambycidae, etc. With the collections there is a fine working library. — Hugh
B. Leech, California Academy of Sciences, San Francisco.

The principal speakers of the evening were Mr. Richard Main and Mr. Herman
Real, of San Jose State College. Their illustrated talk was entitled “5,000 Miles
Through Baja California.”

Coffee and other refreshments were served during a social hour i'n the entomology
rooms following the meeting. — M. R. Gardner, Acting Secretary.

Three Hundred and Forty-Second Meeting

The 342nd meeting was held Friday, 12 November 1971, at 7:45 p.m. in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park,
San Francisco, with President Blanc presiding.

Members present (32) : J. D. Anderson, P. H. Arnaud, Jr., J. W. Bass,
M. Bentzien, R. G. Blair, F. L. Blanc, T. Briggs, R. M. Brown, R.
Bushnell, D. C. Dailey, D. G. Denni'ng, A. R. Dutton, J. G. Edwards, F. Ennik,
W. E. Ferguson, M. R. Gardner, A. R. Gillogly, L. Green, J. T. Hjelle, D. Jolly,
E. A. Kane, R. L. Langston, H. B. Leech, R. Lem, A. E. Michelbacher, C. B.
Philip, F. C. Roberts, H. I. Scudder, R. B. Stecker, K. Taylor, R. Tassan, S. C.
Williams.

Visitors present (25) : G. M. Bass, Fern Blanc, L. Campbell, T. Delgman,
L. Denning, A. Edwards, S. Ferguson, B. Gushkowitz, L. S. Hawkins, D. Jamieson,
P. Jamieson, A. Jung, E. Kemz, A. H. Kreuger, M. A. Kwok, C. LaPoint, L. B.
Mak, M. Michelbacher, R. L. Peterson, R. Poole, H. W. Robinson, R. P. Swaitzell,
L. Tassan, D. Tiemann, R. Zaiate.

The minutes of the previous meeting held 15 October 1971 were summarized.

The following names were proposed for membership: Student members: Albert
Jung, Carolyn Rees, Robert Wong, Steven Haskett; Regular members: Fred C.
Roberts, Lyndon Hawkins, Dean Jamieson, and Raymond L. Peterson.

The following notes were presented:

Contiguous Areas of Arizona and Pacific Slope Floras in Northern Baja
California, Mexico. — Quercus turbinella Green is a common shrubby oak of the
Southwestern U. S. and was described from “'northern Baja California,” but Weld
(1960) in Cynipid Galls of the Southwest indicated he didn’t think Q. turbinella

JANUARY 1972]

PROCEEDINGS

75

occurred in Baja because he had never seen it or it’s distinctive cynipid fauna
there. However, it does occur throughout the Sierra de Juarez of northcentral
Baja California, Mexico. In two papers on the interrelationships between Quercus
dumosa Nutt, (a California oak) and Q. turbinella , Tucker (Madrono, 1952, 11:
234-251; 1953, 12: 49-60) stated that Wiggins thought these two oaks were
isolated in Baja California. During a trip to this area in August 1969, I found
members of the Arizona flora including Q. turbinella, Artemisia tridentata Nutti.,
Pinus quadrifolia Park, and Quercus dunnii Palmer, ( = Quercus palmeri Engelm.)
directly adjacent to representatives of the California flora including Adendostoma
fasciculatum H. & A. and Quercus dumosa. Within five miles southwest of El
Condor, Baja California, (approx. 32° 25' N. Lat., 116° 9' W. Long.), I found
three areas where Q. turbinella and Q. dumosa occur within two feet of each
other. This would be an ideal area to study the environmental conditions restricting
these two floras and associated faunas.

Because of morphological and ecological similarity between Q. turbinella and
an oak in a relict section of the Great Basin flora in the Inner South Coast Range
of California, Tucker described it as Quercus turbinella calif arnica . Although
Munz (1963), A California Flora, (p. 906) and Benson (1967, Amer. J. Bot.,
54(8) : 1017-1026) accept this classification, Jepson (1966, Manual of the

Flowering Plants of California, p. 274) refers to it as a variety of Q. dumosa.

Most oaks in any single subgenus of Quercus with contiguous distributions have
a large percentage of their total gall-making cynipid wasps in common (between
40 and 100% on California oaks). Tucker (1970, Amer. J. Bot., 57(1): 74-84)
and Tucker and Muller, (1958, Evolution, 12: 1-17) have used cynipid host pat-
terns in the analysis of hybrid oak populations. The distinctive faunas of Q.
dumosa and Q. turbinella in northern Baja California suggest host-parasite rela-
tionships might be an additional means of evaluating the taxonomic status of Q. t.
californica. Since Tucker has shown Q. t. californica to have an overlapping
distribution with Q. dumosa (1953, Madrono, 12 : 49-60) these contiguous areas
of distribution could also provide a means of studying the development of these
oak and cynipid patterns and the role of isolation in the development of Pacific
Slope and Southwestern floras and faunas.— D. Charles Dailey, Sierra College,
Rocklin, California.

Mass movement of Sympetrum corruotum (Hagen) (Odonata: Libel-
lulidae) in central California. — The observations of unidirectional mass move-
ments of the dragonfly Sympetrum corruptum (Hagen) have been reported in
the Kensington-Albany-Berkeley area in central California in recent years in notes
by Turner (1965, Pan-Pac. Entomol., 41: 66-67) and Opler (1971, Pan-Pac.
Entomol., 47: 223). In these references the species corruptum was assigned

to the genus Tarnetrum .. Turner suggested that these dragonflies were “probably
migrating from lakes and ponds in Marin County.”

Records of a flight of Sympetrum corruptum in the fall of 1969 in Marin County
are presented. On 11 October 1969 at 17:15 hours, Daylight Saving time, while
traveling north on Dais Road, just south of the intersection of Sequoia Valley Road
and Muir Woods Road (which is in an area west of the city of Mill Valley),
hundreds of dragonflies were observed flying uphill and eastward. It was possible
to look westward, with the sunlight shimmering on their wings, where they could
be seen for a distance of more than a hundred yards — it was a mignificent sight and
flight. At the road level the dragonflies flew from about 4 to 25 feet in height.

76

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

Looking west and downhill some were flying one hundred and more feet above
the dried grassy fields. Rarely a specimen was observed to settle on some vegeta-
tion. The flight was continuing at 17:40 hours when I had to leave. The afternoon
was warm with only the lightest breeze. A continuation of their flight in an easterly
direction would have brought them into the areas where Turner and Opler had
reported their flights in other years.

Sympetrum corruptum is a species with a very wide distribution. It occurs
from Canada to Honduras and is also known from Asia. The single female speci-
men, collected at the time of the observed flight, was confirmed as belonging
to this species by Dr. D. R. Paulson of the University of Washington, in April 1970.
Dr. Paulson reported that he and other Odonata specialists now prefer to include
corruptum in the genus Sympetrum Newma'n: — Paul H. Arnaud, Jr., California
Academy of Sciences, San Francisco.

On the photocopying of entomological literature. — Apart from its legal
and ethical aspects, photocopying is of real significance to our society. We have
kept a stock of back issues of the Pan-Pacific Entomologist in the expectation
of future sales of single copies and complete sets. Copying machines are now
installed not only in research and business centers but also in many stores, and
are inexpensive to use. If students have library volumes of our journal available,
and photocopy the pages of interest, this may well affect the value of our back
issue holdings, and policy on how many extra copies we should order in the future.

Photocopying can be also of great value of our Historical File. We have many
newspaper clippings, some dating back 75 years and more. Newsprint deteriorates
and discolors rather quickly, but bond paper can be used in “Xerox” machines
so it may be highly desirable to copy such articles, and in the future put photo-
copies rather than the original newsprint into the files.

The following clear and pertinent statements are from the journal Philatelic
Literature Review (Vol. 20, No. 3, 30 September 1971), from John Alden’s letter
to the Editor “On the photocopying of Philatelic Literature,” and comments thereon
by the editor, Charles J. Peterson, to whom I an indebted for permission to quote.
Page 133 (Alden) “. . . Readers should perhaps also be reminded that the photo-
copying of copyrighted material, though often ignored in practice, is still illegal.
Although the American library profession has attempted to have incorporated in
American copyright law a ‘fair use’ provision to permit an individual to make such
copies, it has not yet been passed. The economic effect on publishers caused by
the electrostatic copying (by ‘Zerox’ or other comparable processes) have become
increasingly serious, and have led to the devising of paper which will forestall
such copying. Potential . . . clients . . . might bear in mind that in many
countries — particularly the United Kingdom — registration is not required to
secure copyright: publication is in itself sufficient.” Page 134 (Peterson)
“. . . There are broad questions here involving principles of copyright law and
of personal ethics: Is it all right for a non-profit organization to make photocopies,
but not business firms? Can an individual make a copy for himself, but not for a
third party? What about the copying machines which are now being installed
in public libraries as a customer service: isn’t this tactic endorsement of the
practice?” — Hugh B. Leech, California Academy of Sciences, San Francisco.

The Australian sod fly Altermetoponia rubriceps (Macquart) in Marin
County, California. — Altermetoponia rubriceps was first recorded from the New
World by E. L. Kessel (1948, Science, 108(2813) : 607-608) on the basis of speci-

JANUARY 1972]

PROCEEDINGS

77

mens from San Francisco. It proved to be already widespread within the city; in
a mimeographed article, “A study of the Australian sod fly” (1959, Occas. Pap.
No. 1. Student Section, Calif. Acad. Sci., p. I— VII) D. C. Rentz cited its economic
importance in Australia and New Zealand, and gave field notes for San Francisco,
including localities. Part of this appeared in somewhat different form in the
Proceedings of the 272nd Meeting of the Pacific Coast Entomological Society in
the Pan-Pac. Entomol., 37(1): 65. Subsequently it has occurred in numbers at
Berkeley, Contra Costa County, and presumably in adjacent areas on the east
side of San Francisco Bay. It is still very numerous in San Francisco, and in 1971
the spring brood congregated in such numbers on the taller grass stems as to make
black patches on many lawns.

On 9 October 1971, adults were abundant at Black Point, Marin County, at a
parking area and boat ramp under the western end of the bridge by which High-
way 37 crosses the Petaluma River at tidewater. The day was hot and almost
windless; at noon A. rubriceps singletons and some pairs in copulation were
flying in all directions and could easly be picked out of the air by hand. This
locality is in the north western corner of San Pablo Bay, only about 20 miles air-
line north of San Francisco, but Dr. Kessel thinks the species has not been
recorded from Marin before. It is probable that it occurs in adjacent southern
Sonoma County, and in Napa and Solano counties just east of there. — Hugh B.
Leech, California Academy of Sciences, San Francisco.

The evening’s main speaker was Dr. Henry W. Robinson, Professor of
Parasitology, San Jose State College, who spoke on “Epidemiologic Aspects of
Parasitic Infections in Developing Nations — Some Personal Reflections.”

Coffee and other refreshments were served during a social hour in the entomology
rooms following the meeting. — M. R. Gardner, Acting Secretary.

Three Hundred and Forty-Third Meeting

The 343rd meeting was held Friday, 17 December 1971 at 7:45 p.m. in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park,
San Francisco, with President Blanc presiding.

Members present (26) : F. G. Andrews, P. H. Arnaud, Jr., J. W. Bass, R. G.
Blair, F. L. Blanc, J. A. Chemsak, H. V. Davies, D. G. Denning, A. R. Dutton,
F. Ennik, W. E. Ferguson, J. R. Gabel, L. Gormley, E. A. Kane, H. B. Leech, P. A.
McClelland, R. Main, R. L. Peterson, C. B. Philip, J. A. Powell, H. G. Real,
F. C. Roberts, R. E. Stecker, R. W. Thorp, M. S. Wasbauer, S. C. Williams.

Visitors present (11) : Nancy Blair, Anna-Marie Bratton, Kathy Cheap, L.
Hawkins, Sandy Jordan, Pat McClelland, S. Montague, L. Roberts, Mollie Schneider,
R. Swaitzell, Joyce Thorp, Barbara Towbridge, Barbara VanKirk.

The minutes of the meeting held 19 November, 1971 were summarized.

Drs. Stecker and Williams introduced the following guests: Anna-Marie Bratton,
Kathy Cheap, Sandy Jordan, Mollie Schneider and Barbara Trowbridge.

The following names were proposed for membership: Mr. Don Dilley, Mr.
Charles Farrell (student membership) .

The following notes were presented :

Distribution of Enlinia calif ornica (Diptera: Dolichopodidae) in
Oregon. — The genus Enlinia contains some of the smallest species of Dolicho-
podidae. The small size of the species — about a millimeter in total body length —

78

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 1

can account for their infrequent collection. It was in the Proceedings of the
meeting of 20 December 1968 (1969, Pan-Pac. Entomol., 45(1): 79) that I

reported the first occurrence of this genus in the western Nearctic. 1970, Robinson
and Arnaud (1970, Occas. Pap. Calif. Acad. Sci., No. 83: 2-7) described Enlinia
calif ornica from one locality in central California — from a spring in Arroyo Ojo
de Agua, at Redwood City, California. It is now possible to report a major
extension in the distribution of this species by recording it from Oregon. On 1
July 1969, Mr. Kenneth Goeden, of the Oregon State Department of Agriculture,
collected a single male 10 miles north of Imnaha, in Wallowa County, at a waterfall.
Wallowa County is in the north-eastern portion of Oregon, and is partly bordered
by the states of Washington and Idaho. — P. H. Arnaud, Jr., California Academy
of Sciences, San Francisco.

Butterflies at flowers of Escallonia rubra (Saxifragaceae) in central
California. — In October 1971, my wife, Madelaine Milliet-Arnaud mentioned
that she had seen a flowering shrub in the Strybing Arboretum and Botanical
Garden, Golden Gate Park, San Francisco, that was especially attractive to butter-
flies. On 22 October at noon, we visited the area. It was noted that at one large
shrub, which attained a height of about fourteen feet, at least 15 Monarchs,
Danaus plexippus (Linnaeus), nymphalids, pierids, and hesperiids were visiting
its reddish flowers. The plant was identified by Dr. Elizabeth McClintock as
Escallonia rubra (Ruiz and Pavon) Persoon. It is a native of Chile. The genus
Escallonia is widely distributed in the Andes and in eastern South America.
There are several species of Escallonia that are grown in the cool coastal regions
of California, and they are widely planted in Golden Gate Park. Escallonia rubra
can be recommended as a butterfly plant for gardens. — Paul H. Arnaud, Jr.,
California Academy of Sciences, San Francisco.

President Blanc called for reports from the Chairmen of the standing committees.

Mr. Hugh B. Leech, chairman of the Historical Committee stated that the fol-
lowing items were received during 1971 :

From new members of the society, personal data sheets; over the years these
membership records will form one of our most valuable accessions. Through
the kindness of Dr. John Thomas, botanist at Stanford University and at the
California Academy of Sciences, a large number of original pen and ink drawings
of scale insects, lice, etc., by the late Professor G. H. Ferris; there are also a
number of zinc cuts of his figures, and drawings by others, which were used
in the journal Microentomology. In addition there is an old (undated) student
notebook containing the late L. H. Knoche’s notes on Professor Vernon Kellogg’s
lectures in entomology; Mr. Knoche was a botanist who left a very large collection
of plants to Stanford. Through the kindness of Mrs. J. Birchim, we received a
file of correspondence and notes from the effects of the late Jim Birchim; they
relate chiefly to his interests in Diptera and Orthoptera. Also added are a few
photographs of entomologists, gift of Dr. E. S. Ross, and a miscellany of newspaper
clippings and small items.

Dr. P. H. Arnaud, Jr., read the treasurer’s financial report and Mr. H. Vannoy
Davis, chairman of the auditing committee reported that the financial records of
the Society for 1971 are in good order.

President Blanc announced his appointment to the publication committee for
1972 through 1974 of Dr. J. A. Powell and Mr. M. R. Gardner.

Mr. M. R. Gardner, chairman of the nominating committee presented a slate

JANUARY 1972]

PROCEEDINGS

79

of candidates for offices in the Society for 1972: Dr. D. G. Denning, President;
Dr. J. A. Chemsak, President-Elect; Dr. M. S. Wasbauer, Secretary; Dr. P. H.
Arnaud, Jr., Treasurer. There were no nominations from the floor. The nominees
were unanimously elected to office for 1972.

The principal speaker of the evening was the outgoing president of the Society,
Mr. F. L. Blanc who presented his presidential address entitled “Human in-
fluences on endemic and exotic insect distributions in California.”

A social hour was held in the entomology rooms following the meeting.: — M. S.
Wasbauer, Secretary.

ENTOMOLOGICAL EXCHANGES AND SALES

Space is available to members and non-members who wish to advertise entomo-
logical exchanges, sales, announcements, etc. We can offer an ad in four issues
for $1.50 per line. Ads will be arranged in order of their receipt. Contact the
Advertising Manager, W. H. Lange, Department of Entomology, University of Cali-
fornia, Davis, California 95616.

Lepidopterist needed to describe new species in genus Comadia (Cossidae). All
stages, life history, and preliminary manuscript available; genus needs revision.
R. P. Allen, California Department of Agriculture, Sacramento 95814.

Need 2x2 color slides of economic insects and damage other than California;
borrow and loan for duplicating; exchange duplicates. R. P. Allen, Calif. Dept, of
Agriculture, Sacramento 95814.

SHARP & MUIR: THE COMPARATIVE ANATOMY OF THE MALE GEN-
ITAL TUBE IN COLEOPTERA. The classic 1912 monograph and six other papers
by the same authors; 304 pp., 43 pis., bound. $10.00. An essential book for all
coleopterists. Entomological Society of America, 4603 Calvert Road, College Park,
Maryland 20740.

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pointed in ADVANCES IN SUGARBEET PRODUCTION: PRINCIPLES AND
PRACTICES, just published by the Iowa State University Press. The control of
insects, plant diseases, nematodes and weeds are discussed in addition to other
phases of production. Available at $12.50 from bookstores or from the Iowa
State University Press.

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APRIL 1972

Vol. 48

No. 2

THE

Pan-Pacific Entomologist

HUCKETT — Notes on Bigot’s North American type-specimens at the Uni-
versity Museum, Oxford (Diptera: Anthomyiidae, Muscidae) 81

CHEMSAK — Review of the genus Cirrhicera Thomson (Coleoptera: Cerambyc-

idae) 86

LINSLEY — The robber fly Callinicus calcaneus (Loew) as a predator on
Andrena omninigra Viereck (Diptera: Asilidae; Hymenoptera:

Andrenidae) 94

JVtOSER AND ROTON — Reproductive compatibility between two widely sepa-
rated populations of Py emotes scolyti (Acarina: Pyemotidae) 97

PARKIN, PARKIN, EWING, AND FORD— A report of the arthropods col-

lected by the Edinburgh University Galapagos Islands Expedition, 1968

100

SPILMAN — A new genus of jumping shore beetle from Mexico (Coleoptera:

Limnichidae) 108

GILLOGLY— A new species of My strops from Costa Rica (Coleoptera:

Nitidulidae) 116

BROTHERS — A new species of Thaumalea from California (Diptera: Thau-

maleidae) , 121

COHEN AND ALLEN — New species of Baetodes from Mexico and Central

America (Ephemeroptera: Baetidae) 123

EDMUNDS AND KOSS — A review of the Acanthametropodinae with a

description of a new genus (Ephemeroptera: Siphlonuridae) 136

BOHART AND GRISSELL — Nesting habits and larva of Pulverro monticola

(Hymenoptera: Sphecidae) 145

CHEMSAK — A new seed inhabiting cerambycid from Costa Rica (Coleoptera)

150

BOOK NOTICES 85, 96, 120

NOTICE 107

SOCIETY ANNOUNCEMENT— Usinger Autobiography 135

SCIENTIFIC NOTES 144, 149

SAN FRANCISCO, CALIFORNIA • 1972

i

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

THE PAN-PACIFIC ENTOMOLOGIST

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The Pan-Pacific Entomologist

Vol. 48

April 1972

No. 2

Notes on Bigot’s North American type-specimens
at the University Museum, Oxford

(Diptera: Anthomyiidae, Muscidae)

H. C. Huckett
Riverhead, New York 11901

Bigot (1885, 1886, 1887) in his descriptive work on new or little
known species of Diptera included several that were recorded from
North America and that occur in the neighboring region of Mexico.
This material, with its type-specimens, has been in the possession of the
Verrall Collection at Newmarket, at which time it was reviewed by Stein
(1907) along with other exotic taxa. Recently that part of the Bigot
material containing anthomyiid and muscid forms, with the exception
of three nominal species ( vide infra), has been donated by the late Mr.
J. E. Collin, in whose custody the Verrall Collection had remained, to
the Elope Department of Entomology at the University Museum, Oxford,
along with his own extensive collection of British Diptera.

Through the curtesy of Professor G. C. Varley and the staff at the
Hope Department of Entomology I was given access to the collections
for the purpose of studying the types of Bigot’s North American species
belonging to the families Anthomyiidae and Muscidae, and was provided
with the necessary facilities for their examination. To Mr. D. M.
Ackland I am particularly indebted for assistance in guiding me to the
various sections of the collection related to my current work, and for
drawing my attention to the questionable status of Nemopoda obscuri-
pennis Bigot, classed with the Acalyptratae.

The types of Bigot’s species have been transferred to separate trays
headed by the large original dark-edged label, on which the specific
name is written in a large formalized hand. Beneath the name and to
the left is written in smaller lettering the locality, e. g. “Am. Sept. ( Mt .
Roche ),” and to the right “]. Bigot.”

The type-specimens, with few exceptions as duly noted, possess a label
attached to their pins, on which the name of the species is repeated in

The Pan-Pacific Entomologist 48: 81-85. April 1972

82

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

ink in Collin’s handwriting, followed by “EX COLL. BIG.” in large
print.

I have noted various discrepancies in the labelling of species and with
earlier records concerning the spelling of names, the sex cited, and in
the number of specimens present, for sake of avoiding ambiguity or
error.

The Bigot names are listed in alphabetical order accompanied by a
reference to their publication date and page.

cmthracina, Anthomyia. (1885, p. 298). 2$, 42, from the Rocky
Mountains, all as treated by Ackland (1968) and belonging to the type
series, of which one female remains undetermined. The named speci-
mens are identical with Calythea separata Malloch.

anthrax, Limnophora. (1885, p. 274). 1$, from Mexico. The male
is notable for the brown calyptrae. The species belongs to the genus
Spilogona Schnabl.

argentina, Ophyra. (1885, p. 302). 12, from Buenos Ayres, agrees
in all particulars with females of Ophyra aenescens (Wiedemann), as
has been corroborated by studies of Oliveira (1941) .

calopus, H.l . (1885, p. 275) . 12, from Mexico, and named in writing
H. calopoda on Bigot’s type label, to which is added “Type” in Collin’s
handwriting. Stein (1907, p. 215) gives the genus as Hydrophoria , in
compliance with the context. The specimen is conspecific with Bitho-
racochaeta leucoprocta (Wiedemann) .

dentata, Homalomyia. (1885, p. 284). 1 $, from the Rocky Moun-
tains, agreeing in all respects with males of Limnophora narona
(Walker).

flavicaudata, Hylemyia. (1885, p. 299). 1$, from Washington Ter-
ritory, having the name H. flavicauda written on Bigot’s type label. The
specimen in conspecific with Hylemya alcathoe (Walker) .

fulviventris, Spilogaster. (1885, p. 291). IS, from California, a
variant of the species Helina troene (Walker), having the thorax gray,
with humerals and scutellum partly reddish tinged, abdomen mainly
fulvous and with two pairs of weak spots.

fulvus, Spilogaster. (1885, p. 289). 1 S, from Washington Territory,
having abdomen, one hind leg and much of one wing missing. The
type specimen has head, thorax and legs fulvous, thus markedly differing
from associated species within Hebecnema Schnabl.

mexicana, Homalomyia. (1885, p. 284). 3S, from Mexico, all con-
specific, and may be recognized readily as belonging to the species
Fannia canicularis (Linnaeus).

monticola, Anthomyia. (1885, p. 297). 6$, 2 2, from the Rocky

APRIL 1972] HUCKETT BIGOT’S N. A. TYPE-SPECIMENS

83

Mountains, all as treated by Ackland (1968) and belonging to the type
series, of which one female is recognized as Delia platura (Meigen) .
As noted by Ackland, the species monticola and anthracina of Bigot are
conspecific, hence the name monticola, due to claims of page precedence,
may be accepted as the valid name for the species. Both taxa are identical
with Calythea separata Malloch.

nigricauda, Hydrophoria. (1885, p. 276) . 2 c?, from the Rocky Moun-
tains, one of which has lost its antennae. Both specimens have basal
segments of abdomen reddish testaceous, and the tubular haustellum
together with the slender labellum one and a half times as long as height
of head, thereby differing from its congeners Neohylemyia tenuirostris
(van der Wulp) and N. mallochi Huckett, that have a shorter haustellum
and a uniformly gray abdomen. The calyptrae and wings of the type
specimens of N. nigricauda possess a brownish tinge. The species be-
longs to the genus Neohylemyia Malloch.

obscuripennis , Nemopoda. (1886, p. 392) . 12, from California, with
the name N. obcuripennis written on Bigot’s type label, and the sex cited
as a male in the description. The type specimen is in poor condition,
owing chiefly to the covering of much of the abdomen in a pinkish mold.
The remaining parts except the forelegs that are missing can with due
care be perceptible, and indicate clearly that the specimen is a female
and is conspecific with Chelisia elegans Stein, N. SYN.

oculijera, Anthomyia. (1885, p. 299). 1$, from Baltimore. The
dark brown marks on the whitish mesonotum are striking and may be
described in some detail. The longish oblong presutural marks becloud
the planes of dorsocentral bristles, extending to anterior border of
mesonotum and ending caudad shortly before reaching the transverse
suture: on postsutural region a band or fascia adjoining the transverse
suture that medially encroaches as a slight wedge between the caudal
pair of presutural acrosticals, laterad the fascia extends to prealar
bristles, and caudad to the third pair of postsutural dorsocentrals, from
thence as a tongue to base of wings. Scutellum entirely black on dorsum.
A chitinous fold or lobe extends distinctly mesad beyond apex of each
process on fifth abdominal sternum.

rubifrons, Anthomyia. (1885, p. 297). 3 2, from Chile, with the
name written A. rubrifrons on Bigot’s type label. The specimens closely
resemble the female of Delia platura (Meigen) , from which I disassociate
them owing to the presence of two anterior sternopleural bristles.
Collin (in letter, 28 March 1951) has indicated that these specimens are
in all probability females of the male A. sancti-jacobi described by Bigot
on the previous page (1885, p. 296), and that all had arrived in the

84

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

same lot from Chile. Stein (1907, p. 285) denoted the species under the
name Chortophila cilicrura Rondani.

rupecula, Homalomyia. (1885, p. 285). 2$, from the Rocky Moun-
tains, agreeing in all particulars with the male of Delia platura
(Meigen) . The name is written H. rupicola on Bigot’s type label.

siphonina, Proboscidomyia. (1885, p. 267). 2$, from the Rocky
Mountains, that are without the usual Collin label attached to pines of
type specimens. The males possess an extremely long slender tubular
proboscis and minutely haired arista, otherwise, in habitus, resembling
the species of Neohylemyia Malloch.

spinipes, Chortophila. (1885, p. 279). 12, from the Rocky Moun-
tains, agreeing closely with the female of Delia platura (Meigen) , and
with which I regard it to be conspecific. The description of the species
accords the sex to be a male.

To Mr. A. C. Pont of the British Museum I am indebted for informa-
tion concerning three nominal species from the Rocky Mountains that
Bigot (1887) described in a later contribution, the types of which are
deposited in the British Museum (Natural History). Two of these taxa
have recently been commented on by Pont (1970) in an article dealing
with the European species of Myospila Rondani.

anthomydea, Curtonevra. (1887, p. 614). 1$, with midlegs missing,
is conspecific with Myospila meditabunda (Fabricius) of authors.

nigriceps, Curtonevra. (1887, p. 615) . 3 S , the antennae and midlegs
lacking in one specimen, one mid and one hind leg missing in the two
others, all conspecific and agreeing with Myospila meditabunda
(Fabricius) of authors.

flavipennis , Musca. (1887, p. 605) . 1$, 12, both in poor condition,
but clearly recognizable as belonging to Musca domestica Linnaeus. The
female is without a locality label.

Literature Cited

Ackland, D. M. 1968. The world species of Calythea Schnabl. and Dzied.

(Dipt., Anthomyiidae) with notes on Bigot’s types. Entomol. Mon. Mag.,
104: 135-144, 23 figs.

Bigot, J. M. F. 1885. Dipteres nouveaux ou peu connus. 25 e partie, XXXIII:

Anthomyzides nouvelles. Ann. Soc. Entomol. Fr., (1884) ser. 6 4:
263-304.

1886. Dipteres nouveaux ou peu connus. 29 e partie (suite), XXXVII: 2 e .
Essai d’une classification synoptique du groupe des Tanypezidi ( mi ld)
et description de genres et d’especes inedite. Ann. Soc. Entomol. Fr.,
ser. 6 6: 369-392.

1887. Dipteres nouveaux ou peu connus. Leptidi, Muscidi. Bull. Soc. Zool.
Fr., 12: 581-617.

APRIL 1972] HUCKETT— bigot’s N. A. TYPE-SPECIMENS

85

Oliveira, S. J. de. 1941. Sobre Ophyra aenescens (Wied.). Arq. Zool. Sao Paulo,
2, Art. 15 pp. 341-355, 3 pis.

Pont, A. C. 1970. Myospila hennigi Gregor and Povolny, 1959 (Dipt., Muscidae) ,
new to Britain and notes on the European species of Myospila Rondani.
1856. Entomol. Mon. Mag., 106: 111-113.

Stein, P. 1907. Revision der Bigot’schen und einiger von Macquart beschriebenen
aussereuropaischen Anthomyiden. Z. Syst. Hymenopt. Dipt., 7 heft 3
pp. 209-217 ; ibid heft 4 pp. 273-293.

BOOK NOTICE

Catalogue of the Thysanoptera of the World (Part 1). By C. F. Jacot-
Guillarmod. 20 February 1970. Annals of the Cape Provincial Museums,
Natural History, vol. 7, pt. 1. Pp. iii -|- 216 (numbered pages 217 and 218
are blank pages). Soft cover. R5.00 (about US$ 7.00).

In November 1970 Dr. Howell V. Daly, Jr., mailed from Grahamstown, a copy
of Mr. Jacot-Guillarmod’s catalog for review in the Pan-Pacific Entomologist.
It is a pleasure to do so, for this is an important work that is well done. This

first volume of a projected monumental work of possibly six or more parts when

completed, is dedicated to Dr. J. C. Faure, under whose guidance it was
started some 30 years ago. It begins (pp. i-iii) with a brief introduction
including “Scope and Method.” The general classification followed is that of
Priesner with slight modification. Part one covers the suborder Terebrantia
families Aeolothripidae, Mesothripidae, Paleothripidae, Permothripidae, Mero-
thripidae, Heterothripidae, and Uzelothripidae. Fifty genera (24 are monotypic)
and 339 valid species are cataloged. Both fossil and recent species are included.
For each valid name, the original description and all subsequent publications

are cited; for each genus the type species is cited; for each species the

location of type, distribution, type locality, and habitat are cited; invalid names
are cross-indexed. This publication, in this economically important order, should
have an extensive use. — Paul H. Arnaud, Jr., California Academy of Sciences,
San Francisco, 94118.

86

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Review of the Genus Cirrhicera Thomson

(Coleoptera: Cerambycidae)

John A. Chemsak

University of California, Berkeley 94720

The hemilophine genus Cirrhicera Thomson is comprised of a number
of rather pretty, moderate-sized species occurring from about central
Mexico to Panama. Thomson (1857) included three Mexican species
(one erroneously cited as “Patrie: Venezuela”) and another in 1860
from the same country. Bates (1881) added four species from Mexico
and Guatemala and one from Panama (1885). The remaining two
known species were described from Mexico by Gahan (1892).

Specimens of Cirrhicera are rather rare in collections and there are
at hand only 92 specimens representing eight species. The types or
determined material of all of the unrepresented species have been ex-
amined at the British Museum (Natural History) .

These studies were undertaken during a National Science Founda-
tion sponsored study on North American Cerambycidae (Grant GB-
4944X) . The following institutions and individuals are gratefully
acknowledged for making material available: American Museum of
Natural History; British Museum (Natural History) ; California Acad-
emy of Sciences; J. M. Campbell; Canadian National Collection;
Cornell University; Field Museum of Natural History, Chicago; Los
Angeles County Museum; Museum of Comparative Zoology, Harvard;
Ohio State University; United States National Museum; University of
California, Davis; and University of Kansas. Special thanks are given
to Celeste Green for preparing the illustrations.

Genus Cirrhicera Thomson

Cirrhicera Thomson, 1857, Arch, entomol. 1: 309; Thomson, 1860, Class, de

cerambycides, p. 63; Thomson, 1864, Systema cerambycidarum, p. 128;
Lacordaire, 1872, Genera des coleopteres, 9(2) : 892; Bates, 1881, Biologia

Centrali-Americana, Coleoptera, 5: 213.

This genus is characterized by the tufts of black hairs on the fifth
and sixth antennal segments. The third antennal segment is longer
than the first and the fourth shorter than the third but longer than the
first. The distal segments are short. The eyes are finely faceted and
almost separated, the two lobes connected by a single row of facets. The
pronotum is cylindrical, transverse and not laterally produced. The

The Pan-Pacific Entomologist 48: 86-93. April 1972

APRIL 1972]

CHEMSAK — REVIEW OF CIRRHICERA

87

prosternal process is narrow, arcuate and expanded apically. The elytra
have strong costae extending from the humeri making the epipleurae
almost vertical. The legs are slender, tibiae lacking a sinus, and the
tarsal claws are bifid.

Type species. — Hemilophus leuconotus Laporte (Thomson designa-
tion, 1864) .

Eleven species are presently known.

Key to Species of Cirrhicera

1. Elytra without pale pubescent spots, pale pubescent patches either

covering most of surface, transverse, or absent 2

Elytra with pale, pubescent spots usually at base and behind middle 5

2(1). Elytra with pubescent fasciae covering most of surface or basally and

medially transverse 3

Elytra uniformly densely pubescent with black spots at middle and at
apical one-third; pronotum densely grayish pubescent except for nar-
row, median, longitudinal line. Length, 9-13 mm. Mexico to El

Salvador cinereola Bates

3(2). Elytra with pubescent fasciae extending over most of surface; pronotum

with lateral pubescence same color as that of elytra 4

Elytra black with broad yellow pubescent bands transverse, one basal,
one post-median; pronotum orange pubescent laterally. Length, 15-16

mm. Mexico conspicua Gahan

4(3). Abdomen with last sternite emarginate at apex; antennae usually dark.

Length, 9-17 mm. Mexico leuconota (Laporte)

Abdomen with last sternite rounded at apex; antennae testaceous.

Length, 10-12 mm. Costa Rica to Panama panamensis Bates

5(1). Elytra without two elevated tubercles at base 6

Elytra with two strongly elevated tubercles at base; antennae with scape
densely fringed with long pubescence beneath. Length, 8.5 mm

Mexico and Guatemala cristipennis Bates

6(5). Elytra with two pairs of pubescent spots, one basal and one post-median ._ 7
Elytra with two pubescent spots behind the middle only; color black.

Length, 9 mm. Mexico nigrina Thomson

7(6). Elytra with basal pubescent spots sutural, lying behind scutellum 8

Elytra with basal pubescent spots well separated, lying on basal margin.

Length, 8-12 mm. Nayarit to Veracruz, Mexico basalts Gahan

8(7). Elytra with pubescent spots not circled with black 9

Elytra with pubescent spots circled with black, apices deeply emarginate,
margins acute. Length, 7-11 mm. Southern Mexico and Guatemala ....

championi Bates

9(8). Pubescent spots yellow; long erect hairs absent or sparse on pronotum

and elytra 10

Pubescent spots white; long erect hairs numerous on pronotum and
elytra. Length, 7.5 mm. Veracruz, Mexico niveosignata Thomson

10(9). Abdomen without pubescent spots; antennae with basal segments dark,

88

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48 , NO. 2

fifth and sixth segments black. Length, 10-11 mm. Guatemala

longifrons Bates

Abdomen with yellow pubescent spots; antennae pale throughout, seg-
ments five and six brownish. Length, 6.5-13 mm. Mexico to Panama _
sallei Thomson

ClRRHICERA CINEREOLA Bates
(Fig. 1)

Cirrhicera cinereola Bates, 1881, 5: 215.

The body entirely clothed with grayish to ochraceous appressed
pubescence. The elytra have four dark spots at the apical one-half.
Type locality. — Guatemala, near the city.

Additional records. — 1 $ , 1 $ , Chiapas, Mexico, Pacific slope Cordellerus,
800-1,000 M (L. Hotzen T9) ; 1 $ , San Jeronimo, Volcan Tacana, Chiapas, Mexico, 9
September 1970 (E. C. Welling) ; Id , Mexico (F. C. Bowditch) ; Id , 1$, Antigua,
Guatemala, October 1965 (N. L. H. Krauss) ; Id, Chicacoa Such., Guatemala,
2,000 ft., 22 June 1965 (J. M. Campbell); Id, Ahuachapan, El Salvador, 18
August 1960.

Cirrhicera conspicua Gahan
(Fig. 2)

Cirrhicera conspicua Gahan, 1892, 1892: 269, pi. 12, fig. 13.

This species is very distinctive by the orangish lateral bands of the
pronotum and by the yellow transverse bands of the elytra, one basal
and one post-median.

Type locality. — Guerrero, Mexico.

Known only from the type series (male, female) .

Cirrhicera leuconota (Laporte)

(Fig. 3)

Hemilophus leuconotus Laporte, 1840, 2: 489.

Cirrhicera leucronota, Thomson, 1857, 1: 309.

Cirrhicera leuconota, Thomson, 1860, p. 64; Thomson, 1864, p. 128; Bates, 1881,
5: 213.

Ground color brown to dark brown with the elytra extensively clothed
by pale pubescence. The median dark band of the pronotum is broad
and the antennae dark. The abdomen has the last sternite emarginate
apically in both sexes. The extent of the pale patch of the elytra is some-
what variable and usually emarginate toward the suture at the middle.
Type locality. — Mexico.

Material examined.- — Mexico: 4dd, 12, Orizaba, Veracruz, 12-22 August
1961 (R. & K. Dreisbach) ; 4<d d, Fortin de las Flores, Veracruz, 10 June 1959
(H. E. Evans), 6 July 1963 (W. A. Foster) ; 62 9, Cordoba, Veracruz, 8 July

APRIL 1972] CHEMSAK REVIEW OF CIRRHICERA 89

Fig. 1. Cirrhicera cinereola Bates. Fig. 2. C. conspicua Gahan. Fig. 3. C.
leuconota (Laporte) . Fig. 4. C. panamensis Bates.

1961 (D. H. Janzen), 8 October 1941 (DeLong, Good, Caldwell & Plummer), 23
July 1936 (C. H. Seevers) ; 1 2 , El Fortin, Veracruz, 8 July 1941 (H. S. Dybas) ;
12,2 miles NE San Andreas Tuxtla, Lago Encantada, 460 M, Veracruz (B. & B.
Valentine) ; 1 $ , 5 miles W Orizaba, Veracruz, 7 July 1962 (J. M. Campbell) ; 12,
Mexico (Deyer) ; 12, Temescal, Oaxaca, 11, 12 September 1964.

Cirrhicera panamensis Bates
(Fig. 4)

Cirrhicera sallaei var. panamensis Bates, 1885, 5: 429; Chemsak and Linsley, 1970,
43 : 409 (lecto type design.) .

In coloration this species is quite similar to C. leuconota. In addition
to the pale antennae, C. panamensis also has the apex of the last ab-
dominal sternite rounded.

Type locality. — Volcan de Chiriqui, Panama.

Material examined. — 1<5, 6 Km S San Vito, Puntarenas, Costa Rica,
2 May 1967 (D. F. Viers) .

Cirrhicera cristipennis Bates
(Fig. 5)

Cirrhicera cristipennis Bates, 1881, 5 : 214.

This species is easily recognizable by the two prominent tubercles near
the base of the elytra and the highly elevated pronotal disk. The antennae
are densely fringed on the scape and segments three and four also have

90

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Fig. 5. C. cristipennis Bates. Fig. 6. C. nigrina Thomson. Fig. 7. C. basalis
Gahan. Fig. 8. C. championi Bates.

a small fringe. The elytra are finely clothed by very short pubscence
having a lavender cast.

Type locality. — Mexico.

Material examined. — 1$, Orizaba, Veracruz, Mexico, 12-22 August,
1961 (R. & K. Dreisbach) ; IS, Morales, Guatemala, 8 March 1905.

Cirrhicera NIGRINA Thomson
(Fig. 6)

Cirrhicera nigrina Thomson, 1857, 1: 310; Bates, 1881, 5: 214.

The characterization of this species is based on a specimen deter-
mined by Bates in the Salle Collection. The ground color is black with
two rounded pale spots a little behind the middle of the elytra.

Type locality. — Mexico (erroneously cited as Venezuela by
Thomson) .

Cirrhicera basalis Gahan
(Fig. 7)

Cirrhicera basalis Gahan, 1892, 1892 : 269, pi. 12, fig. 5.

The color is pale to dark brown with the pubescent patches of the
pronotum and elytra white. The median dark area of the pronotum
is broad apically, converging toward the base. The basal white patches

APRIL 1972]

CHEMSAK — REVIEW OF CIRRHICERA

91

are placed on each side of the scutellum while the post-median pair is
sutural and usually posteriorly oblique.

The two separated basal spots will separate this species from C. sallei.

Type locality. — Guerrero, Mexico.

Material examined. — Mexico: 12, 6 miles E San Bias, Nayarit, 27 August
1959 (A. S. Menke, L. A. Stange) ; 1 $ , San Bias, 7 August 1964 (W. R. M.
Mason); 12, 18 miles N Tepic, Nayarit, 16 August 1960 (D. C. Rentz) ; 1 <$ ,
Tepic, 13 September 1951 (R. & K. Dreisbach) ; 1 $ , 4 miles W Tepic, 31
August 1961; 12, Barra de Navidad, Jalisco, September 1965 (N. L. H. Krauss) ;
12, Km 108 on Hwy 110 to Manzanillo, Jalisco, 20 July 1956 (R. E. Beer and
party); 1$, Puerto Vallarta, Jalisco, 6 July 1957 (J. A. Comstock); 2 8 $,
Acapulco, Guerrero, 1 August 1934 (C. C. Plummer), 4 October 1945; 1$,
Temascal tepee, D. F., 1931 (G. B. Hinton); 1$, Cordoba, Veracruz, 4 August
1965 (A. B. Lau) .

Cirrhicera championi Bates
(Fig. 8)

Cirrhicera championi Bates, 1881, 5: 214, pi. 15, fig. 12; Chemsak and Linsley,
1970,43: 409 (lecto type design.) .

Very similar to C. sallei but having the yellowish elytral spots
separated but contiguous and outlined in black or dark brown. The
appendages are pale except for segments one, five and six of the antennae.
Type locality. — Zapote, Guatemala.

Material examined. — 4$ 8, San Jeronimo, Volcan Tacana, Chiapas, Mexico,
12, 21 August 1970, 2 September 1970, 10 October 1970 (E. C. Welling); 12,
Moca, Such., Guatemala, 3,000 ft., 16 July 1959 (P. & C. Vaurie) ; 1 2, 15 Km S La
Reforma, S. M., Guatemala, 3 September 1965 (J. M. Campbell) ; 1 2 , 1 Km N
Yecocapa, Chi., Guatemala, 1,500 M, 29 May 1966 (Campbell); 1$, 4 Km N
Palin, Esc., Guatemala, 4,500 ft., 21 June 1966 (Campbell); 12, Finca Moca,
Santa Barbara, Such., Guatemala, 3,000 ft., 12 June 1966 (Campbell).

Cirrhicera niveosignata Thomson
(Fig. 9)

Cirrhicera niveosignata Thomson, 1860, p. 64; Bates, 1881, 5: 214.

This species may be recognized by the small white, almost scale-like
pubescent patches. The basal pair is separated and behind the scutellum
and the post-median pair is contiguous at the suture. The body is
moderately clothed with long erect hairs and the short appressed
pubescence is relatively sparse.

Type locality. — Mexico.

Material examined. — 12 , Tezonapa, Veracruz, Mexico, 8 August
1941 (H. S. Dybas).

92

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Fig. 9. C. niveosignata Thomson. Fig. 10. C. longifrons Bates. Fig. 11. C.
sallei Thomson.

ClRRHICERA LONGIFRONS Bates
(Fig. 10)

Cirrhicera longifrons Bates, 1881, 5: 214; Chemsak and Linsley, 1970, 43. 409
(lectotype design.).

This species is known only from the originally described male and
female. It greatly resembles C. sallei but the two apparently differ by
the lack of pubescent patches on the abdomen and dark basal antennal
segments of C. longifrons. These two species may ultimately prove to
be the same.

Type locality. — Purula, Guatemala.

Cirrhicera sallei Thomson
(Fig. 11)

Cirrhicera sallei Thomson, 1857, 1: 310.

Cirrhicera sallaei, Bates, 1872, 1872: 231; Bates, 1881, 5: 214.

This appears to be the most common and widespread species of
Cirrhicera. The subbasal yellowish patches are irregular and contiguous

APRIL 1972]

CHEMSAIC REVIEW OF CIRRHICERA

93

at the suture. The postmedian pair are variable in size and shape but
also meet at the suture.

Type locality. — Mexico.

Material examined. — Mexico: 1$, Cotaxtla, Veracruz, 26 August 1959 (R. F.
Smith) ; 1 $ , Fortin de las Flores, Veracruz, 6 August 1963 (W. A. Foster) ; 1 d ,
Cordoba, Veracruz (A. Fenyes) ; 1$, Coyame, Lake Catemaco, Veracruz, 10-18
July 1963 (D. R. Whitehead); 1$, Comoapan Falls, Veracruz, 14 July 1963
(Whitehead); 1 $ , Teapa, Tabasco, 11 July 1964 (E. Fisher); 1$, X-can,
Quintana, Roo, 27 July 1967 (E. C. Welling); 1?, 72 miles E La Ventosa,
Oaxaca, 21 July 1963 (J. T. Doyen); Id, 12, Cuernavaca, Morelos, July 1945
(N. L. H. Krauss) . Guatemala: Id, Matias de Galvez, 14-15 August 1965 (P. J.
Spangler) ; Id, Dept. El Progreso, Virgen, 11-12 August 1965 (Flint and Ortiz) .
British Honduras: 12, Belize (Peck) ; 12, Augustine Mt., Pine Ridge, 3-7 July
1963 (C. C. Porter) ; 2d d, M-tee District, 12 June 1906. El Salvador: Id, 12,
6 miles W Quezaltepeque, 12 August 1963 (D. Q. Cavagnaro, M. E. Irwin) ; 12,
San Salvador, 19 September 1960; 12, La Libertad, November 1959 (Krauss);
1 2, Metepan, 5 August 1954. Honduras: 1 2, Zamorano, September 1953 (N. L. H.
Krauss). Costa Rica: 12,4 Km N Canas, Guanacasta, 15 July 1965 (D. Viers) ;
12, Boca de Barranca, Puntarenas, 19 June 1963 (C. L. Hogue); Id, 12,
Piedras Negras; 2d d, 12, Turrialba, 21, 29 May 1951 (0. L. Cartwright), 28
May 1962 (H. Ruckes) ; Id, Barranca, Puntarenas, 24 July 1929 (F. Nevermann) ;
12, Hamburgfarm, Reventazon, 25 March 1935 (Nevermann); 12, Costa Rica.
Panama: 2d d, Sumit, Canal Zone, June, July 1953 (Krauss) ; 12, Porto Bello,
18 February 1912 (A. Busck) .

Literature Cited

Bates, H. W. 1872. On the longicorn Coleoptera of Chontales, Nicaragua.
Trans. Roy. Entomol. Soc. London, 1872: 163-238.

1879-1886. Biologia Centrali-Americana, Coleoptera, Longicornia, 5: 1-436.

Chemsak, J. A. and E. G. Linsley. 1970. Additional designations of lectotypes
of Neotropical Cerambycidae in the collections of the British Museum
(Natural History). J. Kans. Entomol. Soc., 43: 404-417.

Gahan, C. J. 1892. Additions to the Longicornia of Mexico and Central America,
with notes on some previously recorded species. Trans. Roy. Entomol.
Soc. London, 1892: 255-274.

Lacordaire, T. 1872. Genera des coleopteres. (Paris). 9(2): 411-930.

deLaporte, F. L. N. de C. 1840. Histoire naturelle des animaux articules, vol. 2,
564 pp. Paris.

Thomson, J. 1857. Description de cerambycides nouveaux ou peu connus de ma
collection. Arch. Entomol., 1: 291-320.

1860. Essai d’une classification de la famille des cerambycides et materiaux
pour servir a une monographic de cette famille, 404 pp. Paris.

1864. Systema cerambycidarum ou expose de tous les genres compris dans la
famille des cerambycides et families limitrophes. Mem. Soc. Roy. Sci.
Liege, 19: 1-540.

94

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

The Robber Fly Callinicus calcaneus (Loew) As A Predator on

Andrena omninigra Viereck

(Diptera: Asilidae; Hymenoptera: Andrenidae)

E. Gorton Linsley
University of California, Berkeley 94720

In an earlier note, (Linsley, 1944), I reported on the predatory be-
havior of 22 individuals from a population of Callinicus calcaneus
(Loew) foraging on a hillside near Oakhurst, Madera County, Cali-
fornia, which was densely clothed with the common ground cover of
the lower Sierra Nevada yellow pine forests known as mountain misery
or bear-clover ( Chaemaebatia foliolosa Benth.). The robber flies were
capturing bees which were taking pollen and/or nectar from
Chaemaebatia blossoms. The prey in this sample consisted entirely of
solitary bees of the families Megachilidae and Andrenidae ( Osmia spp.
and Andrena spp.). Among the captures, brightly colored Osmia out-
numbered dull colored Andrena by approximately 3:1, although sweep
samples of Chaemaebatia blossoms yielding 536 individual bees indicated
that Andrena outnumbered Osmia by about 5:1. It appeared that the
asilids were selectively capturing the brilliant or metallic colored Osmia.
The flies were foraging in the brightly sunlit areas among the trees
where most of the bees were working and their prey were captured
while in flight between flowers.

It has been suggested previously that bee- and wasp-killing robber
flies tend to locate a suitable foraging site and remain there, selecting
their prey either fortuitously or on the basis of size, coloration, or
some other identifiable attribute, depending upon the extent and range
of choice and prior experience (cf. Linsley, 1960). Further evidence
that this may be true in the case of Callinicus calcaneus was obtained
on 2 June through 5 June 1970 on a hillside 1.5 miles east of Railroad
Flat, Calaveras County, California. Here, as in the Madera County
locality, the robber flies were foraging over Chaemaebatia foliolosa.
However, unlike the earlier situation the observed prey, with one excep-
tion, were exclusively females of Andrena ( Onagrandrena ) omninigra
omnigra Viereck, a wholly black species which was taking nectar from
blossoms of those Chaemaebatia plants which were shaded by scattered
pine trees ( Pinus -ponder osa ) . They were the dominant bees visiting
Chaemaebatia at this time but by no means the only ones. The others,
although relatively few in numbers, were confined to sunny areas and

The Pan-Pacific Entomologist 48: 94-96. April 1972

APRIL 1972] LINSLEY CALLINICUS PREYING ON ANDRENA

95

involved both sexes of two species of brightly colored Osmia and two
Andrena species, the females of which were taking pollen. However,
based on collections from the same hillside in the previous year, these
must have been early seasonal arrivals because in late June of 1969
Osmia and Andrena were abundantly active both in individuals and
species, with much the same species composition as that found at the
Oakhurst site.

These observations suggest that when Callinicus calcaneus adults
emerged (the individuals seen still had their wing margins and body
pubescence intact) the first prey encountered were the densely black
Onagrandrena, which were probably seeking nectar from shaded rather
than sunlit plants in order to minimize integumental heat absorption
while feeding in the exposed cup-like flowers. Presumably having
learned to exploit bees in this situation the robber flies retained this
behavior pattern while the Onagrandrena remained abundant, in spite
of the increasing availability of suitable prey on plants in adjacent
sunny sites. Regrettably, the locality could not be revisited after an
interval of ten days or two weeks to observe the behavior of Callinicus
at that time.

Altogether, 15 female Onagrandrena were taken from Callinicus by
observing their capture, which invariably took place when the bees
were in flight. The robber flies were swept gently into the net shortly
after they settled on a plant with their prey, and were allowed to drop
the then dead bee in the net and escape with a minimum of disturbance.
Under these circumstances marking was not feasible and it is not possible
to say with certainty whether or not the same robber fly was captured
more than once, although the presumption is high that this may have
been so, since only one or two individuals of Callinicus were visible in
the observation area at any one time. Also, the Onagrandrena were
localized to shady spots in an area about 65 yards long by 50 yards wide
near a dirt road, although Chaemaebatia extended on both sides of the
road for hundreds of yards. However, Callinicus are fast fliers and
cover more territory than many other asilid bee predators. They moved
in and out of this foraging site, both when disturbed and undisturbed.

It should be noted that Andrena ( Onagrandrena ) omninigra is a
Clarkia (Onagraceae) oligolege, yet none of the females had traces of
Clarkia pollen in their scopae and, as would be expected, none were
taking pollen from Chaemaebatia (Rosaceae). No Clarkia plants were
evident near the site where the females were working and the only
colony of Clarkia noted in prior years, a substantial population of C.
speciosa , was slightly more than one air mile away on the other side of

96

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

two 300 to 500 ft. ridges. On this date these plants were examined but
they were not yet in bloom. Although they had been sampled twice in
previous years, other Clarkia oligoleges were abundant but no Onagran-
drena were present in the collections. The localized occurrence of the
females in an extensive area of Chaemaebatia, and the capture of a male
nearby, suggested that they might be coming from an adjacent nest site,
but a careful search failed to reveal one if it was in the vicinity.

Diurnally, the earliest Callinicus with Onagrandrena prey was ob-
served at 8:30 a.m. PST, the latest at 4 p.m. Most were taken shortly
before noon (11-11:30 a.m.) and at mid-afternoon (3-3:30 p.m.) al-
though the site was visited intermittently during several days, it was
not under continuous surveillance on any one day.

The identification of two female Callinicus calcaneus captured as
voucher specimens was kindly confirmed by E. I. Schlinger.

Literature Cited

Linsley, E. G. 1944. Prey of the robber fly, Callinicus calcaneus Loew (Diptera:
Asilidae). Pan-Pac. Entomol., 20: 67-68.

1960. Ethology of some bee- and wasp-killing robber flies of southeastern
Arizona and western New Mexico (Diptera: Asilidae). Univ. Calif.
Publ. Entomol., 16: 357-392, pis. 48-55.

BOOK NOTICES

The following two facsimile reprint editions have been issued by E. W.
Classey, Ltd.

Centurie de Lepidopteres de l’Ile de Cura. P. H. Poey. 1832. E. W. Classey,
Ltd., 97 p., 20 color plates, -f- xii. Reprinted 1970. $30.00

This reprint contains a foreword by Lt. Col. C. F. Cowan explaining the
arrangement of the Centurie , bibliographical problems and dates of publication.
An annotated table of contents also is included.

An Illustrated Essay on the Noctuidae of North America. A. R. Grote
1882. E. W. Classey, Ltd., 85 p., 4 color plates, -f- 17 p. foreword. Reprinted
1971. $16.95.

The foreword by Dr. R. S. Wilkinson is an extensive biography of A. R. Grote.
This volume also includes a five page reprint of an essay entitled “A Colony of
Butterflies” which was first printed in 1876, American Naturalist, 10:129-132.

These books are available from the North American distributor: Entomological
Reprint Specialists, P. 0. Box 77971, Dockweiler Station, Los Angeles, California
90007. — R. W. Thorp, University of California, Davis, 95616.

APRIL 1972]

MOSER & ROTON — PYEMOTES SCOLYTI

97

Reproductive Compatibility Between Two Widely Separated
Populations of Pyemotes scolyti

(Acarina : Pyemotidae)

John C. Moser and Lawrence M. Roton

Southern Forest Experiment Station, USD A Forest Service, Pineville,

Louisiana 71360

Pyemotes scolyti (Oudemans) (1936) is an external parasite of
Scolytus spp., and apparently occurs throughout the range of Scolytus.
It was described from specimens reared on Scolytus multistriatus
(Marsham) in elm near Arnhem, Netherlands, and designated by
Krczal (1959) as the genotype.

Specimens of Pyemotes scolyti reared from galleries of Scolytus
ventralis LeConte in Abies grandis collected near Moscow Mountain,
Idaho, were recently forwarded to us for identification by Alan A.
Berryman and Bernard Scott. Although the material was morpho-
logically inseparable from that taken in galleries of Scolytus mul-
tistriatus in Ulmus americana from Delaware, Ohio (det. E. A. Cross),
the possibility still remained that the two populations might be cryptic
species similar to those found for spider mites by Boudreaux (1963).

Methods

Five mothers were reared from each population and five female
and male progeny from each mother were selected for cross matings.
Thus, 25 virgin females reared from 5 mothers of the “ventralis”
strain were mated with 25 males from the “multistriatus” strain and
their hybrid progeny tallied. A similar cross was then made using
females of the “multistriatus” strain and males of “ventralis” strain.

Since the first individual born is usually a male, care had to be
exercised in the cross matings to see that these and any subsequent
males were immediately removed so as not to interfere with the
mating activities of males from the opposite strain. Test males were
identified by a small paint droplet on a seta (Moser and Roton, 1970) .

Results and Discussion

The data in Table 1 show little reproductive isolation between the
two populations, and indicate that they are indeed the same species.
However, some genetic breakdown may have occurred in the male
“multistriatus” X female “ventralis” cross. Although the progeny

The Pan-Pacific Entomologist 48: 97-99. April 1972

98

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Table 1 . Number of Fj. progeny of “ventralis” and “multistriatus”
strains of Pyemotes scolyti.

Progeny (Fi)

Standard

Strain n Mean deviation Maximum Minimum

9

8

$

8

$

8

9

8

Ventralis (mated)

20

94.5

2.1

30.5

1.0

152

5

40

1

Ventralis (unmated)

5

-

63.8

-

46.8

-

135

-

21

Multistriatus (mated)

20

120.6

2.0

20.8

1.6

168

4

59

1

Multistriatus (unmated)

8 Multistriatus

5

—

23.6

—

4.8

—

31

—

19

X

$ Ventralis

8 Ventralis

25

106. 0 1

l.l 2

30.8

0.4

174

50

2

1

X

$ Multistriatus

25

96. 9^

1.3 s

28.5

0.6

168

48

3

1

1 Hybrid females.

2 “Ventralis” males.

3 “Multistriatus” males.

count was normal, males did not appear until about 75 percent of
the females had been born. In addition, one preliminary rearing
resulted in 9 of 20 mothers producing normal numbers of females,
but no males. Production of all-female progeny in Pyemotes parvis-
colyti Cross and Moser was induced by restricting the feeding period
of mothers to 3 days (Moser, Cross, and Roton, 1971). In Pyemotes
scolyti, the virgin females are normally fertilized as they emerge
tail first from the birth canal. The male is embedded head first in
the canal and mating occurs as the female abdomen tip slides past
the male abdomen. Males extracted from the birth canal behaved
in a manner similar to that described by Krczal (1959). They mated
with virgin females, but only with difficulty. They reentered the
canal as soon as possible, usually mating with only a few of the
available females. Normally, males leave the birth canal when they
are pushed out by the next male to be born. Occasionally, males
left the canal for no apparent reason, but in both cases, they usually
died within 12 hours, and never were observed to reenter the canal.
In at least 3 cases where virgin females were present when males
left the canal, no matings took place. Hence, any females born before
the first male emerge unfertilized, and probably remain so even when
males are born later. Thus, in the male “multistriatus” X female
“ventralis” matings, there was a 75 percent loss in reproductive
potential of progeny.

APRIL 1972]

MOSER & ROTON PYEMOTES SCOLYTI

99

Virgin females of both populations were reluctant to attack the
host, Scolytus multistriatus larvae. When they did, swelling was
usually minimal, resulting in a reduced number of progeny, all of
which were male. As Boudreaux (1963) showed in his experiments
with tetr any chid mites, rearing through the F 2 generation is not
necessary to show reproductive isolation in species exhibiting haplo-
diploid parthenogenesis (arrhenotoky) such as we observed. Since
the Fi males carry the same genes as the female parent, the F 2
generation is really a backcross of the Fi generation and not a true
F 2 cross. Therefore, the F 2 generations will exhibit an average of
characters closer to the average original female parent, and suc-
ceeding generations will even be more biased.

Cross (1965) notes that many pyemotids, due to their small size,
are cosmopolitan within their ecological limits, and that greater
morphological variation may occur between nearby localities than
between continents. Perhaps this phenomenon also applies to breeding
barriers between widely separated populations of Pyemotes scolyti.

The “multistriatus” population is probably of European origin and
introduced with the elm bark beetle sometime prior to 1909. The
“ventralis” population may or may not be native, but the formidable
ecological barriers between the host beetles make it unlikely that
this population was recently derived from the “multistriatus” popu-
lation.

Acknowledgments

We thank Drs. H. B. Boudreaux and E. A. Cross for interest and
advice on methodology. We also thank Dr. Alan Berryman, Mr.
Bernard Scott, Dr. Bruce H. Kennedy, and Dr. John W. Peacock for
collecting the mites and bark beetles used in the study.

Literature Cited

Boudreaux, H. B. 1963. Biological aspects of some phytophagous mites. Ann.
Rev. Entomol., 8: 137-154.

Cross, E. A. 1965. The generic relationships of the family Pyemotidae
( Acarina : Trombidiformes) . Univ. Kans. Bull., 45: 29-275.

Krczal, H. 1959. Systematik und Okologie der Pyemotiden. Beitr. Syst. Okol.
Mitteleur. Acarina, 1(2): 385-625.

Moser, J. C., E. A. Cross, and L. M. Roton. 1971. Biology of Pyemotes
parviscolyti (Acarina : Pyemotidae) . Entomophaga, 16(4): 367-379.
Moser, J. C., and L. M. Roton. 1970. Tagging mites with aerosol paint. Ann.
Entomol. Soc. Amer., 63: 1784.

Oudemans, A. C. 1936. Neues iiber Pedicidoides Targ. Tozz. 1878. Festschr.
fiir Prof. Dr. Embrik Strand, 1: 391-404.

100

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

A Report on the Arthropods Collected by the Edinburgh
University Galapagos Islands Expedition, 1968 1

Patricia Parkin , 2 D. T. Parkin , 2 A. W. Ewing , 3 and H. A. Ford 4

The Galapagos Archipelago has attracted much scientific study since
Charles Darwin’s visit in 1835 first drew the attention of the world’s
biologists to its potential. The initial studies were of collections
obtained for description and taxonomic determination of the plants
and animals present among the islands. Linsley and Usinger (1966)
list 21 expeditions of various sizes and eight individual collectors
who worked in the islands for greater or lesser periods of time.

The primary objects of non-taxonomic study have been the verte-
brates, notably the birds (Lack, 1947; Bowman, 1961, 1963; and
many others), while the terrestrial invertebrates have been compar-
atively neglected (important exceptions being Curio, 1966, and Linsley,
1966) . Even distributional data of invertebrates is fragmentary. Hence,
casual collecting by non-specialists can yield valuable information,
and new species can even be found in routine sampling of insects
around the lights of the Research Station on Santa Cruz.

The authors of this paper worked in the Galapagos Archipelago
from July to November 1968. Arthropods were collected whenever
time could be spared from principal research projects. The material
is consequently highly non-random, being concentrated around the
Research Station in Academy Bay with supplementary material from
the highlands of Santa Cruz. Additional collections were made on
Santiago and Floreana, with only a few specimens being taken on
several other islands.

This paper lists, for the interest of students of Galapagos entomology,
the material which we collected. Details are given of the islands,
locality, habitat zones and date. The number of specimens is also
given together with the depository if other than the Royal Scottish
Museum, Edinburgh. We are extremely grateful to the authorities
who identified the material. Their names are given below, and all
identifications and notes or comments in the checklist are initialled
for the authority concerned.

1 Contribution No. 142 from the Charles Darwin Research Station.

2 Present Address: The Genetics Laboratory, School of Botany & Zoology, University Park,
Nottingham, England.

3 Present Address : Department of Biological Sciences, Portsmouth Polytechnic, Portsmouth,
England.

4 Present Address: Department of Biology, University of Stirling, Bridge of Allan, Stirling,
Scotland.

The Pan-Pacific Entomologist 48: 100-107. April 1972

APRIL 1972] PARKIN ET AL . — GALAPAGOS ARTHROPODS

101

It is interesting to note that Linsley & Usinger (1966) list 618
species of insects from the Galapagos Archipelago reported in the
literature prior to 1966. We collected 72 of these, approximately
11.5%. To obtain such a high proportion of the known species in
such a short time suggests either that we collected in the same
localities as previous workers and duplicated their samples, or that
the fauna is very impoverished and many species are widespread.
An argument in support of the latter alternative is furnished by the
consideration that we amassed 80 species from individual islands,
of which no less than 38 are island records new to Linsley & Usinger’s
list. Our authorities were unable to put names to 8, leaving 30
records of previously known insects on new islands-about 42% of
our material. This suggests strongly that the Galapagos insect fauna
is limited in number of species, but that many species occur on
several islands.

In the systematic list which follows, an asterisk in front of the
species name indicates that the species is not listed by Linsley &
Usinger (1966). An asterisk in front of the island name signifies
that the species is not recorded from that island in the same work.
All dates refer to 1968.

Class Arachnida
Scorpiones: Vejovidae

Hadruroides lunatus L. Koch. Santiago: At settlement in James Bay, Arid
Zone, 18 August, 1; Waterhole by James Bay, Arid Zone, 18 August, 1;
Floreana: base of highest volcano under stone, 1,200 ft. SW side, Transition
Zone, 31 August, 1.

Scorpiones : Buthidae

Centrums princeps Karsch. Santa Cruz: East of Research Station, coastal scrub,
Arid Zone, 4 August, 1.

Araneae : Argiopidae

Gasteracantha insulana Thorell. Santa Cruz: East of Research Station, 30 yards
from sea, ca 40 ft., Arid Zone, 4 August, 1 female.

All Arachnida determined by D. J. Clark, (NH) 1970, who adds
that these species have been recorded from the Galapagos Is. by Banks
(1902, Proc. Wash. Acad. Sci ., 49-86).

Class Insecta
Dermaptera

Euboriella annulipes Lucas. (=r Anisolabis bormansi Scudder, tests A. Brindle) .
(*) Floreana: 2,100 ft., at top of volcano, grassland, short vegetation,
Transition Zone, 31 August, 1 nymph. (Det. A.B. 1970).

102

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Dictyoptera : Blattidae

Periplaneta australasiae Fabricius. Floreana: 31 August, 1 (Det. D.R.R. 1969).

Dictyoptera: Pycnoscelidae

Pycnoscelus surinamensis Linnaeus. (*) Floreana: base of main volcano, three
miles from Black Beach, Arid Zone, 31 August, 1. Santa Cruz: half way
along old trail from Puerta Azora-Bella Vista, Transition Zone, 8 September,
2 juvs. (Det. D.R.R. 1969).

Saltatoria: Tettigoniidae

Nesoecia cooksoni Butler. (*) Santiago: on volcano in James Bay, Arid Zone,
17 August, 1 male (Det. D.R.R. and retained by BM (NH)).

Saltatoria : Gryllidae

Cryptoptilum sp. (*) Santiago: on volcano in James Bay, Arid Zone, 17 August,
1 female (Det. D.R.R. 1969) .

Mantodea : Acrididae

Schistocerca melanocera Stal. (*) Santiago: in crater of salt mine, James Bay,
Arid Zone, 15 August, 1 female & 1 juvenile; at Settlement in James
Bay, Arid Zone, 18 August, three males. (Det. V.M.D. 1969) .

Halmenus choristopterus Snodgrass. Floreana: 30 August, 1 female. (Det.
V.M.D. 1969) .

Halmenus robustus Scudder. Santa Cruz: on vegetation near La Caseta, Humid
Zone, 1. (Det. V.M.D. 1969) .

Odonata: Aeschnidae

Anax amazili Burmeister. (*) Santa Cruz: La Caseta, Humid Zone, 15 October,
one male. (Det. A.R.W. 1970) .

Hemiptera : Lygaeidae

Heraeus pacificus Barber. Santiago: 11 August, 1 female (Det. G.M.B. 1970).

Neuroptera: Hemerobiidae

Megalmus darwini Banks. Santa Cruz: Research Station, Arid Zone, 25
September-14 October, six (Det. A.R.W. 1970).

Neuroptera: Chrysopidae

Chrysopa nigripilosa Banks. (*) Santa Cruz: Research Station, Arid Zone,
12-27 October, 25. (*) Santiago: near hut, James Bay, Arid Zone, 14

August, 1 ; settlement, Arid Zone, 22 September, 1. (Det. A.R.W. 1970) .

Chrysopa sp. near galapagoensis Banks. ( :|! ) Floreana: Wittmer Farm, Humid
Zone, 29 August, 1. (*) Santa Cruz: Research Station, Arid Zone, 22
September, 1. (*) Santiago: Humid Zone, 23 September, common, 1
specimen taken. (Det. A.R.W. 1970) .

Neuroptera : Myrmeleontidae

Myrmeleon perpilosus Banks. Santa Cruz: Research Station, Arid Zone, 21
August-28 October, 6 males, 11 females, 2 larvae. (*) Santa Fe: came
to light, 14 September, 1 male. (Det. A.R.W. 1969).

APRIL 1972] PARKIN ET AL . — GALAPAGOS ARTHROPODS

103

(*) Brachynemurus dcirwini Stange. Santiago: James Bay settlement, 21
September, 1 female, 1 larva. (Det. A.R.W. 1969) .

Lepidoptera : Pieridae 5

Phoebis sennae Linnaeus. San Cristobal: 600 ft. in Humid Zone below village,
1 August, 5 females. Santa Cruz: half mile west of Research Station, on
coast by brackish pool, Arid Zone, 22 August, 2 females. Santiago: James
Bay, around flowering shrubs growing between lava blocks, Arid Zone, 14
& 21 August, 9 males & 3 females; James Bay, at settlement, Arid Zone,
21 September, 1 male.

Lepidoptera : Danaidae

(*) Danaus gilippus ssp. near strigosus Bates. San Cristobal: near village,
600 ft., Humid Zone, 1 August, 4 males & 4 females.

Lepidoptera : Hesperiidae

Urbanus dorantes galapagensis Williams. San Cristobal: near village, 600 ft.,
Humid Zone, 1 August, 1 male. Santa Cruz: La Caseta, Humid Zone,
16-18 October, 10 males.

Lepidoptera: Spiiingidae

Erinnyis ello encantada Kernbach. Santa Cruz: Research Station, at light,
Arid Zone, 4 August, 1 female.

Erinnyis obscura conformis Rothschild & Jordan. Santa Cruz: Research Station,
Arid Zone, 19 August-21 October, 2 males & 1 female.

Pholus labruscae yupanquii Kernbach. Santa Cruz: Research Station, Arid Zone,
10 August, 1 male.

Celerio lineata florilega Kernbach. Santa Cruz: Research Station, Arid Zone,
28 October, 1 female.

Lepidoptera: Arctiidae

Utetheisa galapagensis Wallengren. Santa Cruz: Research Station, at light, Arid
Zone, 3-5 August, 5 females; 25 September-3 October, 1 male & 6
females; Hornemann Farm, near Bella Vista, 650 ft., Humid Zone, 9
October, 2 females (Det. A.H.H. 1970) .

TJtetheisa ornatrix Linnaeus. (*) Santa Cruz: La Caseta, Humid Zone, 15
October, 1 female.

Lepidoptera: Noctuidae: Agrotinae

Agrotis ipsilon Hufnagel. (*) Santa Cruz: Research Station, Arid Zone, 27
September, 1 male.

Lycophotia oceanica Schaus. (*) Santa Cruz: Research Station, at light, Arid
Zone, 5 August, 1 female; 21 September-12 October, 1 female & 1 male.

Lepidoptera: Noctuidae: Hadeninae

Mythima sequax Franclemont. Santa Cruz: Hornemann Farm, near Bella
Vista, 650 ft, Humid Zone, 9 October, 1 male. (American authors put
this in Pseudaletia Franclemont).

6 All Lepidoptera determined by E. C. Pelliam-Clinton unless otherwise stated.

104

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Lepidoptera: Noctuidae: Amphipyrinae

Magusa sp. (*) Santa Cruz: Research Station, Arid Zone, 21 August, 1 male;
4 October, 2 males. ( Teste A.H.H. 1970, the Magusa of Galapagos is 'not
M. orbifera (Walker), but is undescribed).

Platysenta apameoides Guenee (= ebba Schaus) . (*) Santa Cruz: Research

Station, Arid Zone, 6 October, 1 female; Hornemann Farm, near Bella
Vista, 650 ft., Humid Zone, 9 October, 1 female. (Teste A.H.H. 1970, the
species formerly known as P. apameoides and so listed by Linsley &
Usinger (1966) is P. plagiata (Walker)).

Harrisonia williamsi Schaus. Santa Cruz: Research Station, Arid Zone, 22
August-14 October, 6 females.

Amyna insularum Schaus. Santa Cruz: Research Station, Arid Zone, 26
September-4 October, 1 male & 2 females.

Elaphria dubiosa Schaus. Santa Cruz: Research Station, Arid Zone, 4-21
October, 4 females & 1 male, (described by Schaus as a Rivula and so
listed by Linsley & Usinger (1966), in Catocalinae-A.H.H., 1970).

Lepidoptera: Noctuidae: Catocalinae-including Ophiderinae

Melipotis harrisoni Schaus. Santa Cruz: Research Station, Arid Zone, 5
August-12 October, 2 females & 1 male.

(*) Lyncestis sp. near acontioides (Guenee). Santa Cruz: Research Station,
Arid Zone, 12-13 October, 1 male & 1 female ( Teste A.H.H. 1970, the
species near acontioides should be separated from Melipotis ) .

Epidromia zephyntis Schaus. This species occurs in two forms in both sexes;
the forms appear to be inseparable by genitalia. Typical (dark) form-Santa
Cruz: Research Station, at light, Arid Zone, 5 August, 2 females; 13
October, 1 female. Light form-Santa Cruz: Research Station, at light,
Arid Zone, 25 September-12 October, 3 males & 1 female.

Psorya hadesia Schaus. (*) Santa Cruz: Research Station, at light, Arid
Zone, 3 August, 1 female; 3-4 October, 1 male & 1 female.

Mods incurvalis Schaus. Santa Cruz: Research Station, Arid Zone, 4 October,
1 female.

(*) Glympis sp. Santa Cruz: Research Station, Arid Zone, 25 September-14
October, 3 males & 2 females. (Det. A.H.H. 1970).

Lepidoptera: Geometridae

(*) Eupithecia leleupi Herbulot. Santa Cruz: Research Station, Arid Zone,
12 October, 1 female. (Det. D.S.F. 1970) .

Camptogramma stellata Guenee. Santa Cruz: Research Station, at light, Arid
Zone, 3 August-6 October, 12 females.

Cosymbia impudens Warren. Santa Cruz: Research Station, at light, Arid
Zone, 3 August-4 October, 2 males & 1 female. Santiago : Settlement, Arid
Zone, 23 September, 3 females.

(*) Semiothisa cruciata Herbulot. Santa Cruz: Research Station, Arid Zone,
26 September, 2 females (1 without abdomen). (Det. D.S.F. 1970).

Lepidoptera: Pyralidae

(*) Hymenia perspectalis Hubner. Santa Cruz: Hornemann Farm, 650 ft., near
Bella Vista, Humid Zone, 8 October, 1 female.

APRIL 1972] PARKIN ET AL . — GALAPAGOS ARTHROPODS

105

(*) Stemorrhages sp. near lustralis (Guenee). Santa Cruz: Research Station,
Arid Zone, 12 October, 1 male. (Det. M. Shaffer 1970) .

Piletocera bujalis Guenee. Santa Cruz: Research Station, 3 October, 1 male.

Beebea guglielmi Schaus. (*) Santa Cruz: Research Station, at light, Arid
Zone, 4 August, 1 female.

Lepidoptera: Tortricidae

Crocidosema plebeiana Zeller. (*) Santa Cruz: Research Station, Arid Zone,
3 October, 1 female (without head) .

Lepidoptera : Gelechiidae

Arislotelia naxia Meyrick. (*) Santa Cruz: Research Station, Arid Zone, 3
October, 1 male.

Lepidoptera : Blastobasidae

Blastobasis crotospila Meyrick. (*) Santa Cruz: Research Station, Arid Zone,
26 September, 1 female.

Lepidoptera: Hyponomeutidae

Atteva hysginiella Wallengren. Santiago: James Bay, near hut, Arid Zone,
14 August, 1 female.

Coleoptera : Carabidae

Selenophorus galapagoensis G.R. Waterhouse. (*) Pinta: Arid Zone, 22 September.
(Det. P.M.H.).

Coleoptera: Oedomeridae

Alloxacis collenettei Blair. Santa Cruz: Research Station, 14 October, 2 (Det.
by C.M.F. von H. as Oxacis collenettei) .

Coleoptera: Elateridae

Physorinus quirsfeldi Mutchler. Santa Cruz: Research Station, Arid Zone,
12-14 October, 2. (Det. C.M.F. von H. 1970) .

Coleoptera: Coccinellidae

Cycloneda sanguinea Linnaeus. Santa Cruz: On Mora vegetation, about 1.5
miles west of La Caseta in the tortoise reserve, Wet Zone, 2; Research Station,
Arid Zone, 9 August, 1. (Det. R.P.D. 1970).

Coleoptera : Rliipiphoridae

(*) Genus and species unknown at the British Museum (Natural History).
Santa Cruz: Research Station, Arid Zone, 28 August, 1; 13-14 October, 2.

Coleoptera: Tenebrionidae

Stomion galapagoensis G.R. Waterhouse. Floreana: 2.5 miles east from Black
Beach along track, 1100 ft., Arid Zone, 3; On top of volcano, 2,100 ft.,
in grass and short vegetation, 31 August, 4; 3 miles east of Black Beach, at
foot of final cone of volcano, 1,500 ft., Arid Zone, 31 August, 2. (Det.
M.J.D.B. 1970).

Stomion laevigatum G.R. Waterhouse. Santiago: On top of volcano in James
Bay, Arid Zone, 11 August, 1. (Det. M.J.D.B. 1970).

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Stomion rugosum Van Dyke. Pinta: Arid Zone, 22 September, 1. (Det.
MJ.D.B. 1970).

Ammophorus bifoveatus G.R. Waterhouse. (*) Santiago: On top of volcano in
James Bay, Arid Zone, 11 August, 3. (Det. M.J.D.B. 1970).

Pedonoeces bauri Linell. (*) Floreana: 3 miles east of Black Beach, at foot
of final cone of volcano, 1,200 ft., Arid Zone, 31 August, 1. (Det.
MJ.D.B. 1970).

Pedonoeces opacus Van Dyke. (*) Santa Cruz: 1 mile east from Research
Station, 150 yards from coast, Arid Zone, 4 August, 1 (Det. MJ.D.B. 1970).

CoLEOPTERA: BOSTRICHIDAE

Amphicerus cornutus galapaganus Lesne. Santa Cruz: Research Station, Arid
Zone, 23 October, 1 (Det. C.M.F. von H. 1970).

COLEOPTERA: CERAMBYCIDAE

Eburia lanigera Linell. (*) Santa Cruz: Research Station, Arid Zone, 12-13
October, 2. (Det. R.T.T. 1970) .

Estoloides galapagoensis Blair. Santa Cruz: Research Station, Arid Zone, 13-14
October. (Det. R.T.T. 1970) .

Acanthoderes galapagoensis Linell. Santa Cruz: Research Station, Arid Zone,
13 October, 1. (Det. R.T.T. 1970) .

CoLEOPTERA : CURCULIONIDAE

Pantomerus galapagoensis Linell. (*) Floreana: 3 miles east of Black Beach,
in dry scrub, 1,500 ft., Arid Zone, 31 August, 1. (Det. R.T.T. 1970).

Diptera : Tipulidae

Limonia ( Geranomyia ) tibialis Loew. Santa Cruz: Media Luna, in grassland,
Wet Zone, 18 October, 1 female. (Det. A.M.H. 1970) ; Research Station,
at night, Arid Zone, 20 August, 1 male. (Det. A.R.W. 1970) .

Diptera : Culicidae

Aedes ( Ochlerotatus ) taeniorhynchus Wiedemann. Santa Cruz: Research Station,
found dead on lab. bench in morning, Arid Zone, 5 August, 1 female.
(Det. A.H.H. 1970).

Diptera : Tab anidae

Tabanus vittiger Thomson. Santa Cruz: Research Station, Arid Zone, 2
September, 1 female. (Det. H.O. 1970).

Diptera : Calliphoridae

Sarcophaga sp. Santa Cruz: Research Station, 2 August, 1 male.

Hymenoptera : Apidae

Xylocopa darwini Cockerell. Santa Cruz: found near brackish pool, 0.5 mile
west of Research Station, Arid Zone, 22 August, 1 female. (Det. A.R.W. 1970) .

Hymenoptera: Mutillidae

Photopsis sp. ( :,: ) Santiago: settlement, Arid Zone, 23 September, 1 male.
(Det. M.C.D. 1970).

APRIL 1972] PARKIN ET AL . — GALAPAGOS ARTHROPODS

107

Acknowledgments

The Edinburgh University Galapagos Islands Expedition was financed
principally by The Percy Sladen Trust, The British Museum (Natural
History) , Shell International Marine Limited, The Carnegie Trust
for the Universities of Scotland, and Edinburgh University. Accom-
modation was provided at the Charles Darwin Research Station, and
we are most grateful to the Director, Mr. Roger Perry, and to Dr.
Tjitte de Vriess for making us so welcome.

The authorities who performed the identifications are: G. M. Black
(Hemiptera), M. J. D. Brendell (Coleoptera) , A. Brindle (Dermaptera) ,
D. J. Clark (Arachnida) , M. C. Day (Hymenoptera) , V. M. Dirsch
(Mantodea), D. S. Fletcher (Lepidoptera) , P. M. Hammond (Cole-
optera), C. M. F. von Hayek (Coleoptera), A. H. Hayes (Lepidoptera),
A. M. Hutson (Diptera), H. Oldroyd (Diptera), E. C. Pelham-Clinton
(Lepidoptera), R. D. Pope (Coleoptera), D. R. Ragge (Orthoptera) ,
R. T. Thomson (Coleoptera), A. R. Waterson (Odonata, Neuroptera,
Hymenoptera and Diptera) .

Literature Cited

Bowman, R. I. 1961. Morphological Differentiation and Adaptation in the
Galapagos Finches. Univ. Calif. Publ. Zooh, 58: 1-302.

1963. Evolutionary Patterns in Darwin’s Finches. Calif. Acad. Sci. Occas.
Pap., 44: 107-140.

Curio, E. 1966. Farbung und Reheverhalten Dreier Raupenformen eines
Schwarmes. Umsch. Wiss. Tech., 66: 475.

Lack, D. 1947. Darwin’s Finches. Cambridge Univ. Press, 208 p.

Linsley, E. G. 1966. Pollinating Insects of the Galapagos Islands, in The
Galapagos: Proceedings of The Symposium of the Galapagos Inter-
national Scientific Project, ed. R. I. Bowman. Univ. Calif. Press, 318 p.
Linsley, E. G. and R. L. Usinger. 1966. Insects of the Galapagos Islands. Proc.
Calif. Acad. Sci., 33: 113-196.

NOTICE

An annual Directory of Environmental Consultants, will be published beginning
in 1972. Environmentally concerned professionals interested in having their
name and short resume appear in the Directory should send a No. 10, self-
addressed, stamped envelope to: Directory of Environmental Consultants, P. 0.
Box 8002, University Station, St. Louis, Missouri 73108.

108

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

A New Genus and Species of Jumping Shore Beetle

from Mexico

(Coleoptera: Limnichidae)

T. J. Spilman

Systematic Entomology Laboratory, Entomology Research Division,

Agr. Res. Serv., USDA 1

In the fall of 1966 William G. Evans of the University of Alberta
collected insects, pseudoscorpions, and chilopods from the intertidal
zone of the western coast of Mexico. As he says in his introductory
report (1968), he studied about one thousand miles of Mexican main-
land coast from Guaymas in Sonora to Barra De Navidad in Jalisco;
this coast and that of the southern tip of Baja California and the
Hawaiian Islands comprise a distinct marine littoral faunal zone,
which Evans designates the Subtropical Zone. His objective was,
among other things, to compare the ecology of the Mexican coast
with that of others he had studied. However, an interesting by-
product of his trip came at the southernmost point when he discovered
a remarkable new genus and species of intertidal beetle.

These little beetles, about two millimeters long, dark, compact and
somewhat streamlined, have long hind legs bearing heavy long setae
or spurs. Their morphology, given below in formal descriptions,
places them in the subfamily Thaumastodinae of the Dryopoid family
Limnichidae. Evans’ beetles were aggregated in groups of six to
twelve in high and mid-tide rock crevices, and they jumped an inch
or two when disturbed but were otherwise slow moving. One other
member of the subfamily, Martinius ripisaltator Spilman, lives in the
marine intertidal zone of Cuba and also jumps. Although all other
Thaumastodinae do live near water, either fresh or salt water, none
of those near fresh water has been recorded as being a jumper.

Heretofore this subfamily contained three genera: Acontosceles
Champion, with one species in India and another in the Philippines;
P seudeucinetus Heller, with a single species in the Philippines and
Malay Penninsula; Martinius Spilman, with one species in the Canal
Zone and another in Cuba. I reviewed the taxonomy and morphology
of the subfamily (1959) and then added the Cuban species (1966).
In the first article I tried to indicate similarities and relationships
of the three genera and came to the conclusion that P seudeucinetus

1 Mail address: c/o U. S. National Museum, Washington, D. C. 20560

The Pan-Pacific Entomologist 48: 108-115. April 1972

APRIL 1972]

SPILMAN JUMPING SHORE BEETLE

109

is an intermediate genus, but it is much more closely related to
Martinius than to Acontosceles. In other words, Acontosceles stands
well apart.

This new genus, described below under the name Mexico , has five
outstanding characteristics. First, the antenna is very odd. At first
glance the general shape of the antenna seems similar to that of
P seudeucinetus , but the number of articles appears to be 10 instead
of the normal 11 because the second and third articles seem to be
one. However, what appears to be a mere line or constriction on the
apical third of this seemingly single article is actually a division or
j oint, and the morphology becomes obvious when the two are separated.
The second article is large, the third small (Fig. 8). The apex of
the second is concave, and the base of the third has an opposing
concavity and a slender medial stalk; that stalk is actually the base of
the third and fits into a hole in the center of the concavity of the
second. I wonder about the function of such an arrangement; why
should there be opposing, close-fitting concavities on neighboring
antennal segments? One other genus, Martinius , also has an odd
antenna, but that oddity is not in its shape but in its number of
articles — it has only seven. Both Acontosceles and P seudeucinetus
have normal antennae of eleven articles. Second, Mexico has a
sharply margined, deep, bare antennal groove immediately above the
mandible, just wide enough to contain the antenna (Fig. 2). The
groove does not exist in other genera. This groove is located in the
broad, shallow, setose depression that is present in other genera;
I had previously called that broad depression the antennal groove.
Third, the eyes of Mexico are widely separated, very nearly vertical
in position, and suboval (Fig. 2). However, this genus is not alone
in having distinctive eyes ; the eyes of each genus are differently shaped
and arranged. The eyes of Mexico are the most widely separated in the
subfamily, and their shape and position are intermediate between
those of Acontosceles and P seudeucinetus. Fourth, the prosternum
of Mexico , the longest in the subfamily (Fig. 1), is one and a half
times as long as the prosternum of P seudeucinetus and twice as long
as that of Martinius. Fifth, the metasternum is long (Fig. 1), almost
as long as that of Acontosceles ; this structure is very short in
P seudeucinetus and Martinius. Also, the metacoxae of Mexico are
moderately oblique, of moderate size, and have moderately large
plates. In Acontosceles the metacoxae are only slightly oblique, are
relatively small, and have small plates ; in P seudeucinetus and Martinius
they are very oblique, are very large, and have very large plates. The

110

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Figs. 1-7. Mexico litoralis, new species. Fig. 1. Beetle, ventral view. Fig.
2. Head, anterior view. Fig. 3. Antenna. Fig. 4. Labium, aboral view. Fig.
5. Hind leg, anterior view. Fig. 6. Male parameres and pars basalis, dorsal
view. Fig. 7. Male penis, dorsal view; outline of cross section at right.

APRIL 1972]

SPILMAN JUMPING SHORE BEETLE

111

length of the metasternum is inversely proportional to the size of
the metacoxae in all four genera.

The majority of comparisons made in the generic description of
Mexico indicates a close relationship to P seudeucinetus and Martinius.
However, I feel that the five characteristics discussed in the preceding
paragraph indicate that Mexico is intermediate between Acontosceles
on the one hand and P seudeucinetus and Martinius on the other. That
relationship, surely only a conjecture, might be expressed in a linear
manner as follows, with the number of hyphens being directly propor-
tional to distance of relationship: Acontosceles— Mexico— Pseudeucine-
tus-Martinius

Each time I look critically at members of this subfamily some
different facet of functional morphology catches my eye. In my
first paper (1959) I commented on several structures and their
functions; in my second (1966) I discussed the functional significance
of position of setae on the metatarsi. Now I am able to report on the
morphology of the closure of the cavity formed by the elytra and
abdomen. Closure is effected by three structures: a lateral seal, a
lateral clasp, and an apical clasp. First, a lateral seal is formed by a
dorsally projecting, continuous, longitudinal, thin, sclerotized plate or
flange on each side of the dorsum of the abdominal segments (Fig.
10). This plate is actually a dorsal projection of all the combined
sternites, and the lightly sclerotized tergites are continuous with the
plate. The plate fits tightly against the underside of the lateral edge
of the elytra when the latter are at rest. Second, a lateral clasp
(Fig. 11) is formed by a very small, thin, dorsal projection on the
lateral edge of the metacoxa. That projection fits into a very small,
very narrow and short slit on the lower edge of the elytral pseudo-
pleuron. Third, an apical clasp is formed in two different ways in
the subfamily. The clasp is similar in P seudeucinetus, Martinius, and
Mexico (Fig. 12). The last visible abdominal sternite has a notch
on each side of the midline of the posterior border and a slightly
expanded process at the apex of that notch. The elytral apex has a
truncate process, and the lateral surface of that process is concave.
When the combined elytra are closed against the abdomen, the
processes of the elytral apices fit between the processes of the abdominal
sternite; the medial surface of each abdominal process fits snugly
into the concavity of the lateral surface of the elytral process. However,
the apical clasp in Acontosceles is different (Fig. 13). The apex of
the elytron is slightly projected, but the projection is not parallel
sided nor is it truncated. Proximad to the apex of the elytron the

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Fig. 8-12. Mexico litoralis, new species. Fig. 8. Male sternite 9, dorsolateral
view. Fig. 9. Male tergite 9, dorsolateral view. Fig. 10. Abdomen in region
of 2nd visible sternite, diagrammatic cross section; a-lateral plate, b-dorsal
surface, c-ventral surface. Fig. 11. Lateral area of left elytron and abdomen,
showing lateral clasp opened; a-lateral clasp, b-elytral epipleuron, c-elytral
pseudopleuron, d-metacoxa. Fig. 12. Apex of elytra and abdomen, showing
apical clasp opened. Fig. 13. Acontosceles hydroporoides Champion. Apex of
elytra and abdomen, showing apical clasp opened.

APRIL 1972]

SPILMAN JUMPING SHORE BEETLE

113

lateral border is incurved and from that incurved section a small
vertical flange projects downward. The posterior border of the last
visible abdominal sternite does not have notches or processes, but it
is slightly rolled upward or recurved on its lateral thirds. When the
combined elytra are closed against the abdomen, the recurved border
of the sternite presses against or grips the flanges of the elytra.

Each of these three structures has a distinct function in the common
cause of closure. The lateral seal forms a tight seal, keeping water
from the subelytral cavity. The lateral clasp aligns the borders of
the elytra and abdomen, preventing lateral movement of the elytra.
The apical clasp functions as a locking mechanism and as a seal in
the area where movement of the apical abdominal sternite must be
possible. All three structures serve to maintain the integrity of the
subelytral cavity.

Key to the Genera of Thaumastodinae

1. Tarsal formula 4-5-5. Eye acutely angulate ventrally. Antenna inserted

very near eye Acontosceles Champion

Tarsal formula 4-4-4. Eye evenly arcuate or broadly angulate ventrally.
Antenna inserted near mandible and distant from eye 2

2. Antenna with 7 articles Martinius Spilman

Antenna with 11 articles or apparently with 10 articles 3

3. Antenna with articles 2 and 3 seemingly united, with continuous border

(Fig. 3) . Elytra with lateral border serrate posteriorly

Mexico Spilman, new genus

Antenna with 11 obvious articles, all articles distinct. Elytra with lateral
border entire posteriorly Pseudeucinetus Heller

The four genera in the Thaumastodinae are compared in each
character in the following description of the new genus. If the
character mentioned is identical or similar to that same character in
Acontosceles , an “(A) ” is placed after the character. A “(P) ” indicates
similarity or identity to Pseudeucinetus , and an “(M)” indicates
Martinius. If the character is unique or greatest in Mexico, the
exclamation mark “ ( ! ) ” is used.

Mexico Spilman, new genus

Head (Fig. 2) with face weakly convex in lateral aspect (PM) ; epistomal
suture present (PM). Eyes (Fig. 2) subovate, dorsolateral, widely separated (!).
Antennal groove (Fig. 2) broad, margined dorsally, with shiny, deep, acutely
margined groove in ventral third of broad groove (!). Antenna (Fig. 3) inserted
near mandible (PM) ; short, with 11 articles (PA) ; middle segments sub-
moniliform (PM) ; with club of four articles (!) ; 2nd article large, 3rd small,
2nd and 3rd almost fused and together appearing as one ( ! ) . Labrum with
lateral borders evenly arcuate (PM). Mandible with apical slender part not
offset from lateral border of main body of mandible ( ! ) ; lacinia mobile longer,

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its apex at level of apex of mandible proper ( ! ) ; without elevation at base of
lacinia mobile ( ! ) . Maxilla slender, with galea simple apically (PM) . Labium
(Fig. 4) with postmentum as wide as long (PM) ; ligula with broad, blunt
lateral projections (!); 2nd palpal article longer (PM) and much broader
than 3rd (!). Pronotum evenly convex (PM). Prosternum (Fig. 1) relatively
long anterior to procoxal cavities, longest prosternum in tribe ( ! ) . Mesosternum
(Fig. 1) on same level as prosternal process (PM) , with distinctly bordered
depression for reception of prosternal process (M), linear in area anterior to
each mesocoxa (PM) . Metastemum (Fig. 1) evenly convex, short anterior to
metacoxae, slightly expanded laterally, posterior borders moderately converging
medially, becoming triangular between coxae (in all respects intermediate
between A and PM) . Metepistemum (Fig. 1) very narrow, widest anteriorly,
becoming linear posteriorly (PM) . Hind tibia (Fig. 5) with many heavy spurs
(PM). Tarsal formula 4-4-4 (PM). Foreleg not exhibiting sexual dimorphism
(A) . Metacoxa (Fig. 1) oblique, large, its length equal to distance between
it and mesocoxa (A) . Elytra with lateral border posteriorly serrate (M) ; apex
with truncate process (PM) (Fig. 12). Abdomen (Fig. 1) with first visible
stemite very long laterally and very short medially where large hind coxae
encroach (PM) ; with border of ultimate visible sternite bidentate (PM) (Fig.
12). Metendosternite with stalk not bulbous anteriorly (PM); vertical plate
not thickened on dorsal border (PM) ; anterior tendons not visible (PM) . Male
with aedeagus (Fig. 6) with tegmen ventral to penis when retracted within
abdomen (M) ; with pars basalis symmetrical, not sclerotized dorsally, forming
trough for penis (M) ; with parameres obviously longer than basal piece (!)
and not acuminate apically (PM) ; sternite nine (Fig. 7) symmetrical (M) ;
sternite eight (Fig. 8) small, modified V-shaped (PM).

Type-species: Mexico litoralis Spilman, new species.

The generic name Mexico should be considered to be in the third
declension: Mexico, Mexiconis. The name would be similar to Dido
and Leo; it is masculine.

Mexico litoralis Spilman, new species

Dorsal surface covered with dense, fine, setigerous punctures. Setae on dorsal
surface of head similar to those on pronotmn and elytra except directed anteriorly;
setae on anterior surface of head strongly arched and directed ventrally. Setae
on pronotum and elytra of three kinds. First kind very dense, short, fine,
appressed, and brown, and covering most of surface. Second kind very dense,
short, broad, appressed, and silvery or white, and forming irregular bands or
spots on elytra; bands or spots obscured when lighted from certain directions.
Third kind evenly dispersed among other kinds, dense, longer, fine, recurvate,
and brown. Ventral surface covered with very dense, fine, setigerous punctures;
setae whitish yellow, posteriorly directed, appressed, and fine. Ultimate visible
sternite with four rows of very long, coarse, brown, curved setae, each row
with four or five setae; setae not very obvious because of length of very dense
finer setae. Antenna clothed with long whitish setae; much longer setae
sparsely distributed and especially noticeable on apex of last article. Legs with
dense whitish setae. Tarsi with pair of course setae on apex of each article
except distal article. Sexual dimorphism not visible.

APRIL 1972]

SPILMAN — JUMPING SHORE BEETLE

115

Eyes with overall surfaces continuous with surfaces of head; dorsal separation
of eyes equal to 0.8 length of eye; anterior borders widely divergent ventrally
in anterior view and almost straight, posterior borders strongly curved;
dorsally and ventrally evenly arcuate. Antenna attaining middle point of procoxa;
with club of 4 articles; article 1 large, subglobular, article 2 long, 3 short,
3 to 7 becoming gradually broader, 8 to 10 broad, submoniliform, 11 elongate.
Pronotum with anterior border moderately incurved; anterior angles only
moderately projecting; lateral borders evenly arcuate; posterior border sinuate
on lateral thirds and with broad, posteriorly projecting, truncate lobe on middle
third. Elytra parallel-sided in anterior half, evenly arcuate in posterior half,
together forming semicircle at apex. Surface transversely evenly convex and
longitudinally weakly convex; pseudo pleural carina, or angulation of dorsal
surface with pseudopleuron, visible throughout its length in dorsal view, forming
lateral border of elytra; pseudo pleural carina with approximately 10 small
serrations on posterior half; apex of each elytron with small truncated projection
which lies medial to apical projections on ultimate sternite of abdomen; pseudo-
pleuron wide anteriorly, gradually narrowing posteriorly, visible throughout its
length.

Length: 1. 7-2.0 mm; width, 0.9-1. 1 mm.

Holotype, USNM No. 71706, Allotype, and 13 Paratypes, Tenacatita
Bay, Jalisco, Mexico, 19°16'45"N., 104°48'30 // W., W. G. Evans, 4
November 1966. Evans in a letter to me said, “. . . they hopped like
flea beetles. I found the beetles, aggregated in groups of 6 to 12, in
crevices of high tide-zone and splash-zone rocks on the protected
south end of Tenacatita Bay. These rocks were scattered irregularly
on a shingle beach. Collembola, as yet unidentified, were also found
on the surface of these rocks as well as in crevices, but algae were not
present.”

Acknowledgment. — Of course, this study would have been im-
possible without William George Evans. His scientific ability, his
generosity, and his gentlemanliness make working with him a pleasure.

Literature Cited

Evans, W. G. 1968. Some intertidal insects from western Mexico. Pan-Pac.
Entomol, 44(3) : 236-241.

Spilman, T. J. 1959. A study of the Thaumastodinae, with one new genus and
two new species (Limnichidae) . Coleopt. Bull., 13(4): 111-122.

1966. A new species of Martinius from Cuba (Coleoptera: Limnichidae).
Coleopt. Bull., 20(4) : 123-125.

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

A New Species of My strops from Costa Rica

(Coleoptera: Nitidulidae)

Lorin R. Gillogly

San Pedro, California 90731

Nitidulid beetles of the genus Mystrops Erichson are found in the
New World from Mexico south to Argentina. Not a great deal is
known of their life histories and food preferences. At the present
time three species are known definitely from feather-leafed palms,
where the larvae and adults are found in the blossoms feeding on
pollen. The beetles may occur in great numbers on the male in-
florescence. If disturbed, they fly around it like a swarm of gnats.

Bondar (1940) reported an agricultural pest, Mystrops fryi Grouvelle
(M. palmarum Bondar), in blossoms of Cocos coronata Mart, and
Cocos nucifera Linnaeus. From male flowers of the palm Cocos
romanzoffiana Chamille, he collected another species, Mystrops bondari
Gillogly.

The queen palm, known variously as Cocos plumosa Hook., or
Arecastrum romanzoffianum Beccari, is associated with a third species.
A large number of specimens of Mystrops heterocera Sharp were
collected with a long-handled net from blossoms of the queen palm
on 28 August 1965 by the author. The locality was Mexico, near
Acayucan on the highway across the Isthmus of Tehuantepec.

A new species of Mystrops infesting palm blossoms is herein described
and figured, with illustrations by G. M. Gillogly. Because of its
potential economic importance, the nitidulid beetle will merit further
observations and studies.

Mystrops costaricensis Gillogly, new species

Male. — (Fig. 1). Oblong, strongly convex, surface finely reticulate, moderately
shiny but surface obscured by rather dense, recumbent, golden pubescence,
color uniformly testaceous except for black coarsely faceted eyes. Scutellum
enclosed by somewhat darker triangular area when viewed at certain angles.

Head transverse, with dense, coarse, close lying golden pubescence obscuring
surface. Eyes coarsely faceted, prominent. Front nearly quadrate, only slightly
narrower in front of eyes than behind. Labrum (Fig. 4) very deeply bilobed,
indentation nearly attaining clypeus, lobes fairly evenly rounded. Mandibles
(Fig. 3) with outer margin rather evenly curved, tip almost sharp with inner
margin crenate; two teeth at base of inner margin of tip, somewhat obscured
by beard; molar area rasplike, with larger teeth toward tip and becoming
rather small at base. Maxillae (Fig. 5) with lacinia broad, truncate, inner
angle produced into definite spur, outer angle acute but not attenuate, tip

The Pan-Pacific Entomologist 48: 116-120. April 1972

GILLOGLY — A NEW MYSTROPS

117

APRIL 1972]

Figs. 1 and 2. My strops costaricensis. Fig. 1. Adult male. Fig. 2. Female.
Elytra tips attenuate.

heavily bearded; palpi with first segment minute, twice as wide as long,
second segment transverse, nearly elbowed, about three times as long and three
times as wide as first segment, third segment nearly square in outline being
somewhat narrower at tip than at base; terminal segment elongate, nearly twice
as long as first three segments together, widest at basal one-fourth narrowing
to tip and base. Labium (Fig. 6) with ligula spatulate; paraglossae broad,
obliquely truncate tips, strongly bearded; palpi, first segment minute, second
clavate, third thick, slightly sigmoidal; submentum bilobed distally with small
protuberance in middle of sinuation, strongly indented on each side near base.
Antennae short, reaching to mesosternum, pubescence fine with few erect hairs
on each segment; club distinct, of three equal segments. Pro thorax almost
rectangular, nearly twice as wide as long, anterior margin straight, a little
narrower than base; sides narrowly margined, hardly explanate, evenly arcuate

118

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

3 4

Figs. 3. and 4. Mystrops costaricensis. Adult male. Mouthparts. Fig. 3.
Mandible. Fig. 4. Labrum.

from apex to three quarters then strongly retracted to distinct obtuse angle;
base narrowly margined, lightly sinuate at each side of scutellum, more strongly
sinuate near hind angles which project slightly to rear; surface rather closely
moderately finely punctate with pubescence somewhat less dense than on head;
texture of surface smooth between punctures. Scutellum triangular, twice as
wide as long, surface distinctly finely reticulate. Elytra length to width as
1.3 to 1, sides parallel, tips rounded, sutural angles rounded, pubescence and
punctation more sparse than on prothorax. Pygidium convex, evenly curved,
strongly closely pubescent. Prosternum convex, sparsely finely punctate, lightly
pubescent, surface strongly finely reticulate, process narrow, depressed behind
coxae. Metasternum sparsely punctate, strongly finely reticulate; axillary space
large, extending along episternal suture nearly to hind coxae. Abdominal
segments of equal length, rather pubescent, surface reticulate. Supplementary
segment not visible from above, hardly visible from below but usually to be seen
from behind.

Female. — Punctation more sparse than male, pubescence finer. Tips of elytra
strongly attenuate (Fig. 2) . Pygidium tip simple. Antennae similar to those
of male.

Length. — 1.4 to 1.7 mm. Width: 0.6 to 0.8 mm.

Holotype male and allotype Guapiles, Limon, Costa Rica, altitude
300 m, 16 November 1968 are deposited in the California Academy
of Sciences. Paratypes are deposited in the collections of the University

APRIL 1972]

GILLOGLY — A NEW MYSTROPS

119

Figs. 5 and 6. Mystrops costaricensis. Adult male. Mouthparts. Fig. 5.
Maxilla. Fig. 6. Labium.

of Costa Rica; the Inter-American Institute of Agricultural Sciences
at Turrialba, Costa Rica; American Museum of Natural History; and
the United States National Museum.

Type and allotype and all 322 paratypes (159 males and 163
females) were taken with the assistance of Ingeniero Francisco Mata-
moros from the same male inflorescence of Elaeis guineensis Jacq.,
African oil palm, on the afternoon of 16 November 1968 at Guapiles
in the state of Limon. The oil palms are grown at the Experiment
Station Los Diamantes, a cooperative project of the Department of
Agriculture of Costa Rica with the A.I.D. program of the United
States.

Mystrops costaricensis falls in the key (Gillogly, 1955) at couplet
#3 with M. discoidea Murray from which it may be separated by
its short antennae. The antennae of M. discoidea are nearly as long
as the body in both sexes.

120

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

My visit to the station was arranged through the kindness of Dr.
Vargas of the Department of Agriculture of Costa Rica. I gratefully
acknowledge the counsel and assistance of the ecologists at the Tropical
Science Center of San Jose, Costa Rica: Joseph A. Tosi, Jr. and
Dr. L. R. Holdridge.

Literature Cited

Bondar, G. 1940. Notas Entomologicas da Bahia. V. Rev. Entomol. (Rio de
Janeiro), 11: 199-214.

Gillogly, L. R. 1955. A review of the genus My strops Erichson (Coleoptera,
Nitidulidae) . Rev. Brasil. Entomol., 3: 191-204.

BOOK NOTICES

The following bulletins have been published recently by the Research Division,
Virginia Polytechnic Institute and State University.

The Insects of Virginia:

No. 3. A systematic review of the genus Culicoides (Diptera: Ceratopogoni-
dae) of Virginia with a geographic catalog of the species occurring
in the eastern United States north of Florida. Frank V. Battle and
E. Craig Turner, Jr. Res. Div. Bull. 44, V.P.I. and State Univ., 129
p., 36 pis. 1971.

No. 4. Shield bugs (Hemiptera; Scutelleroidea, Scutelleridae, Corimelaenidae,
Cydnidae, Pentatomidae) . Richard L. Hoffman. Res. Div. Bull. 67,
V.P.I. and State Univ., 61 p., 16 figs. 1971.

Studies on the Morphology and Systematics of Scale Insects:

No. 3. Morphology and systematics of Cerococcus parrottii (Hunter) with
notes on its biology (Homoptera: Coccoidea: Asterolecaniidae) James
0. Howell, Michael L. Williams, and Michael Kosztarab. Res. Div.
Bull. 64, V.P.I. and State Univ., 23 p., 5 pis., 1 map. 1971.

No. 4. Morphology and systematics of the adult females of the genus
Lecanodiaspis (Homoptera: Coccoidea: Lecanodiaspididae) . James 0.
Howell and Michael Kosztarab. Res. Div. Bull. 70, V.P.I. and State
Univ., 248 p., 29 photos, 5 maps, 40 pis., + viii. 1972.

The above publications are available on an exchange basis from the Mailing
Room, Research Division, Hutcheson Hall, Virginia Polytechnic Institute and
State University, Blacksburg, Virginia 24061. — R. W. Thorp, University of
California, Davis, 95616.

APRIL 1972]

BROTHERS — A NEW THAUMALEA

121

A New Species of Thaumalea from California

(Diptera: Thaumaleidae)

Donald R. Brothers

Biological Sciences Department, San Jose State College, California 95114

The genus Thaumalea Ruthe is represented in North America by
five relatively rare species (Stone in Stone et al. 1965, Schmid 1970).
Of these, only one species, T. fusca (Garrett) , has been cited as possibly
occurring in California (Wirth and Stone in Usinger 1963). Cole
(1969) cites a specimen “near fusca ” from Bodie, Mono County,
California.

In an attempt to determine the identity of Thaumalea specimens
collected from Santa Clara County, California, it was discovered that
the terminalia of the male specimens differed considerably from those
of an unpublished illustration of the T. fusca holotype. The ninth
tergite of T. fusca is void of elongated ventral projections, and the
dististyles are without setose lobes or terminal elongated processes.
The following description of the undescribed species is published to
make the name available for taxonomic studies of this genus.

Thaumalea santaclaraensis Brothers, new species

(Figs. 1, 2)

Male. — Body length 2. 5-3.0 mm. Thorax brown. Abdomen brownish black.
Antennae 12 segmented; pedicels very large, with long setae; flagellar segments
1, 2, 3, 4, 6, and 8 each bearing long, stout seta. Tip of each terminal segment
of antennae also bears two long setae. Wing length 2.75-3.25 mm, uniform
light brown; uniformly covered with microtrichia ; veins brown; macrotrichia
on veins C, R, and Ri; 8-10 on vein R, and 10-12 on vein Ri. Halteres
elongate, yellow. Legs light brown, darkened distally. Ninth tergite brown,
heavily chitinized, about as long as broad, tapering and slightly round apically.
Medial ventral edges of ninth tergite produced inward on both sides, forming
elongated projections. Basistyles about as long as broad, sparsely and coarsely
setose. Dististyles about same length as basistyles; each with prominent setose
basal lobe; outer margin bearing two setae; apical end with stout tooth and ex-
panded, elongated, curved process. Parameres simple, rather slender; tips
slightly pointed inwards.

Female. — Body length 3-4 mm. Valves of genital plate setose and pointed;
tips each bearing two setae. Posterior corners of ninth tergite rounded. Other
structures as for male.

Holotype male and allotype, from Alum Rock Park, Santa Clara
County, California, elevation 900 feet, 31 July 1971, and 31 May
1971 respectively, (D. R. Brothers). Thirteen male and eight female

The Pan-Pacific Entomologist 48: 121-122. April 1972

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

2

Fig. 1 and 2. Thaumalea santaclaraensis Brothers, new species. Fig. 1. Male
terminalia, ventral view. Fig. 2. Female terminalia, lateral view.

paratypes were collected from the type locality during June and
July, 1971, (D. R. Brothers).

The holotype male and allotype female will be deposited in the
California Academy of Sciences, San Francisco, California; paratypes
at the California Academy of Sciences, and the U. S. National Museum,
Washington, D. C.

Terminalia illustrations are of mounted paratypic material.

Acknowledgments

I thank Dr. P. H. Arnaud, Jr. of the California Academy of
Sciences for providing an illustration of the male terminalia of the
T. fusca holotype.

Literature Cited

Cole, F. R. 1969. The flies of western North America. Univ. Calif. Press,
Berkeley, 693 pp.

Schmid, F. 1970. Quelques Thaumaleides nouvelles ou peu connues (Diptera,
Thaumaleidae) . Natur. Can., 97 : 499-510.

Stone, A. 1965. Family Thaumaleidae. p. 120. in Stone, A., et al. 1965. A
Catalog of the Diptera of America, North of Mexico. U. S. Dep.
Agr., Agr. Handb. 276., 1,696 pp.

Wirth, W. W. and A. Stone. 1956. Chapter 14. Aquatic Diptera. pp. 372-482.

in Usinger, R. L. (ed.) . 1956. Aquatic insects of California. Univ.
Calif. Press, Berkeley, 508 pp.

APRIL 1972]

COHEN & ALLEN NEW BAETODES

123

New Species of Baetodes from Mexico and Central America

(Ephemeroptera: Baetidae)

Sandra D. Cohen and Richard K. Allen 1

California State College, Los Angeles, 90032

The genus Baetodes , described by Needham and Murphy (1924), was
based upon nymphs of two species collected in Brazil. Baetodes
serratus was described, named, and designated as the genotype, and
the other was described and reported as Baetodes Nymph No. 1.
Traver (1943) reported the adult stage of the genus when she
described B. spiniferum from a male imago and a female subimago
collected in Venezuela. In this same paper, she reported an undescribed
species of Baetodes from Mexico, and in 1944, published additional
records of B. serratus from Brazil. Edmunds (1950) reported a
nymphal record of the genus from Texas, and a second Mexican
record from Culinevera (Cuernavaca? ) , south of Mexico City. Demoulin
(1955) described B . itatiayanus from nymphs collected in Brazil, and
Packer (1966) was the first to report the genus from Central America
when he published records from twelve localities in Honduras. Mayo,
in 1968, described two additional species, B. levis and B. spinae ,
from nymphs collected in Ecuador.

This report includes descriptions and names for nymphs of nine
species of Baetodes collected in Mexico, Guatemala, El Salvador, and
Honduras, and the number of named species now stands at fourteen.
The characters most useful in distinguishing the species of Baetodes
in the nymphal stage are: (1) the presence or absence, the number,
and the degree of development of the median tubercles on the pronotum,
metanotum, and abdominal terga; (2) the presence or absence of
coxal gills; (3) the color on the abdominal gills; and (4) the color
on the thoracic and abdominal segments. Coxal gills are of limited
taxonomic value. In some species, they are always present, in others
they are always absent, but in others they may be present or absent.

Baetodes adustus Cohen and Allen, new species

Nymph. — Length: body 5. 0-6.0 mm; caudal filaments 6.0-7.0 mm. General
color brown with darker brown, red, reddish brown and pale markings. Head
brown with darker brown markings ; occiput with sparse setae. Thoracic nota brown
with darker brown markings; pronotum with a dorsal, median, posterior
elevation ; metanotum with moderately developed dorsal, median, posterior tubercle

^^The research upon which this report is based was supported by National Science Foundation.

The Pan-Pacific Entomologist 48: 123-135. April 1972

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

(Fig. 1) ; thoracic nota with sparse setae; thoracic sterna pinkish red; legs
brown with pale markings; femora brown with variable pale markings; tibiae
brown with thin, pale longitudinal streak; tarsi brown with pale markings and
brown at apices; coxae with finger- like gills; tarsal claws with 7 denticles.
Abdominal terga usually brown to reddish brown, color often darker along
anterior margin of segments; abdominal terga 1-8 with median tubercle;
tubercles moderately developed on terga 1-7, small on tergum 8 (Fig. 1) ; abdominal
terga with sparse setae; abdominal gills pale, often tinged with pinkish red;
abdominal sterna brown with reddish brown and pale markings; abdominal
sterna 1-6 brown, suffused with red and with reddish brown lateral markings;
sterna 7-10 brown with pale median macula, and reddish brown lateral markings.
Caudal filaments light brown.

Holotype , mature female nymph, Stream 5 mi. S. Ciudad Mendoza,
Vera Cruz, Mexico, 7 November 1968, R. K. Allen. Paratopotypes:
2 male and 3 female nymphs, same data as holotype, 1 male and 1
female in collection University of Utah, Salt Lake City, others in
collection California State College at Los Angeles. Paratype: 1 female
nymph, Rio Jamapa, 32 mi. N. E. Coscamatipec, Vera Cruz, Mexico,
14 July 1966, R. K. Allen, in collection California State College at
Los Angeles.

Remarks. — Baetodes adustus is superficially similar to B. pallidus
n. sp. from Mexico and Honduras. Both species bear median abdominal
tubercles on segments 1-8, a moderately developed metathoracic
tubercle and only a small elevation on the posterior margin of the
pronotum, and the coxae with finger-like gills. Mature nymphs of
B. adustus are longer than those of B. pallidus , and the abdominal
terga are brown to reddish brown and without pale markings (Fig. 1) .

Baetodes caritus Cohen and Allen, new species

Nymph. — Length: body 3.0-4.0 mm; caudal filaments 4.0-5. 0 mm. General
color brown with dark brown, reddish, and pale markings. Head brown, sutures
pale and pale around compound eyes; occiput with sparse setae. Thoracic nota
brown; pronotum without dorsal, median, posterior elevation; metanotum with
barely discernible dorsal, median, posterior elevation; thoracic nota with sparse
setae; thoracic sterna reddish with numerous pale dots; legs brown with pale
markings; femora brown with pale median and basal maculae (Fig. 2) ; tibiae
light brown to brown; tarsi pale, brown at apices; coxae with small finger-like
gill; tarsal claws with 8 denticles. Abdominal terga brown with dark brown
and reddish brown markings; abdominal terga 1-3 usually brown; terga 4-9
often brown with dark brown anterior margins; terga 6-9 usually brown with pale
median longitudinal stripe; tergum 10 light brown; terga 1-3 each with barely
discernible, median elevation; abdominal terga with sparse setae; abdominal
gills pale; abdominal sterna pale to light brown with reddish brown lateral
markings; sternum 1 reddish with paired large submedian white dots; sterna

APRIL 1972]

COHEN & ALLEN NEW BAETODES

125

Fig. 1. Baetodes adustus, mature male nymph, dorsal view

2-6 pale to light brown with lateral reddish brown markings, with white dots;
sterna 7-10 light brown (Fig. 6) . Caudad filaments light brown.

Holotype , mature female nymph, Rio Latoma at km 181 ON Hwy.
2, Guatemala, 24 October 1968, R. K. Allen. Paratopotype: 1 male

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

nymph, same data as holotype, in collection California State College
at Los Angeles. Paratypes: 1 female nymph, Rio Hondura at Ciudad
Carrita, 5 mi. E. El Salvador/Honduras border, Honduras, 28 October
1968, R. K. Allen; 1 female nymph, Rio Mizata, 27 mi. W. La Libertad,
El Salvador, 28 October 1968, R. K. Allen; 1 female nymph, Rio
Sesecapa near Mapastepec on Hwy. 200, Chiapas, Mexico, 20 July 1966,
R. K. Allen; 1 female nymph, Stream 7 mi. N. Arriago, on Hwy. 190,
Chiapas, Mexico, 20 July 1966, R. K. Allen; 1 female nymph, Rio
Papagayo on Hwy. 95, Guerro, Mexico, 16 November 1968, R. K.
Allen; 1 male nymph, Rio Amacuzac at Huajintlan on Hwy. 95,
Morelos, Mexico, 14 November 1968, R. K. Allen; 3 female nymphs,
Rio San Antenco at Tonala, Chiapas, Mexico, 1 November 1968, R. K.
Allen; 2 male and 1 female nymph, Rio Piedras Negras at Piedras
Negras near Poza Rica, Vera Cruz, Mexico, 12 November 1968, R. K.
Allen; 1 female nymph, Rio Hondura at Ciudad Carrita, ca. 5 mi. E.
El Salvador, 28 October 1968, R. K. Allen; above paratypes in
collection California State College at Los Angeles; 1 female nymph,
Escula Agricola Panamericana, Dept. El Paraiso, Honduras, 26 October
1964, J. S. Packer; 1 male and 1 female nymph, Stream 8 km. E.
Danli, El Paraiso, Honduras, 29 August 1964, J. S. Packer; above
paratypes in collection University of Utah, Salt Lake City.

Remarks. — Baetodes caritus appears to be most closely related to
B. deficiens n. sp. Both species have reduced median abdominal
tubercles, and similar dorsal abdominal color patterns. Baetodes caritus
is distinguished from the latter species by the following characters:
(1) femora brown with pale maculae (Fig. 2) and (2) thoracic
sterna reddish with numerous pale spots (Fig. 6) .

Baetodes deficiens Cohen and Allen, new species

Nymph. — Length: body 3. 5-4.5 mm; caudal filaments 5.0-6.0 mm. General
color light brown to pale with dark brown markings. Head light brown,
darker brown around median ocellus; occiput with sparse setae. Thoracic
nota light brown with brown markings; mesonotum usually with brown transverse
marking across anterior margin; pronotum without median posterior elevation
or tubercle; metanotal tubercle barely discernible; thoracic nota with sparse
setae; thoracic sterna pale; legs pale with brown markings; coxae usually with
gills; femora pale with brown basal, median, and usually subapical maculae,
and with dark brown to black marking at each apex (Fig. 3) ; tibiae and tarsi
pale; tarsal claws with 7 denticles. Abdominal terga brown with dark brown,
reddish brown, and pale markings; abdominal terga 1-3 reddish brown with
pale to light brown posterior margins; terga 4-6 with dark brown to reddish
brown markings at anterior margin ; terga 7-9 brown, usually with dark
brown anterior spots; tergum 10 pale; terga 1-3 each with barely discernible

APRIL 1972]

COHEN & ALLEN NEW BAETODES

127

Figs. 2-3. Left femora of Baetodes nymphs. Fig. 2. B. caritus. Fig. 3. B.
deficiens. Fig. 4, Baetodes adustus, tarsal claw. Fig. 5, Baetodes adustus,
nymphal caudal filaments. Figs. 6-12. Abdomens of Baetodes nymphs. Fig. 6.
B. caritus, ventral view. Fig. 7. B. deficiens, lateral view. Fig. 8. B. fuscipes,
dorsal view. Fig. 9. B. fuscipes, lateral view. Fig. 10. B. inermis, dorsal view.
Fig. 11. B. inermis, lateral view. Fig. 12. B. inermis, ventral view.

128

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

median elevation (Fig. 7) ; terga with long setae; abdominal gills pale;
abdominal sterna pale with brown markings; sterna 1-7 usually with sub-
median anterior transverse streaks; sterna 6-9 with sublateral longitudinal
streaks; sternum 10 pale. Caudal filaments pale.

Holotype , mature female nymph, Rio Clarrita at San Morano on
Hwy. 4, Dept. El Paraiso, Honduras, 29 October 1968, R. K. Allen.
Paratypes: 1 male and 4 female nymph, Rio Amacuzac at Huajintlan
on Hwy. 95, Morelos, Mexico, 14 November 1968, R. K. Allen; 1
male and 2 female nymphs, Rio Choluteca, Tegucigalpa, Honduras, 29
October 1968, R. K. Allen ; 1 female nymph, Rio San Marcos at Apapan-
tilla 8 mi. S. E. Villa A. Camacho, Vera Cruz, Mexico, 12 November 1968,
R. K. Allen; 2 male and 3 female nymphs, Rio Piedras Negras at
Piedras Negras near Poza Rica, Vera Cruz, Mexico, 12 November 1968,
R. K. Allen; 2 male and 1 female nymphs, Rio Papagpyo near Tierra
Colorado, Guerrero, Mexico, 16 November 1968, R. K. Allen; 1 male
and 1 female nymph, Stream 9 mi. S. Rio Grande de Santiago, Jalisco,
Mexico, 17 October 1968, R. K. Allen; above paratypes in collection
California State College at Los Angeles; 7 female nymphs, Dept. El
Paraiso, Escula Agricola Panamericana, Rio Yeguace, Honduras, 26
October 1964, J. S. Packer; 1 female nymph, Dept. Choluteca, Stream
near Choluteca on Pan Americana Hwy., Honduras, 10 October 1964,
J. S. Packer, above paratypes in collection University of Utah, Salt
Lake City.

Remarks. — Baetodes deficiens is distinguished from B. caritus , an
apparently closely related species, and all other described species of
Baetodes by the following combination of characters: (1) abdominal
terga without obvious tubercles, and with only raised elevations of
segments 1—3; (2) femora pale with brown maculae (Fig. 3) ; and

(3) thoracic sterna pale.

Baetodes fuscipes Cohen and Allen, new species

Nymph. — Length: body 3.0-4.0 mm; caudal filaments 3.0-4.0 mm. General
color brown and reddish brown with pale and reddish markings. Head brown
with irregular dark brown markings; occiput with sparse setae. Thoracic nota
brown with dark brown markings; pronotum with moderately well-developed
median posterior elevation; metanotum with moderately developed median
tubercle; thoracic nota with sparse setae; thoracic sterna reddish with numerous
pale spots; coxae without gills; legs brown with pale and dark brown markings;
femora brown with pale L-shaped marking, and dark brown at apices; tarsal
claws with 7 denticles. Abdominal terga reddish brown with pale margins;
terga 1-3 dark brown; terga 4-8 brown; terga 9-10 light brown (Fig. 8) ;
terga 1-7 with median tubercles; terga 1-6 with moderately developed tubercles;
terga 7 with poorly developed tubercles (Fig. 9) ; terga with sparse setae;

APRIL 1972]

COHEN & ALLEN — NEW BAETODES

129

abdominal gills pale; abdominal sterna reddish with brown to pale markings
and pale spots; sterna 1-4' reddish with numerous pale spots; sterna 5-9
reddish, brown to pale medially; sternum 10 pale. Caudal filaments light brown.

Holotype , male nymph, Stream 5 mi. S. Ciudad Mendoza, Vera
Cruz, Mexico, 7 November 1968, R. K. Allen. Paratypes: 2 female
nymphs, Rio Jamapa 3 mi. N. E. Coscomatipec, Vera Cruz, Mexico,
8 November 1968, R. K. Allen, in collection California State College
at Los Angeles; 1 female nymph, 10 mi. W. Juticalpa, Rio Julicalpa,
Dept. Olancho, Honduras, 6 November 1964, J. S. Packer, in collection
University of Utah, Salt Lake City.

Remarks. — The nymph of B. fuscipes is similar to B. adustus and B.
pallidus n. sp. in the number and development of the median, dorsal,
abdominal tubercles, and in the development of the metanotal tubercle.
The former species differs as the posterior pronotal tubercle is well
developed and the coxae are without gills. This species is further
distinguished by the color characters of the abdominal terga.

Baetodes inermis Cohen and Allen, new species

Nymph.— Length 4.0-5.0 mm; caudal filaments 5.5-6.5 mm. General color
light brown to brown with darker brown and reddish brown markings. Head
light brown to brown with darker brown markings; head often with dark
brown band between ocelli; occiput with sparse setae. Thoracic nota brown
with dark brown markings; pronotum without posterior median elevation;
metanotum with poorly developed median tubercle; thoracic nota with sparse
setae; thoracic sterna pale usually with dark circular lines around bases of
legs; legs pale with brown markings; femora pale with brown markings; tibiae
and tarsi pale, often brown at apices; coxae with or without gills; tarsal claws
with 7 denticles. Abdominal terga pale to brown with dark brown and reddish
brown markings; terga 1-6 usually pale with brown to reddish brown anterior
markings triangular in shape on posterior segments; terga 7-9 brown with
dark brown anterior markings; terga 5-9 often with disjunct transverse reddish
brown triangular marking; tergum 10 brown (Fig. 10) ; terga 1-7 with poorly
developed median tubercle; tubercles of segments 1-6 small, tubercles barely
discernible on segment 7 (Fig. 11) ; terga with sparse setae; abdominal gills pale;
abdominal sterna pale with reddish brown markings; sterna 1-7 usually with
thin transverse anterior reddish brown marking ; sternum 8 often with faint
transverse marking; sterna 9-10 usually pale (Fig. 12). Caudal filaments pale.

Holotype , mature female nymph, Rio San Marcos near Ciudad
Victoria, Tamaulipas, Mexico, 24-25 November 1968, R. K. Allen.
Paratopotypes: 12 male and 45 female nymphs, same data as holotype,
1 male and 3 female nymphs in collection of University of Utah, Salt
Lake City; California Academy of Sciences; and Canadian National
Collection, Ottawa; remainder in collection of California State College
at Los Angeles. Paratypes: 20 male and 32 female nymphs, Rio

130

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

Amacuzac at Huajintlan on Hwy. 95, Morelos, Mexico, 14 November
1968, R. K. Allen; 1 female nymph Stream 15 mi. N. Ayoquezco,
Oaxaca, Mexico, 20 October 1968, R. K. Allen; 1 male nymph, Rio
Grande 3 mi. S. Gualatao, Oaxaca, Mexico, 6 November 1968, R. K.
Allen; 14 male and 16 female nymphs, Rio La Pasion at Tizapan
El Alto, Jalisco, Mexico, 16 October 1968, R. K. Allen; 1 female nymph,
Rio Atoyac at Ayoquezco, Oaxaca, Mexico, 22 October 1968, R. K.
Allen; 1 female nymph, stream at Hacienda Guadalupe, Jalisco,
Mexico, 17 October 1968, R. K. Allen; 1 female nymph, Rio Papagayo
on Hwy. 95, Guerrero, Mexico, 16 November 1968, R. K. Allen, above
paratypes in collection California State College at Los Angeles.

Remarks. — Baetodes inermis is readily distinguished from all de-
scribed species of the genus by the distinctive color pattern on the
abdominal terga, and by the thin transverse lines across the anterior
margin of the abdominal sterna.

Baetodes noventus Cohen and Allen, new species

Nymph. — Length: body 3. 5-4.5 mm; (caudal filaments broken). General
color brown to reddish brown with dark brown and reddish markings. Head
unicolorous brown ; occiput without setae. Thoracic nota brown with dark
brown markings; pronotum with well developed dorsal, median tubercle;
metanotum with a well developed dorsal median tubercle (Fig. 17) ; thoracic
nota without setae; thoracic sterna light brown, suffused with red; legs
light brown with pale and dark brown markings; femora brown with pale basal
and apical maculae, and dorsal margin and apices dark brown; tibiae and
tarsi light brown, apices of tarsi brown; coxae with finger- like gills; tarsal
claws with 6 denticles. Abdominal terga brown to reddish brown, middle
segments often pale with brown markings; abdominal terga 1-9 with median
tubercle; terga 1-7 with well developed tubercles, tubercles moderately developed
on terga 8-9 (Fig. 16-17) ; abdominal terga without setae; abdominal gills
pale, often suffused with red; abdominal sterna pale, suffused with red and
with brown markings; sterna 1-5 pale, suffused with red; sterna 6-9 pale with
sublateral brown markings; sternum 10 pale. (Caudal filaments broken).

Holotype, mature female nymph, Rio Mizata, 27 mi. W. La Libertad,
El Salvador, 28 October 1968, R. K. Allen. Paratypes: 1 female
nymph, Rio Clarrita at San Morano on Hwy. 4, Dept. El Paraiso,
Honduras, 29 October 1968, R. K. Allen; 1 female nymph, Rio
Latoma at km. 182 on Hwy. 2, Guatemala, 24 October 1968, R. K.
Allen, above paratypes in collection California State College at Los
Angeles; 1 female nymph, Campamento Galera at bridge on Hwy. 3,
Dept. Olancho, Honduras, 7 November 1964, J. S. Packer; 1 male and
1 female nymph, 38 km E. Zamorana on Hwy. 4, Dept. El Paraiso,
Honduras, 31 October 1964, J. S. Packer; 1 male nymph, Stream

APRIL 1972]

COHEN & ALLEN NEW BAETODES

131

Figs. 13-15. Left femora of Baetodes nymphs. Fig. 13. B. pallidus. Fig. 14.
B. pictus. Fig. 15. B. tritus. Figs. 16-23. Abdomens of Baetodes nymphs.
Fig. 16. B. noventus, dorsal view. Fig. 17. B. noventus, lateral view. Fig. 18.
B. pallidus, dorsal view. Fig. 19. B. pallidus, lateral view. Fig. 20. B. pictus,
dorsal view. Fig. 21. B. pictus, lateral view. Fig. 22. B. tritus, dorsal view.
Fig. 23. B. tritus, lateral view.

132

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

E, Guaimaca on Hwy. 3, Dept. Francisco Morazan, Honduras, 6
November 1964, J. S. Packer, above paratypes in collection University
of Utah, Salt Lake City.

Remarks. — The nymph of Baetodes noventus is distinguished from
all described species of Baetodes by the following combination of
characters; (1) abdominal terga with median tubercles on segments
1-9; (2) well developed median, dorsal, tubercles on the pronotum,

metanotum, and abdominal segments 1-7; and (3) coxae with gills.

Baetodes pallidus Cohen and Allen, new species

Nymph. — Length: body 3. 5-4.5 mm; (caudal filaments broken). General
color light brown and reddish brown and black marldngs. Head light brown
with pale transverse band between compound eyes and reddish brown transverse
band between lateral ocelli; occiput with sparse setae. Thoracic nota light
brown with brown markings; pronotum with small dorsal, median elevation;
mesonotum with reddish brown transverse marking across anterior margin ;
metanotum with moderately developed dorsal, median tubercle; thoracic nota
with setae; thoracic sterna pale; legs light brown with pale and brown
markings; femora light brown with pale basal macula, elongate median macula,
and apical dark brown mark (Fig. 13) ; tibiae and tarsi light brown, apices
of tarsi brown; coxae usually with finger- like gills; tarsal claws with 6
denticles. Abdominal terga reddish brown with black and pale markings;
terga 1-3 reddish brown with pale median macula, and darker reddish brown to
black posterior margin ; terga 4<-9 pale with black transverse markings along
anterior and posterior margins; tergum 10 usually pale (Fig. 18) ; abdominal
terga 1-8 with a median tubercle; terga 1-7 with moderately developed tubercles;
tubercle barely discernible on tergum 8 (Fig. 19) ; abdominal terga with sparse
setae; abdominal gills pale; abdominal sterna light brown. (Caudal filaments
broken) .

Holotype, mature male nymph, Rio Clarrita at San Morano on
Hwy. 4, Dept. El Paraiso, Honduras, 29 October 1968, R. K. Allen.
Paratypes: 1 male nymph, Stream 7 mi. N. Arriaga on Hwy. 190,
Chiapas, Mexico, 20 July 1966, R. K. Allen; 1 female nymph, Stream
9 mi. S. Rio Grande de Santiago, Jalisco, Mexico, 17 October 1968,
R. K. Allen, above paratypes in collection California State College at
Los Angeles; 1 female nymph, Stream 10 mi. E. Guainaca on Hwy. 3,
Dept. Fransico Morazan, Honduras, 6 November 1964, J. S. Packer;
1 female nymph, Stream 38 km E. Zamorano on Hwy. 4, Dept. El
Paraiso, Honduras, 29 August 1964, J. S. Packer; 1 female nymph,
Escula Agricola Panamericana, El Paraiso, Honduras, 26 October
1964, J. S. Packer, above paratypes in collection University of Utah,
Salt Lake City.

Remarks. — This species has many characters in common with
Baetodes adustus but is distinguished from it by size and color.

APRIL 1972]

COHEN & ALLEN NEW BAETODES

133

Mature nymphs of B. pallidus are small (3. 5 — 4.5 mm in length),
and the abdominal terga have distinctive reddish brown, black and
pale markings.

Baetodes pictus Cohen and Allen, new species

Nymph. — Length: body 5.5-6.5 mm; caudal filaments 6.0-7. 0 mm. General
color brown with dark brown, yellow, and reddish brown markings. Head
brown, frontoclypeal area pale; occiput with sparse setae. Thoracic nota
brown with dark brown markings; pronotum with a posterior median elevation;
metanotal tubercle small; thoracic nota with dense setae; thoracic sterna
pale; coxae without gills; legs brown with pale markings; femora brown with
large pale median marking (Fig. 14) ; tibiae and tarsi brown, tarsi brown at
apices; tarsal claws with 6 denticles. Abdominal terga brown with reddish
brown markings; terga 1-9 brown with broad reddish brown anterior transverse
band, narrow reddish brown posterior transverse band, and reddish brown median
macula; tergum 10 brown (Fig. 20) ; abdominal terga 1-7 with poorly developed
median tubercle; tubercles barely discernible on terga 8-9 (Fig. 21) ; setae
moderately dense; abdominal gills pale; abdominal sterna yellow with pale
markings; sternum 1 pale; sterna 2-9 yellow with pale margins, posterior
segments often with pale median longitudinal stripe; sternum 10 pale. Caudal
filaments yellowish.

Holotype, nearly mature female nymph, Stream 5 mi. S. Ciudad
Mendoza, Vera Cruz, Mexico, 7 November 1968, R. K. Allen.

Remarks. — Baetodes pictus nymphs are superficially similar to
those of B. noventus and B. pallidus. They are distinguished from B.
noventus as the body tubercles are smaller in size, and the coxae are
without gills. They are distinguished from B. pallidus as the middle
abdominal terga bear a dark median macula, and the abdominal sterna
are yellow.

Baetodes tritus Cohen and Allen, new species

Nymph. — Length: body 6. 0-8.0 mm; caudal filaments 8.0 10.0 mm. General
color light brown with brown and reddish brown markings. Head light brown
with irregular brown markings; frontoclypeal area pale; occiput with setae.
Thoracic nota light brown with brown and pale markings; pronotum without
dorsal, median, posterior elevation or tubercle; mesonotum with moderately
developed dorsal, median, posterior elevation ; metanotum with well developed
tubercle (Fig. 23) ; thoracic nota with setae; thoracic sterna pale to light
brown; legs light brown with reddish brown markings; femora light brown
with variable reddish brown markings; femora often with large subbasal reddish
brown macula and submarginal reddish brown streak near dorsal margin (Fig.
15) ; tibiae and tarsi light brown and suffused with reddish brown; coxae without
gills; claws with 7-8 marginal denticles. Abdominal terga light brown to brown
with reddish brown markings; terga 1-9 each with reddish brown marking
across anterior margin of segment; abdominal marking often absent on posterior

134

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

segments, and marking often disjunct in young specimens; abdominal terga
1-10 with a well developed median tubercle (Fig. 22-23) ; abdominal terga with
setae; abdominal gills reddish with pale margins; abdominal sterna light brown
with dark brown paired sublateral longitudinal streaks. Caudal filaments brown.

Holotype , male nymph, Rio Tecolapan near Santiago Tuxtla
on Hwy. 180, Vera Cruz, Mexico, 16 July 1966, R. K. Allen. Para-
types: 17 male and 16 female nymphs, Rio San Marcos near Ciudad
Victoria, Tamaulipas, Mexico, 24-25 November 1968, R. K. Allen,
1 male and 1 female nymph in each of the following collections:
University of Utah, Salt Lake City; California Academy of Sciences;
Canadian National Collection, Ottawa; V. K. Mayo, Tucson, Arizona;
remainder in collection of California State College at Los Angeles. 1
male nymph, Stream at Santa Isabel, 12 mi. above Arriaga on Hwy.
190, Chiapas, Mexico, 23 October 1968, R. K. Allen; 4 female nymphs,
Rio Latoma at km 182 on Hwy. 2, Guatemala, 24 October 1968, R. K.
Allen; 2 female nymphs, Rio Amacuzac at Huajintlan on Hwy. 95,
Morelos, Mexico, 14 November 1968, R. K. Allen; 1 male nymph, Stream
at Hacienda Guadalupe, Jalisco, Mexico, 17 October 1968, R. K. Allen,
1 male and 1 female nymphs, Rio Sesecapa near Mapastepec on Hwy.
200, Chiapas, Mexico, 20 July 1966, R. K. Allen; 11 male and 14
female nymphs, Stream 9 mi. S. Rio Grande de Santiago, Jalisco,
Mexico, 17 October 1968, R. K. Allen, above paratypes in collection
California State College at Los Angeles; 1 female nymph, Stream 8
km El Paraiso, Honduras, 29 August 1964, J. S. Packer, in collection
University of Utah, Salt Lake City.

Remarks. — Baetodes tritus appears to be the most widely distributed
Mexican Baetodes as it is known from central Mexico to Honduras.
This species is also the most easily recognized as the abdominal terga
bear well developed median tubercles on segments 1-10.

Acknowledgments

We thank G. F. Edmunds, Jr., University of Utah, for the loan of
the J. S. Packer collection, and Jerry Battagliotti, California State
College, Los Angeles, for preparing the included illustrations. Type
specimens are deposited in the California Academy of Sciences.

Literature Cited

Demoulin, G. 1955. Une Mission biologique Beige au Bresil Ephemeropteres.

Bull. Inst. Roy Sci. Natur. Belg., 31: 1-32.

Edmunds, G. F., Jr. 1950. New records of the mayfly genus Baetodes, with
notes on the genus. Entomol. News, 61: 203-205.

APRIL 1972]

COHEN & ALLEN NEW BAETODES

135

Mayo, V. K. 1968. Two new species of the genus Baetodes from Ecuador. Pan-Pac.
Entomol., 44: 251-257.

Needham, J. G. and H. E. Murphy. 1924. Neotropical mayflies. Bull. Lloyd
Libr., 24: 1-79.

Packer, J. S. 1966. A preliminary study of the mayflies of Honduras. Ceiba,
12 : 1 - 10 .

Traver, J. R. 1943. New Venezuelan mayflies. Bol. Entomol. Venez., 2: 79-98.
1944. Notes on Brazilian mayflies. Bol. Mus. Nac. Rio de Janeiro, Zool.,
22: 1-53.

SOCIETY ANNOUNCEMENT

The Pacific Coast Entomological Society announces the publication
of a new book in its Memoir Series: Volume 4: Robert Leslie
Usinger: Autobiography of an Entomologist.

This unique book, written by one of the outstanding entomologists
of our time, relates personal experiences and episodes from boyhood
through his professional career. Included are accounts of research
expeditions to various countries.

Edited by Drs. E. G. Linsley and J. L. Gressitt, the cloth-bound
book consists of 343 pages, 38 illustrations, and dust jacket. The
book should be available by 15 August 1972, price $15.00' (postage
extra and 5% state sales tax to be added for California residents).
Members of the PCES will receive a 20% discount.

Orders for this limited edition (1,000 copies) may be placed with:

Dr. Paul H. Arnaud, Jr., Treasurer
Pacific Coast Entomological Society
c/o California Academy of Sciences
Golden Gate Park
San Francisco, California 94118

Please make checks payable to the Pacific Coast Entomological Society.

136

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

A Review of the Acantliametropodinae with a Description

of a New Genus

(Ephemeroptera: Siphlonuridae)

George F. Edmunds, Jr. and Richard W. Koss 1

University of Utah, Salt Lake City, 84112, and Johns Hopkins University, Baltimore

The subfamily Acanthametropodinae is one of the least known mayfly
groups and a reasonable understanding of the taxon is available now
for the first time. The subfamily was established by Edmunds (in
Edmunds, Allen and Peters, 1963) for Acanthametropus and an un-
described genus. These forms were known only as nymphs. Siphluriscus
chinensis Ulmer (1920) was the first acanthametropodine to be de-
scribed, and it remains known only from the six adult specimens from
China used in the original description. Acanthametropus nikolskyi
Tshernova (1948) was described from an immature nymph from the
Amur River. Burks (1953) described Acanthametropus pecatonica (as
Metreturus ) from two nymphs collected in Illinois in 1926 and 1927.
Metreturus was erected as a new genus only because Tshernova’s paper
was then unknown to American mayfly workers. Edmunds and Allen
(1957) synonymized Metreturus Burks with Acanthametropus Tsher-
nova. Both Tshernova and Burks placed Acanthametropus as allied
to Ametropus, but Edmunds and Traver (1954) and Edmunds and
Allen (1957) placed the genus in the Siphlonuridae. Edmunds, Allen
and Peters (1963) recorded Acanthametropus from the Savannah
River, Georgia-South Carolina, based on three nymphs. Comparison
of the three Savannah River and two Illinois specimens does not
allow a decision as to whether or not there is more than one species
in the United States.

Tshernova (1967) described the Lower Jurassic Stackelhergisca
from a well preserved nymph from Siberia and an associated wing
fragment. Both have characters of the extant acanthametropodines.
The fossil nymph indicates that the legs are all directed posteriorly
and the mouthparts are for carnivorous feeding. The fork of MP is
deep and asymmetrical in the wing fragment; this is a character of
Acanthametropodinae, but it is not exclusive to that subfamily.

The genus Analetris, described below, was found first in 1947 as
a partial nymph in the stomach of a fish collected from the Green
River at Hideout Canyon (Edmunds, 1957). One nymph subsequently

-'■This study was supported by a grant from the National Science Foundation.

The Pan-Pacific Entomologist 48: 136-144. April 1972

APRIL 1972] EDMUNDS & KOSS — ACANTHAMETROPODINAE

137

was collected by G. R. Smith and G. G. Musser from the Green River
at Buckboard Flats, Sweetwater Co., Wyoming in July 1959. Since
1947 many hours were expended in an effort to collect additional
specimens from the Green River and its larger tributaries. In 1962-63
the Green River was inundated with water stored behind Flaming
Gorge Dam and hope for additional specimens dimmed. In 1968 the
junior author was successful in locating a population in a stretch of the
Blacks Fork River from near Granger, Wyoming, to the crossing of
the Blacks Fork River at Interstate Highway 80, about 7 miles east of
Little America. He was able to rear one subimago $ which enabled
us to recognize that Siphluriscus was an Acanthametropodinae. It is
not known how many miles of the Blacks Fork River support Analetris,
but it is relatively short because in many summers the river is almost
dry before it reaches Flaming Gorge Reservoir. Most of the river is
not readily accessible, but it is in increasing danger from oil well
drilling, agricultural development, dam building and pollution. In
fact, five additional collecting attempts have produced no more spec-
imens. Lehmkuhl (1970) collected a series of nymphs of the same
species in the South Saskatchewan River. This river is also in danger
from pollution.

In the Acanthametropodinae, then, there are about 55 known spec-
imens of 4 genera, of which 7 are imagos and subimagos. Analetris
is the best represented genus but its known habitats are endangered.
More than 40 miles of the Green River where Analetris presumably
occurred have already been submerged behind Flaming Gorge Dam.
The failure to collect more specimens is a result of a habitat in which
it is difficult to collect, the shy and extremely fast swimming behavior
of the nymphs, and inadequate collecting techniques.

Subfamily Acanthametropodinae

Adult. — Tornus of forewings near mid length; vein MP deeply forked; anal
field elongate. Hind wings % or more as long as fore wings; vein MP deeply
forked, base less than 14 as long as fork. Fore tarsi of $ 2% to 3 times
as long as tibiae. Subgenital plate of $ deeply excavated medially. Terminal
filament at least 2 mm long.

Nymph. — Mouthparts of carnivorous type; mandibles with long sharp incisors;
maxillae with fang-lilce spines; third segment of labial palpi reduced. Legs
directed posteriorly when dead (or when swimming). Fore and middle pairs
of legs with femora broadest in basal one-third; tarsi ca. % as long as tibiae;
claws long, varying from % to 1% times as long as tarsi; hind legs with margins
of femora subparallel, femora Yq or less as wide as long; claws longer than
tibiae and tarsi combined; tibiae, tarsi and claws bowed inward. Posterolateral
projections present on abdominal segments 1-9. Cerci with long setae on

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mesal margins; terminal filament with long setae on both lateral margins.

Keys to the two genera known as adults and two known as nymphs
are not necessary. Adults of Analetris have the three caudal filaments
subequal in length while in Siphluriscus the terminal filament is much
shorter than the cerci. Acanthametropus nymphs have conspicuous
lateral projections on the head, prothorax and metathorax and a median
tubercle on each abdominal tergum; these are not found in Analetris.
It is possible that Siphluriscus, known only from adults, and Acan-
thametropus, known only as nymphs, represent a single genus.

Analetris Edmunds, new genus

Subimago $ . — Eyes with ommatidia of fairly uniform size. Fore wings (Fig.
2) with fork of MA about % as long as base; vein MP deeply forked, the
fork over 4 times as long as the base; CuA connected to hind margin by 5
to 7 short crossveins (one forked on one wing, additional intercalary on one
wing not attached basally) . Hind wings (Fig. 3) more than 50% as long as
forewings; MP deeply forked. Tarsi more than twice as long as tibiae; claws
of each pair dissimilar, one rounded apically, one sharply hooked. Penes fused,
rounded apically; forceps base deeply V-shaped (Fig. 7). Terminal filament
subequal in length to cerci.

Nymph (Fig. 1). — Head without lateral or frontal projections; maxillary
palpi present. Fore femora broadest about % distance from base. Abdomen
without median tubercles on terga. Caudal filaments subequal in length. Gills
with margins entire, and with two ventral (posterior) smaller lobes (Fig. 4).

Type species. — Analetris eximia Edmunds, n. sp.

Etymology. From Greek, without a molar (grinder).

Analetris eximia Edmunds, new species

Genus et species incertus, Edmunds, 1954:64

Genus et species novum, Edmunds and Musser, 1960:113

Undescribed genus, Edmunds, 1957:23; Edmunds, Allen and Peters, 1963:10;

Lehmkuhl, 1970:183.

Subimago $. — Length: body 12; wings 12 mm. Head pale, with pair of
brown stripes between eyes; lower portion of eyes gray, upper portion orange.
Prothorax yellowish brown, notum slightly darker. Mesothorax yellowish brown,
with light brown stripe on each side between median line and inner parapsidal
furrows of the notum, continuing to scutellum; medium brown along outer
parapsidal furrows to wing bases, thin darker brown stripe along anterolateral
margins of mesocutellum ; medium brown marks at base of wings. Metathorax
yellowish brown, dark brown streak on each side near apex of scutellum;
medium brown marks at wing bases. Wings pale, longitudinal veins light brown,

Fig. 1. Analetris eximia, nearly mature nymph, dorsal view.

APRIL 1972] EDMUNDS & KOSS — ACANTHAMETROPODINAE

139

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

cross veins pale brown. Fore and hind tibiae and tarsi (Figs. 5 and 6). Legs
yellowish brown; tibiae, tarsi and apex of fore femora medium brown; middle
and hind legs with tibiae and tarsi darker than femora. Apex of each tarsomere
narrowly ringed in brown.

Abdomen largely pale yellowish brown except for tergum one; tergum one
largely medium brown, lateral margins pale, pale transverse lunar mark on
anterior margin each side of midline, and pale spot in middle area of tergum
on each side of midline; markings on terga 2 to 10 rather similar to those
same segments of nymphs (Fig. 1), but more diffuse; median brown band
bounded laterally by darker streaks; diffuse brown triangles lateral of median
stripe, bases on posterior margins, triangles not reaching lateral margins of
terga; darker line along posterior margin of each tergum. Sterna pale. Male
genitalia (Fig. 7) . Caudal filaments light brown at base, becoming paler apically.

Mature nymph. — Length: body 15, caudal filaments 4 mm. Color pale with
markings generally light brown (Fig. 1). Head pale with pair of brown stripes
between eyes from hind margin to ocelli ; ocelli gray ; apices of mandibles
and maxillae brown. Pronotum with narrow transverse brown stripe near front
margin, interrupted at midline; broader transverse stripe near middle of
segment, interrupted at midline and reaching halfway to margin; smaller brown
spot near margin on each side; sternum pale. Mesonotum pale with longitu-
dinal brown stripe laterad of inner parapsidal furrow on each side; stripes on
each side laterad of outer parapsidal furrow, extending to wing pad base;
wing pad bases largely pale; apex of scutellum brown; pleura and sternum pale.
Metathorax pale except for complex brown markings on metascutellum. Legs
pale with diffuse brown band near apex of femur; this band very pale on
middle and hind pair of legs.

Each abdominal tergum with wide brown median stripe containing four
pale dots, and with brown triangles on each side, their bases on hind margin;
dark brown streak near posterolateral corner of each tergum and dark brown
spot near each gill insertion. Sterna pale. Gills pale, washed with brown
dorsally on inner half. Caudal filaments pale basally, becoming increasingly
darker brown toward apices.

Holotype subimago $ and nymphal exuvium of same , Blacks Fork
River at Interstate Hwy. 80, 7 mi. E. of Little America, Sweet-
water County, Wyoming, collected 3 August 1968, emerged 9 August
1968, R. W. Koss and W. P. McCafferty, in collection of University
of Utah. Paratopotypes : 5 nymphs, 6 July 1968, R. W. and D. Koss;
3 nymphs, 3 August 1968, R. W. Koss and W. P. McCafferty; 20
nymphs, 18 July 1968, R. W. Koss. Paratypes: Wyoming: Sweet-
water Co.; Blacks Fork River at Granger, 4 nymphs, 3 August 1968,

Figs. 2-7. Analetris eximia. (Figs. 2,3 and 5 to 7. subimago $ ; Fig. 4.
nymph). Fig. 2. Fore wing. Fig. 3. Hind wing. Fig. 4. Gill 4, dorsal view.
Fig. 5. Fore tibia, tarsus and claws. Fig. 6. Hind tibia, tarsus, and claws. Fig. 7.
Genitalia, dorsal view.

APRIL 1972 ] EDMUNDS

& KOSS

acanthametropodinae

141

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R. W. Koss and W. P. McCafferty; Green River at Buckboard Flats,
1 nymph, 16 July 1959, G. W. Smith and G. G. Musser. Utah:
Daggett Co.; Green River at Hideout Canyon, 1 nymph, 3 September
1947, G. F. Edmunds, Jr. Saskatchewan: South Saskatchewan River
at Lemsford Ferry, 12 July 1970, D. M. Lehmkuhl, 10 nymphs, one
in Canadian National Collection, one in University of Utah collection,
others at University of Saskatchewan.

One or more paratypes in collections of R. W. Koss, Florida A & M
University, California Academy of Sciences, Purdue University and
Institute Royal des Sciences Naturelles, Brussells; others at University
of Utah.

Notes on the Habitat and Biology of Nymphs

Analetris nymphs are known only from warm rivers having a
constantly shifting sand substrate. Other insects found in this habitat
are the nymphs of the mayflies Pseudiron and Ametropus and the
dragonflies Ophiogomphus and Gomphus. The type locality in the
Blacks Fork River is slow-moving (approximately 0.5 ft./sec. on the
surface), shallow and alkaline. The river is mostly 1-2 feet deep in
mid- July and fairly clear. Following rains and in the spring it is
milky to muddy and several times the summer volume. At mid-day
17 July 1968 the water temperature was 21° C (71° F) . The water
temperatures at the Green River collecting sites varied from 19° to
23° C (66° to 74° F) in summer and fall.

The nymphs were most abundant in the fine-grained sand deposits
on the downstream end of deposition zones on the inside curves of
the meandering river. They anchor themselves on the shifting sand
substrate by burying the legs in the sand nearly to the base of the
tibiae by rapidly shifting the body forward and backward. The front
legs are positioned slightly laterigrade in a somewhat anteriorly
directed “pinching” position, while the middle and hind legs are
perpendicular to the substrate surface. Frequently the nymphs rapidly
fan the gills, excavating sand from below and around the body to
create a slight depression in the sand. In nature sand probably buries
the nymphs, and they were observed to remain buried in a laboratory
aquarium with circulating water. When the water was circulating
slowly they were observed to do a “push-up” movement, undulate the
abdomen, free themselves of sand, and fan the gills, after being
buried for 2-4 minutes. These reactions are presumed to be a response
to inadequate oxygen available to the nymphs. In nature such move-
ments may be necessary during low nighttime oxygen concentrations.

APRIL 1972] EDMUNDS & KOSS ACANTHAMETROPODINAE

143

Nymphs survived longer in circulating water than in continually
aerated standing water.

The nymphs often move over the sand searching the surface, perhaps
for suitable substrate or for food, with the maxillary palpi. The
mouthparts are obviously adapted for carnivorous habits and in the
lab they fed on red-blooded Chironomidae, eating them whole.

The nymphs swim rapidly, and they readily avoided hand screens,
rapidly dragged dip nets and a rapidly moved deep-bag net. The
most effective collecting device proved to be a net with an opening
3 feet wide by 1 foot high, and a bag 4 to 5 feet deep. The net was
used in shallow water, and it was hand held on the substrate. The
collector’s foot was used to “herd” or direct the specimens downstream
into the net. This was best accomplished by dragging the foot along
the bottom from an upstream position to a downstream one, starting
from one side of the net opening and gradually moving over the other
side. The net was then raised and shifted over to a new, undisturbed
area, and the process repeated.

When the same net was staked out in the river overnight as a
driftnet, it proved ineffective for collecting nymphs of Analetris and
Ametropus , but an excellent collecion of Pseudiron nymphs was made.

Lehmkuhl (1970, and personal communication) was able to collect
Analetris by rapidly moving a drag net upstream over the substrate
surface.

Literature Cited

Burks, B. D. 1953. The mayflies, or Ephemeroptera, of Illinois. Bull. 111.

Natur. Hist. Surv., 26 (Art. 1) : 1-216, 395 figs.

Edmunds, G. F., Jr. 1954. The mayflies of Utah. Proc. Utah Acad. Sci. Arts
Lett., 31 : 64-66.

1957. The predaceous mayfly nymphs of North America. Proc. Utah Acad.
Sci. Arts Lett., 34: 23-24.

Edmunds, G. F., Jr. and R. K. Allen. 1957. A checklist of the Ephemeroptera of
North America north of Mexico. Ann. Entomol. Soc. Amer., 50:
317-324.

Edmunds, G. F., Jr. and G. G. Musser. 1960. The mayfly fauna of Green River
in the Flaming Gorge Reservoir Basin, Wyoming and Utah. Univ.
Utah Anthropol. Pap. 43: 111-123.

Edmunds, G. F., Jr. and J. R. Traver. 1954. An outline of a reclassification of
the Ephemeroptera. Proc. Entomol. Soc. Wash., 56(5) : 236-240.

Edmunds, G. F., Jr., R. K. Allen and W. L. Peters. 1963. An annotated
key to the nymphs of the families and subfamilies of mayflies
(Ephemeroptera). Univ. Utah Biol. Ser. 13(1): 1-49.

Lehmkuhl, D. M. 1970. Mayflies in the south Saskatchewan River; Pollution
indicators. Blue Jay, 28: 183-186.

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Tshernova, 0. A. 1948. A new genus and species of mayfly from the Amur
Basin (Ephemeroptera, Ametropodidae) . Dokl. Akad. Nauk SSSR, 60
(8) : 1453-1455.

1967. May-fly of the recent family in Jurassic deposits of Transbaikalia
(Ephemeroptera, Siphlonuridae) . Entomol. Obozrenie, 46: 322-326.

Ulmer, G. 1920. Neue Ephemeropteren. Arch. Naturgesch., 85 (11) : 1-80.

SCIENTIFIC NOTE

Mass movements of Nymphalis calif ornica (Boisduval) in the San Fran-
cisco Bay area during 1971 (Lepidoptera : Nymphalidae) . — The California
Tortoise Shell butterfly undergoes periodic or sporadic population outbreaks
and mass movements which have been called migrations in the literature. The
last time such an event took place in the San Francisco Bay area was in
1959-1960 when several aggregations of the butterflies were observed flying in
spring and under bark in winter in Marin County, followed by tremendous
populations developed in June the following season. During intervening years
the species is scarce and may not be a continuous breeding resident here.

During October 1971, I observed movements of N. calif ornica in the Berkeley
Hills, Alameda County, at a site above the Caldecott Tunnel (about 1,400 feet
elevation). On 5 October, a smoggy day with temperatures in the 80’s °F.,
the butterflies passed to the southeast (along the axis of the hills) at a steady
2/minute on a 50 foot sighting line, between 11:45 and 12:05 p.m. (P.S.T.) .
Most flew 3-8 feet above the ground over the roadway which forms a corridor
through the trees at this point; a few flew circuitously or eastward through
the trees. However, on the open ridge east of the trees none could be seen
heading eastward, that is off the ridge axis. 7 October was cooler (77° maximum
in Oakland) with an easterly breeze. The Nymphalis were less numerous and
less directional in their activities, with 45 sighted in aggregated 30 minutes
surveillance beween 11:50 and 12:40 p.m., and only 75% moved in a south-
easterly or easterly direction. On 12 October, another warm, smoggy day (90°
in Oakland) , the flight was greatly increased, with a census of 392 during
periods totaling 35 minutes (11/minute) between 11:30 and 12:15 p.m. Counts
were made on the road and on open ridge knolls to the east and all individuals
moved southeasterly despite a stiff northeasterly breeze. The site was revisited
between 11:30 to 1:00 p.m. on 14 and 28 October, which were clear but much
cooler days, and no N. calif ornica were present.

Ten specimens were taken on 12 October and eight proved to be females,
the ratio perhaps in part the result of sampling error if males are faster, more
erratic fliers. Other butterflies flying in the vicinity included Precis coenia
(Hiibner), Adelpha bredowi (Geyer), Vanessa carye (Hiibner), Danaus plexippus
(L.), and Colias eurytheme (Boisduval). All but P. coenia were present in low
numbers and none showed any tendency for directional movement.

Sightings of individual N. californica were made in Walnut Creek, Contra
Costa County and in Berkeley on the University of California campus, during
the period of 5-12 October, by John Hafernik. Both localities are non-resident
sites for this species. — J. A. Powell, University of California, Berkeley, 94720.

APRIL 1972] BO HART & GRISSELL PULVERRO BIOLOGY

145

Nesting Habits and Larva of Pulverro monticola

(Hymenoptera: Sphecidae)

R. M. Bohart and E. E. Grissell
Department of Entomology, University of California, Davis, 95616

Pulverro Pate is a nearctic genus with seven described species.
Nothing has been published about its biology and very little about
that of related genera such as Ammoplanops and Ammoplanus. No
doubt the small size of these wasps has helped them avoid observation.
The somewhat related genus Microstigmus is known to take Collembola
or Thysanoptera as prey, so it is not surprising that thrips are used
by Pulverro.

The nesting site of Pulverro monticola Eighme was first observed
in July 1968. A detailed study was made in July 1970 and some
additional work was done in the summer of 1971. Many nest entrances
were found in the side banks along 400 m of dirt logging road near
the Sagehen Creek Biological Station of the University of California
at an altitude of about 6,500 feet in Nevada County, California. To
some extent the nests were grouped, with 15 to 20 in a square meter
area, and the entrances were sometimes only a few mm apart. On the
other hand some nests were rather isolated or scattered along the
roadside. The following description of our primary study area will
serve to characterize other sites used by P. monticola. The roadside
mound was about 0.5 m high and covered nearly 2.0 m 2 . The soil was
fine grained intermixed with many small pebbles and a few stones up
to 15 or 20 cm in length. Surface features included several larger
rocks, a few broken branches, and the remains of a small stump. The
mound was sparsely clothed in Lepidium about 15 cm tall. From 1
to 24 July 1970 we located 32 nests in the mound some of which were
started as late as 24 July. The season was considerably retarded by
cold weather in 1971 and the nesting period was advanced to the
approximate period of 15 July to 7 August.

During the middle of the nesting season, nest activity began about
8:45 a.m. Pacific Standard Time as the sun struck the east slope of
the mound and ceased about 4:00 p.m. as sunlight left the west slope.
Female Pulverro nesting on the east slope were active about an hour
earlier in the morning and stopped about an hour earlier in the after-
noon than those on the west slope.

Daily activity began as the female wasp emerged head first and
remained for a few minutes in the sunlight near the entrance. A

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Figs. 1-4. Prepupal larva of Pulverro monticola Eigme, ventral views. Fig. 1.
Whole larva. Fig. 2. Head enlarged; ant. antennal orbit, a.t.a. = anterior
tentorial arm. Fig. 3. Mouthpart area enlarged. Fig. 4. Mandible. Fig. 5.
Vertical section through a nest of P. monticola ; a, entrance on ground slope;
b, completed cell sealed off by a dirt plug; c, bend in burrow where loose
thrips were found; d, nearly complete cell.

vigorous cleaning activity then began in which the wasp went head
first into the nest, kicked out small dirt particles with her legs and
dragged out larger particles in her mandibles. This material accumu-
lated on the slope immediately beneath the entrance and was some-

APRIL 1972] BO HART & GRISSELL PULVERRO BIOLOGY

147

times distributed somewhat by the wasp. This tumulus served as a
convenient marker for the nest entrance just above it.

Underground morphology of the burrows was determined by ex-
cavation of about 20 of them. This process was extremely difficult and
often unsuccessful because of the small burrow diameter (about 2
mm) and the rock and stick inclusions in the soil. Plaster of Paris,
both dry and in dilute suspension, was pipetted into the burrows to
aid in tracing them. This helped considerably but the uneven soil
texture still created problems. Finally, earth at several sites was sifted
before the nesting season in June 1971. The resulting fine and even
grained soil was used by several Pulverro in July of the same year.
Excavation of these nests in early August 1971 was fairly successful.
One nest followed in its entirety appeared to be typical. Its directions
and dimensions are shown in Figure 5. The burrow was briefly
horizontal, then curved downward to a depth of about 85 mm. At
the 60 mm level a short side burrow, plugged with dirt, led to an
indistinct cell containing 21 thrips, the agglutinated remains of about
12 more thrips, and a half-grown wasp larva (Fig. 5b). At the 85
mm level four thrips were lodged loosely in a bend of the burrow
(Fig. 5c). A cell at the end of the short unplugged side burrow which
followed contained 28 thrips but no egg or larva (Fig. 5d) . Judging
from several excavations, the wasp may deposit thrips temporarily
at a bend opposite the cell, and move them to the cell later. This idea
fits well with the provisioning observations described below.

Prior to provisioning, wasps of both sexes were abundant on flowers
of several sorts, particularly those with a short corolla. Examples
were Potentilla gracilis Dougl. ex Hook., Calyptridium umbellatum
(Torr.) Greene, Naina lobbii Gray, Phacelia species, Chamaesaracha
nana (Gray) Gray, and Ligusticum species. Males could be found
at flowers for about two weeks after the onset of nesting, but copulation
was not observed. Provisioning females presumably collected their
prey on the same flowers. Furthermore, they were observed to crawl
down the corollas of Penstemon gracilentus Gray, apparently in search
of prey. Female wasps were first seen provisioning on 4 July 1970.
Thrips were carried singly, venter uppermost and held in the mandibles
of the wasp. Each thrips was grasped by the thorax with its head
projecting anteriorly from the wasp’s mandibles, and the rest of
the thrips lying under the thorax of the wasp. When a wasp was
captured in an aspirator, it walked readily about the tube without
using its legs to hold the prey. The provisioning female hovers
briefly about its nest and then enters directly without alighting.

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Ordinarily the wasp deposits its prey and emerges in 10 to 40 seconds.
However, several timings were as short as three to five seconds.
We presume that in these instances the female merely dropped her
thrips at a bend in the burrow, as suggested above. On leaving the
burrow a female waits momentarily with her head at the entrance,
then walks back and forth in front of the entrance, pauses, and flies
around in several erratic circles before disappearing. Time recorded
for searching during mid day varied from 40 seconds to six minutes,
with about two minutes on the average, based on 30 observations
and five wasps.

Prey of Pulverro monticola appears to be mostly adult thrips but
occasionally a few large immatures are included in the provisions.
Observed Thysanoptera were Frankliniella moultoni Hood and a few
specimens of Aeolothrips fasciatus (Linnaeus). Both of these were
abundant on flowers in the vicinity.

Judging from the observation of many nests, the entrances are
never closed by the wasp while she is absent in search of thrips or
after the nest is complete. This is in agreement with the fact that
female Pulverro have the tarsal rake undeveloped. The entrance is
thus frequently exposed to predators and parasites. Ants of various
sizes have been seen in fair numbers on the nesting site but even
those small enough to enter the burrows have not done so. No
sarcophagid flies have been noted. All of the nyssonine parasites and
most of the chrysidids which visited the site were too large to enter
the burrows. One minute species of Hedychridium was seen to inspect
the nests and could conceivably be a parasite of Pulverro. Soon
after nesting is completed, the burrow entrances are partially blocked
by wind blown dust and doubtless they are eventually sealed off
by rains.

In one of the excavations a prepupa was found. It was in a silken
cocoon at a depth of about 80 mm. The cocoon was egg-shaped,
2.75 mm long and 0.85 mm at its greatest breadth. The outer surface
of the cocoon was covered with sand grains and parts of thrips. The
prepupa was 2.5 mm long and 0.8 mm wide. It was subsequently
sketched while still alive, then partially cleared and mounted on a
slide for microscopic examination. Details are given in Figures 1—4.
Essential features of the larva are as follows: (1) body fusiform
rather than cylindrical, largely smooth, a few scattered microsetae
including some on mouthparts; (2) head capsule 0.40 mm long, 0.44
mm wide; (3) antennal orbits and frontoclypeal suture not distinct,
mouthparts pigmented only toward apex of mandible; (4) labrum

APRIL 1972] BOHART & GRISSELL — PULVERRO BIOLOGY

149

weakly emarginate apically, bearing about 10 scattered microsetae;
(5) mandible with four teeth including a weak basal one, a single
bristle near middle of outer margin; (6) maxilla largely asetose
but with scattered spicules in lacinial area, maxillary palpus stout
and much larger than galea; (7) spinnerets acute apically, not con-
nected by a membrane and not exceeding labial palpi.

SCIENTIFIC NOTE

A host record for Fedtschenkia (Hymenoptera: Sapygidae). — The genus
Fedtschenkia occurs in both western North America and in palearctic Asia.
There has been some question as to its affinities since it resembles certain
Scoliidae, such as Cosila, but male genitalia and other features place it in the
Sapygidae. There has long been speculation about the identity of its host.
All other known hosts of sapygids are bees, particularly megachilids but also
xylocopids.

In 1956 at Tanbark Flat in the San Gabriel mountains of California the
senior author observed a female Fedtschenkia anthracina (Ashmead) entering
a ground burrow which was found to contain a cell with a dead adult of the
eumenid wasp, Pterocheilus trichogaster R. Bohart. Evidence of a relationship
seemed flimsy until D. J. Horning and the junior author excavated a number
of burrows of P. trichogaster on Santa Cruz Island, California late in April of
1969. F edtschenkia were abundant and active as parasites in the nesting area.
Finally, the senior author at Arroyo Seco, Monterey County, California in May
1971 observed F. anthracina females entering and staying for considerable
periods in burrows of P. trichogaster which were provisioning with geometrids
of the genus Hydriomena Hiibner (det. M. R. Gardner). — R. M. Bohart and
R. 0. Schuster, Department of Entomology, University of California, Davis, 95616.

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150

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

A New Seed Inhabiting Cerambycid from Costa Rica

(Coleoptera)

John A. Chemsak

Division of Entomology, University of California , Berkeley, 94720

The number of known seed-infesting Cerambycidae is relatively
small considering the great potential for larval development this habitat
provides. In North America, Paratimia conicola Fisher lives in the
cones of Pinus attenuata and P. bolanderi (Craighead, 1923; Linsley,
1962) ; Ataxia sulcata Fall and Lepto stylus terraecolor Horn in the
seeds of Rhizophora mangle (Craighead, 1923) ; Leptostylus gibbulosus
Bates in seeds of Sapindus (Vogt, 1949). In Latin America, seed-
infestors include Lepturges spermophagus Fisher in Cowpea, Vigna
(Fisher, 1917) ; Baryssinus leguminicola Linell in Enter olobium (Gil-
mour, 1965 has synonymized this species under Lophopoeum timbouvae
Lameere which is known to infest seeds of various legumes as listed
by Duffy, 1960) ; Leptostylus gundlachi Fisher in pods of coralbean,
Erythrina glauca (Wolcott, 1948) ; and Lepturges guadeloupensis
Fleutiaux and Salle has been reported from pods of Acacia by
Wolcott (1948).

The following new species, reared from Diospyros fruits by P. A.
Opler, was made available by D. H. Janzen and is described at this
time to make the name available for other studies. The illustration
was prepared by Celeste Green. This study was conducted during
the course of National Science Foundation Grant GB-31120X.

Leptostylus spermovoratis Chemsak, new species

(Fig. 1)

Male.— Form moderate sized, robust, convex above; ground color reddish
brown, apex of abdomen dark; pubescence dense, short, appressed, mottled
whitish-brown and black; elytra with a broadly triangular black chevron behind
middle. Head with front shallowly convex, subquadrate, deeply impressed between
antennal tubercles; antennal tubercles elevated, divergent; punctation obscured
by pubescence; pubescence mottled, front mostly dark; eyes moderately coarsely
faceted, deeply emarginate, separated above by more than diameter of antennal
scape; mouthparts and base of labrum with a few long erect hairs; antennae a
little longer than body, basal segments mottled with dark brown, segments
from fifth pale, dark annulate at apices, scape extending to about middle of
pronotum, third segment slightly arcuate, longer than first, fourth slightly
shorter than first, remaining segments gradually decreasing in length. Pronotum
broader than long, sides slightly inflated; base broadly impressed, apex more
narrowly impressed; disk with five prominent tubercles, punctures around

The Pan-Pacific Entomologist 48: 150-152. April 1972

APRIL 1972]

CHEMSAK NEW CERAMBYCID

151

Fig. 1. Leptostylus spermovorads Chemsak, $.

tubercles irregular, base with a line of coarse punctures; pubescence gray-
brown, appressed, black basally at sides and middle and at apex on each side
of middle; prosternum narrow, intercoxal process narrower than width of coxa,
expanded behind, coxal cavities closed; mesosternal process broad, abruptly

152

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 2

declivous in front, middle coxae with small tubercles internally; scutellum
black, triangular, rounded behind, sparsely pubescent. Elytra less than twice
as long as broad, tapering apically, broader than pronotum; disk with semi-
circular elevated ridges at base on each side of scutellum, ridges bearing
elevated black pubescent tubercles, dark pubescent tubercles scattered over
surface but forming a broad triangle behind middle; pubescence mottled, sides
with a dark band extending down humeri to middle; apices narrow, truncate.
Legs robust, femora clavate, pale and brown mottled; tibiae each with two
dark bands. Abdomen densely pale pubescent at sides, sparsely pubescent at
middle; last sternite dark, sparsely pubescent, truncate at apex. Length, 10 mm.

Female.- — Form similar. Antennae slightly shorter. Abdomen with last sternite
rounded at apex. Length, 10-12 mm.

Holotype male, allotype and two paratypes (1 male, 1 female) from
Comelco, Bagaces, Guanacaste, Costa Rica, emerged from Diospyros
fruits on 8 January 1971 (P. A. Opler).

The type series varies little in coloration and the elytral pattern
of all the specimens is quite similar.

Literature Cited

Craighead, F. C. 1923. North America cerambycid larvae. Can. Dept. Agr.
Bull., 27(N.S.) : 1-239, 44 pis.

Duffy, E. A. J. 1960. A monograph of the immature stages of Neotropical
timber beetles. Brit. Mus. (Natur. Hist.). 327 pp., 13 pis.
Fisher, W. S. 1917. A new species of longhorn beetle infesting cowpeas from
Mexico. Proc. Entomol. Soc. Wash., 19: 173-174.

Gilmour, E. F. 1965. Catalogue des Lamiaires du Monde. Mus. G. Frey,
Munich, pp. 559-655.

Linsley, E. G. 1962. The Cerambycidae of North America. Part 2. Taxonomy
and Classification of the Parandrinae, Prioninae, Spondylinae, and
Aseminae. Univ. Calif. Publ. Entomol., 19: 1-102, 1 pi.

Vogt, G. B. 1949. Notes on Cerambycidae from the Lower Rio Grande Valley,
Texas. Pan-Pac. Entomol., 25: 175-184.

Wolcott, G. N. 1948. The insects of Puerto Rico. Coleoptera. J. Agr. Univ.
P. R., 32: 225-416.

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Vol. 48 JULY 1972 No. 3

SAKAGAMI — Bumble Bees Collected by the California Academy — Lingnan
Dawn-redwood Expedition to Central West China, 1948 (Hymenoptera:
Apidae) 153

SCHLISING — Foraging and Nest Provisioning Behavior of the Oligolectic

Bee, Diadasia bituberculata (Hymenoptera: Anthophoridae) 175

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kensis at Point Pinos Sewage Outfall, Monterey County, California
(Diptera: Chironomidae) 204

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Ceramica picta (Lepidoptera: Noctuidae) 208

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SCIENTIFIC NOTES 203, 218

SOCIETY ANNOUNCEMENT— Usinger Autobiography 219

SAN FRANCISCO, CALIFORNIA • 1972

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

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The Pan-Pacific Entomologist

Vol. 48 July 1972 No. 3

Bumble Bees Collected by the California Academy — Lingnan
Dawn-Redwood Expedition to Central West China, 1948

( Hymenoptera : Apidae)

Shoichi F. Sakagami
Zoological Institute, Hokkaido University, Sapporo

The present paper is a report on the taxonomic study of the
bumble bees collected by the California Academy — Lingnan Dawn-
redwood Expedition to Central West China (1948), or more familiarly
known as the Metasequoia Expedition. The main purpose of the
expedition was “to collect insects and other animals in the hope
of finding some ancient faunal elements of possible North American
affinity which might have survived with Metasequoia and the other
ancient trees associated with it” (Gressitt, 1953). Bumble bees
are not suitable material to trace such a relation, yet the collected
specimens are valuable because of insufficient information on this
group in Eastern Asia. Numerous names have been published for
the Chinese bumble bees (Wu, 1941), and a zoogeographical con-
sideration was given by Panfilov (1957). Most descriptions of
Chinese forms published prior to 1940 were incomplete. The result-
ing misinterpretations of taxa were serious for any further studies
on this difficult group. Only recently various “forms” are being
restudied accurately by the examination of type specimens (Tkalcu,
1960, 1961a, b, 1968a,b). For the time being, however, we need
more effort to clarify each taxon inhabiting this vast area. For
this purpose, the present study may serve as a milestone for future
synthesis.

Except for a few specimens collected in 1950 from Luichow
Peninsula, Kwangtung Prov. by Dr. J. L. Gressitt, most specimens
were collected by him and Mr. Y. W. Djon during July to September
1948 from the localities as shown in Fig. 1. The map is based upon
a sketch prepared by Dr. Gressitt.

The subgeneric system mainly follows that by Richards (1968).
The synonymic lists include only the original description and im-
portant papers.

The Pan-Pacific Entomologist 48: 153-174. July 1972

154

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48 , NO. 3

Fig. 1 . Map showing localities where the specimens were collected. Hupeh
Province (Lichuan District): Sui-hsa-pa (= Suisapa, 1,000 m, SP), Hsao-ho
(~ Hsiacho, HC), Wang-chia-ying (Wang-ga-ying, WY), Gan-yu-yai (GY),
Song4ung-chi (SL), Leung-ho-lceo (Leung-ho-kow, LK), Szechuan Province
(Wanhsien District): Mo-tau-chi (MT), Lung-chiu-pa (Lunchepa, LP).

Before going further, I would like to express my heartiest thanks
to Dr. Paul H. Arnaud, Jr. and Mr. Hugh B. Leech, California
Academy of Sciences, San Francisco, and Dr. Yoshihiro Hirashima,
Kyushu University, Fukuoka, who gave me the opportunity to study
valuable materials, Dr. J. L. Gressitt, Bernice P. Bishop Museum,
Honolulu, who informed me through Mr. Leech of details about the
localities where the specimens were collected, and Dr. Robbin W.
Thorp, University of California, Davis, who read the manuscript

july 1972]

SAKAGAMI CHINESE BUMBLE BEES

155

and helped me publish it. My sincere gratitude is also expressed
to Dr. Borek Tkalcu, Praha, whose expert knowledge based upon
examinations of many type specimens and whose unselfish answers
to my questions were indispensable to complete the work.

Bombus (Tricornibombus) atripes Smith
Bombus atripes Smith, 1852 : 44, 2 , S .

Megabombus ( Tricornibombus ) atripes , Tkalcu, 1968b: 83, 2, 2, $.

Specimens examined. — WY, 22-23 September, 3 £ ; SL-GY-WY, 19 September,
22,14.

Recently this and the next species were thoroughly redescribed by
Tkalcu (1968b). The hair coat coloration, bright orange brown above,
blackish below, is constant for all specimens examined.

Bombus (Tricornibombus) imitator Pittioni

Bombus ( Tricornibombus ) imitator Pittioni, 1949: 251, 2-

B. ( T .) imitator var. flavescens Pittioni, 1949 : 254, 2 .

Megabombus ( Tricornibombus ) imitator, Tkalcu, 1968b: 90, 2, 2, $•

Specimens examined. — SP, 26 July, 2 2 , 2-3 August, 2 2 , 16 August, 1
2 , 1 $ , 22-30 August, 11 2 ? 1 $ , 12-17 September, 8 2, 3 $ ; HC, August,
1 2 ; LK, August, 1 2 , 31 August, 2 $ , 7 September, 12, 9 September, 1 $ .

These specimens were seemingly already examined by Tkalcu (cf.
Tkalcu, 1968b, p.93). In most specimens the apical half of metasomal
tergum III bears pale ochre orange hairs, occasionally black hairs
are confined to the anterior lateral corners (= flavescens Pittioni,
cf. also Tkalcu, 1968b). Terga III and IV of one male are tawny
yellow orange with sparse admixture of black hairs anteriorly. Black
interalaris of most specimens is poorly defined, and in small workers
often represented by an obscure dark patch.

Bombus (Thoracobombus) opulentus Smith
Bombus opulentus Smith, 1861: 153, 2-

Megabombus ( Agrobombus ) opulentus, Tkalcu, 1968a: brief note.

Specimens examined : — WY, 23 September, 1 2 , 22 September, 1 $ ; GY-WY,
20 September, 1 $ .

The identification of these specimens as B. opulentus , a long for-
gotten taxon, was based upon a brief note by Tkalcu given in the
description of Bombus honshuensis (Tkalcu) and confirmed by him-
self {in litt .) . Because Dr. Tkalcu is preparing the descriptions of
three castes of this species, only some remarks on hair color and
some male characters are given here.

Hair color. — Worker . — Mesosomal disc, upper half of mesosomal side and
metasomal terga I - — - II orange brown. Head excluding sparse orange brown

156

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48 , NO. 3

Fig. 2. Comparison of males of B. opulentus (A ^ D, F) and B. schrencki
(E). A. Antenna; B. Apex of genitalia dorsal view; C. Same dorsolateral view;
D. Lower membrane of basal process of gonostylus; E and F. Sculpture of
posterior part of metasomal tergum IV.

hairs on vertex, lower half of mesosoma, terga IV VI, sterna and legs blackish
to chocolate brown, tergum II laterally broadly blackish. Male. — Generally as
worker but differing in following points: Short plumose hairs on face and gena
pale fulvous; vertex mostly pale orange brown with sparse black hairs; lower
half of mesosoma with more extensive orange brown hairs, leaving narrow dark
chocolate band on venter; coxae and trochanters below with pale brownish
hairs; hairs of tergum II entirely orange brown, slightly paler apically, forming
obscure marginal ciliation; hairs of tergum III entirely blackish in one specimen,
basal half blackish and apically orange brown in another, both with obscure
ciliation; sterna with obscure marginal ciliation.

Male characters. — (cf. Synoptic table of B. unicolor (Friese) rz U, B.
schrencki (Morawitz) = S and B. honshuensis — H in Tkalcu 1968a, p.48) : 1)
Flagellar segments IV X moderately swollen below (Fig. 2 A, = U, =£ S,H) .

2) Punctures bearing hairs on hind tibia dense and homogenous (= S,U, =7 - H) .

3) Forewing darker than in S. and U, with slight iridescence. 4) Penis valve
below with short rectangular denticule ( =U,S , H) . 5) Lateral depression
of gonocoxite basally not distinctly demarcated (Fig. 2 C, = U,S, H) . 6)
Apical contour of gonostylus similar to that of U, inner angle slightly more
pointed though not so conspicuously as in S (Fig. 2 B,C). 7) Lower membrane
of inner process of gonostylus as in S (Fig. 2D). 8) Basal process of lacinia
dorsolaterally well visible (Fig. 2 C, = S,H) . 9) Barba mandibularis honey
yellow (=U,H, y^=S). 10) Erect hairs of scape blackish brown (= U,S, ^ H ) .

Thus, B. opulentus is more closely related to B. schrencki and B.
unicolor than to B. honshuensis. Besides the characters noted above,
the punctures on tergum IV, especially marginally, are better defined

july 1972]

SAKAGAMI — CHINESE BUMBLE BEES

157

& A Jt

a,i h 1 ^ 1 1 /

vz ill * v

Fig. 3. Distribution and color patterns of the four species of the subgenus
Senexibombus Frison. All color patterns shown are of females, except B. senex
f. pallidithorax (IV) known by male only. Bombus kulingensis Cockerell (Central
West China) I. Typical form, II. “ tajushanensis ” Pittioni, III. k. f. quasiflavior
Tlcalcu and “t.” f. flavior Pittioni, IV. “t” f. nigrofasciatus Pittioni, V and VI
a worker and queen described in text; B. bicoloratus Smith (Formosa) I. Typical
form, II. f. fulvolateralis Cockerell, III. f. intermedius Chiu ; B. irisanensis
(Philippines, in variation recorded) ; B. senex Vollenhoven (Sumatra) I. Typical
form, II. f. sumatrensis Cockerell, III. f. ardentior Cockerell, IV, f. pallidithorax
(Frison) ( $ ) . Color codes: A. Black, B. Admixture of blackish hairs, C.
Whitish, D ~ E. Pale to deep yellowish or ochraceous, F. ferruginous. G.
Distribution of bumble bees in E. and SE. Asia.

158

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

and denser in B. opulentus than in B. schrencki (Fig. 2E, F) . Certainly
B. opulentus may be regarded as the Southernmost representative of
the B. schrencki-gvowp.

Bombus (Senexibombus) kulingensis Cockerell
Bombus kulingensis Cockerell, 1917 : 266, $ .

Bombus ( Senexibombus ) tajushanensis Pittioni, 1949: 244, $, £ ; f. nigro-

fasciatus, 246, $ , $ ; f. flavior, 247, $ , 2 ■

Bombus ( Senexibombus ) kulingensis, Tkalcu, 1961a: 46, $ ; f. quasiflavior,

52, $.

Specimens examined. — SP, 26 August, 12 (in very poor condition), 27 August,
1 2 , 29 August, 1 2 •

By the synoptic table of B. tajushanensis and B. kulingensis by
Tkalcu (1961a), the specimens examined straightly go to the lattel.
But Dr. Tkalcu recently informed me that B. kulingensis and B.
tajushanensis are conspecific. Two specimens examined are char-
acterized by their melanism, a queen without pale hairs on meso-
somal dorsum, and a worker with tergum III completely, IV nearly
entirely black. Figure 3 shows the distribution and color patterns
of the four species forming Senexibombus , a small subgenus endemic
to Eastern and Southeastern Asia at the S.E. Asiatic boundary of
bumble bee distribution. The record of B. kulingensis from Western
Hupeh represents, for the time being, the northwestern limit of the
subgenus.

Bombus (Diversobombus) trifasciatus Smith
Bombus trifasciatus Smith, 1852 : 43, 2 , 2-

Bombus ningpoensis Friese, 1909: 675, 2 (cf. Tkalcu, 1961b, 367).

Bombus ( Diversobombus ) ningpoensis, Pittioni, 1949: 251, 2, 2; Tkalcu, 1960:
4, 2 ■

Bombus ( Diversobombus ) ningpoensis subsp. minshanicus Bischoff, 1936: 19, 2*
Specimens examined. — SP, 22-31 July, 232, 1-10 August, 22 2, 11-20
August, 68 2 , 1 5 , 21-31 August, 77 2 , 1-10 September, 12, 2 5, 11-18
September, 20 2 , 15; LK, 12 August, 1 2 , 28-31 August, 5 2 , 1-10 September,
87 2, 15 ; LK-SP, 21 August, 12, 18 September, 2 5 ; HC, 4 August, 12, 12-16
August, 82; WY-SP, 21 July, 12; WY, 22 September, 22; SL-GY-WY, 19
September, 32 ; LP, 29 September, 12 ; MT, 25-26 September, 42 ; Kwangtung
Prov., Luichow Peninsula, 8 September, 1 2 •

This is apparently one of the most abundant and widespread bumble
bee species in Western China. Some structural differences from B.
diver sus Smith, the type species of the subgenus, are as follows :
1) Punctures on ocellocular area above (2, 5) more well defined,
contrasting with interspaces smoother, showing less marked undulations

July 1972]

SAKAGAMI — CHINESE BUMBLE BEES

159

Fig. 4. Comparison of B. trifasciatus (A,C,D,G,J) and B. diversus (B,E,F,H,K).
A,B. Sculpture on ocellocular area (worker) ; C,E. Apex of male genitalia,
dorsal view; D,F. Same, dorsolateral view; J,K. Same, apical view; G,H.
Punctation of male clypeus.

160

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Fig. 5. Comparison of B. trifasciatus (M,N) and B. diversus (L,0) , continued.
L,M. Basal flagellar segments (worker) ; N, 0. Ditto (males) ; P, Q. Parts
used for relative size measurements (cf. in text) .

(Fig. 4A, B). 2) Punctures on metasomal terga (£, $), especially
IV and V, denser, submarginally punctures often nearly as wide
as interspaces. 3) Male clypeus with less coarse punctures and medio-
apical smooth area less contrasting (Fig. 4G, H) . 4) Male antennal
flagellum I slightly longer and II slightly shorter (Fig. 5N, 0). 5)
Inner apex of male gonocoxite relatively well projecting (Fig. 4C— F,
J, K) . 6) Inner angle of male gonostylus more angulate apically
(Fig. 4C-F, J, K). 7) Inner process of male gonostylus shorter and
bifurcate (Fig. 4C, E, J, K) . 8) Outward serration of head of penis
valve less projecting but more continuous (Fig. 4C, E, J, K).

The shape of the inner process of the gonostylus is most critical.
On this character, B. diversus , (Japan), B. ussurensis Radoszkowski
(Japan, Korea, Ussuri) and B. hummeli Bischoff (North China, cf.
Bischoff 1936, p. 19) form one group and B. trifasciatus another,
together with B. wilmani Cockerell (Formosa), B. montivagus Smith
(S.E. Asia, cf. Tkalcu, 1968c), and B. albidopleuralis Friese (Nepal),
suggesting the segregation of two species groups between northern
and southern areas of the range of the subgenus Diversobombus
endemic to East and S.E. Asia.

Using the rich material some variation studies were executed as

July 1972]

SAKAGAMI CHINESE BUMBLE BEES

161

A A

A A A A A A

mm mm

xrmin? wmy mm/

ACA mz ACB

BBC

wmw

mm

CAD ^DAB

Fig. 6. Color variation in B. trifasciatus. a. Mesosomal dorsum; b,c. Meta-
somal terga II and IV ; d. Selected color patterns, I. trifasciatus from original
description; II. f. minshanicus; Others showing representative patterns described
in text.

follows: Hair color. — The variation was studied with respect to
the following three parts, which were most variable, of workers with
relatively intact hair coat. 1) Mesosomal dorsum. Extent of black
interalaris is variable. In most melanic specimens, pale collaris
completely disappears. In the flavinic extreme, interalaris remains
as a relatively narrow band (Fig. 6a, A-D). 2) Metasomal tergum
II varies from completely yellowish to distinctly black banded pos-
teriorly (Fig. 6b, A-D). 3) Tergum IV varies from completely black
to nearly entirely ferruginous, leaving sparse black hairs anterolaterally
(Fig. 6c, A-B).

The frequency distribution of various combinations of color patterns

162

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

(Fig. 3) is as follows: BBB 116, BBC 49, CBB 18, BBD 17, CBC 15,
BCB 14, BAB 9, ABC, DBB each 7, ABB 6, BCC 5, BBA, BDB, CAB,
CBD, ABD, DAC each 2, AAB, ABA, ABD, ACA, ACB, ACC, BAA,
BAC, BAD, BCD, CAA, CAD, CBA, CDD, DAB, DBC each 1. For
each body part, the frequency distribution is as follows: Mesosomal
dorsum, A 20, B 222, C 43, D 11; tergum II, A 23, B 248, C 24,
D 2; tergum IV, A 13, B 193, C 72, D 23, E 4. Pittioni (1949)
referred to the relatively constant color pattern of this species with
variability exhibited mostly in the extent of interalaris. The chromatic
constancy of this species was also suggested by Tkalcu (1960). The
population studied by me seems to be more variable, though not
so obviously as to be called polytypic. The prevalence of type B
in mesosomal dorsum corresponds to subsp. minshanicus described
from S. Kansu, characterized by the extension of interalaris and
resulting diminution of both collaris and scutellaris, together with
the appearance of narrow reddish band at the posterior margin of
tergum IV in some specimens. In the specimens examined the decrease
of collaris often reaches the extreme (type A, complete absence, and
the invasion of reddish hairs on tergum IV is often greater (more
advanced) , whereas the diminution of scutellaris was never observed.
Apparently the population studied shows a tendency to subsp. min-
shanicus-tyipe but it is open to the question whether the species is
divided into more than one distinct subspecies within the range.
A single worker from Luichow Peninsula, Kwangtung Prov. showed
type ABB, and four males with intact coat respectively BBB, BAC,
CAA and DBD.

The variability of mesosomal disc correlates to the body size as
follows :

v -S3

3.81 ~

4.00 /'*■'

4.18 ~

Total

Head width (mm)

3.75 ^

3.94

4.13

4.32

^ 4.37

bees

Color pattern A

2

5

10

3

20

B

15

40

71

79

17

222

C

9

12

9

12

1

43

D

3

1

3

4

11

Melanism appears more frequently in larger specimens. No such
correlation was detected for terga II and IV.

Allometric tendency. — Moure and Sakagami (1962) enumerated
morphological differences between queen and worker bumble bees.
Some of them, especially those of the head, are similar to the allometric
tendency of other bees (cf. Sakagami and Moure, 1965). This

July 1972]

SAKAGAMI — CHINESE BUMBLE BEES

163

feature was studied by using the specimens of B. trifasciatus , with
respect to malar index (length/width of malar space, cf. Fig. 5 Q,
C/D) and occipital index (inter ocellar distance/ocelloccipital distance,
cf. Fig. 5P, B/A). The result is given as follows:

Head width
(mm)

Malar

index

Number of
(specimens)
measured

Occipital

index

Number of
(specimens)
measured

3.25 — 3.57

1.21

(12)

1.10

(14)

3.62 — 3.75

1.18

(14)

1.15

(13)

3.81 — 3.88

1.23

GO)

1.21

( 9)

3.94

1.22

(14)

1.23

(13)

4.00

1.27

(16)

1.29

(16)

4.06

1.26

(12)

1.30

(ID

4.13

1.28

(17)

1.31

(20)

4.19

1.27

(17)

1.34

(13)

4.25

1.28

(17)

1.40

(18)

4.31

1.30

(17)

1.40

(15)

4.38

1.31

(12)

1.43

(11)

4.43 — 4.50

1.29

(10)

1.46

(12)

The gradual increase of the ratio is conspicuous for the occipital
index and, though less remarkable, is also traced in the malar index.
The latter is of particular importance because this index is often
adopted as a useful distinctive character. This allometric tendency
is more conspicuous when queens and small workers are compared.
For instance, the ranges and means of ratios in B. diver sus tersatus
Smith, measured each in five queens and five dwarf workers are
as follows:

Head width (mm) Malar index Occipital index

Queens 4.87— 5.13 (5.04) 1.20— 1.26 (1.24) 1.55 — 1.66 (1.62)

Workers 2.56-2.82 (2.66) 0.95- 1.16 (1.04) 0.65 - 0.88 (0.79)

Bombus (Alpigenobombus) BREVICEPS channicus Gribodo
Bombus channicus Gribodo, 1891 : 116, £ , $ •

Alpigenobombus breviceps channicus , Tkalcu, 1968c: 16, 7, S.

Specimens examined. — MT-LP, 28 September, 1 £ ; LP, 29 September, 1 £ ,

1 $.

Hind tibia and basitarsus are bright chestnut, not yellowish ochre
as in B. b. vicinus (Tkalcu) .

Bombus (Bombus) ignitus Smith

Bombus ignitus Smith, 1869: 207, 2, Tkalcu, 1962: 17, 2, $•

Bombus ( Bombus ) ignitus, Sakagami & Ishikawa, 1969: 185.

Specimens examined. — GY-WY, 20 September, 1 $ ; SL-GY-WY, 19 September,

164 THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Fig. 7. Bombus pyrosoma (male). A. Basal flagellar segments; B and C.
Apex of genitalia dorsal and dorsolateral views.

2 $ ; HC, 8-10 August, 2 £ , 13-15 August, 3 $ ; LK, 9 September, 1 $ ; SP,

3 August, 3 £ •

All specimens have sparse ochraceous hairs on metasomal tergum
II (=var. balteatus Skorikov, 1933 or var. cancellatus Frison, 1935),
and some specimens have similar hairs on tergum I, either anteriorly
or posteriorly or mesosoma anteriorly (= var. subcollaris Skorikov,

1914) .

Bombus (Melanobombus) pyrosoma F. Morawitz
Bombus pyrosoma F. Morawitz, 1890: 349, $.

Bombus ( Lapidariobombus ) pyrrhosoma pyrrhosoma, Bischoff, 1936: 9, $,

s, s.

Bombus ( Lapidariobombus ) pyrrhosoma, Tkalcu, 1961b: 353, £ .

Specimens examined.— WY, 22 September, 1$ ; LK, 7-9 September, 9£ ;
SP, 30 July, 6 August, 2$ ; WY-SP, 21 July, 1$ ; SL-GY-WY, 19 September,
1 £ ; MT, 26 September, 1 $ ; Kwangtung Prov., Luichow Peninsula, 8 September,

IS.

The sculpture on ocellocular area agrees with the figure shown
by Tkalcu (1968a, Fig. 70). Male antenna and genitalia are il-
lustrated in Fig. 7. The hair color of both sexes agrees with that
given by Bishoff (1936) :

July 1972]

SAKAGAMI — CHINESE BUMBLE BEES

165

Worker. — Head, mesosoma and legs dark chocolate brown. Ill-defined collaris
and scutellaris, coxa II below, trochanters, femora basally (to apex in femur
III), tergum I, sterna whitish; other terga orange red (probably bright fer-
ruginous when fresh) . Mesosomal side below broadly whitish with admixture of
dark hairs. Tergum I with sparse admixture of dark hairs.

Variation is seen in the degree of admixture of dark hairs to
whitish parts, resulting in different contrasts of collaris, scutellaris
and pleuralis though collaris is mostly separated from head by a
narrow obscure dark band. A single worker from Luichow Peninsula
possesses pleura nearly whitish with admixture of dark hairs only
medially, otherwise not markedly differing from the other specimens,
that is, not showing color pattern of B. p. honei (Bischoff, 1936) and
without structural difference from the other specimens.

Male. — Head, mesosoma, legs and tergum I ochraceous yellow, other terga
orange red. Sparse dark hairs on vertex, gena, face especially along eyes, and
on interalar areas. Metasoma in poor condition, so that admixture of dark hairs
not precisely examined. Mandibular beard blackish brown, partly with honey
yellow hairs.

Bombus (Pyrobombus) flayescens Smith

Bombus flavescens Smith, 1852: 45, $.

Bombus ( Pratobombus ) flavescens , Pittioni, 1949: 259, $, 2, $•

Specimens examined. — HS, 9 August, 1 $ .

A single specimen of this extremely polytypic species goes to B. /.
f. geei Cockerell in the key by Pittioni.

Hair color. — Chocolate brown ; tergum IV except anterior margin, and
following terga, legs and sterna dirty orange yellow. Mandibular beard pale
orange yellow, short hairs of face whitish to grayish yellow above antenna, more
yellowish below, long facial hairs mostly dark, with admixture of yellowish hairs,
clypeus with yellowish long hairs; small patches of fulvous hairs on terga I
and II laterally and on tergum II medioposteriorly.

Bombus (Pyrobombus) flavus Friese

Bombus pratorum ssp. flavus Friese, 1904: 517, $ ; Friese & v. Wagner 1909:

52, $.

Bombus parthenius var. picipes Richards, 1934: 90, 2 (Tkalcu in litt .) .
Bombus ( Pratobombus ) klapperichi Pittioni, 1949: 266, $ ; f. intermedius:

268, 2 ; f. flavescentiformis : 269, 2 •

Specimens examined. — SP, 20-29 August, 7 2, 2 $ , 17 September, 1 2 ; HS,
9-10 August, 2 2 , 12-16 August, 6 2 ; LK, 31 August, 2 2 , 4-10 September,
1$,52,2$; MT-LP, 28 September, 1 2 ; Kwangtung Prov., Luichow Peninsula,
8 September, 2 2 •

166

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48 , NO. 3

The specimens examined were identified as B. flavus, a long for-
gotten taxon. The species is close to B. modestus Eversmann and
B. beaticola (Tkalcu). Three castes are redescribed below to avoid
any further confusion.

All castes.- — Integument black, hind tibia and all tarsi dark chestnut brown.
Wings pale brown and hyaline, veins dark brown. Hair coat relatively long,
erect and rather uneven.

Queen. — Body 16 mm, forewing 14 mm. Hair color (Fig. lOd, I). — Head dark
to chocolate brown, with short hairs on face and sparsely on occiput fulvo-
testaceous. Interalaris broad and dark to blackish brown. Collaris virtually
absent, represented only by sparse admixture of yellowish hairs. Scutellaris
narrow but distinct, ochraceous yellow as mesosomal side. Metasomal tergum I
entirely and anterior half of tergum II, broadly at middle, narrowly at side,
ochraceous yellow, with sparse admixture of dark brown hairs; remainder of
tergum II, III and anterior two thirds of IV blackish brown; tergum IV
posteriorly, V, and sparse hairs on VI faded ochraceous orange. Sterna I ^ III
marginally pale fulvous, IV ^ V as tergum V, all sterna with admixture of dark
hairs, denser laterally. Legs dark brown, coxae, trochanters and femora basally
below ochraceous yellow.

Structure . — Upper, maximum and lower interorbital distances, 36:46:42. Supra-
orbital punctures (Fig. 8M) 1.0 ~ 1.5 times larger than diameter of facets,

slightly larger than, but essentially similar to those of B. beaticola, dense and
rather irregular; interspaces weakly coriaceous and moderately shining, as wide
as to two times wider than punctures; punctures on paraorbital corridor sparse,
interspaces smooth and shining; postsupraorbital punctures dense, as large as
or smaller than facets, interspaces average narrower than punctures, not
becoming wider along postorbital margin; dorsal furrow distinct; suprafrontal
punctures coarse and sparse, 2.0 ^ 2.5 times larger than facets. Relative measure-
ments of vertex: Ocellocular distance 22, postocellar distance 25, ocelloccipital
distance (cf. Fig. 5P) 25, distance between mid and lateral ocelli 6, diameters
of mid and lateral ocelli 13, 12. Malar space nearly as long as wide, smooth
except sparse, minute and ill-defined punctures on area slightly above mandibular
base. Clypeus below with punctures slightly denser than in B. beaticola (Fig.
8E, F). Labrum as in B. beaticola-, tubercles flat, shining but moderately
coriaceous, with ill-defined punctures except coarser ones on lower slope; median
furrow well depressed, granulate with fine and dense punctures; lamella dis-
tinctly curved, about one-third as wide as labrum. Mandible with basal punctures
much denser than in B. beaticola (Fig. 8A-D) , incisura lateralis distinct.
Length : width ratios of flagellar segments : I (23:13.5), II (16:14), III (18:13.5),
III slightly longer than in B. beaticola (III nearly 1 w) (Fig. 9N, P).
Median impunctate area of mesoscutum smooth and shining, maximum width
about one-third distance between end of median line and posterior margin of
sclerite. Metasomal terga finely tesselate but rather shining especially on IV and
V, where punctures equal or slightly exceed size of facets, well defined and
rather sparse with interspaces about 3 — ’4 times puncture diameter except
marginally where punctures denser and interspaces 2.0 ^ 3.5 times puncture
diameter. Epipygium flat, granulate and weakly elevated, coriaceous with fine

July 1972]

SAKAGAMI — CHINESE BUMBLE BEES

167

Fig. 8. Comparison of B. flavus (A,C,E,G,HJ 5 K)M) and B. beaticola (B,D,F,L).
A,B. Base of right mandible (queen) ; C,D. Base of left mandible (worker) ;
E, F. Punctation on lower part of worker clypeus; G,H. Apex of male
genitalia dorsolateral and dorsal views; J. Inner depression of gonostylus;
K,L. Apex of penis valve; M. Sculpture of ocellocular area (queen).

163

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Fig. 9. Bombus flavus, continued. N,0,P. Basal flagellar segments (queen,
male, worker) ; Q, R. Male hind and mid basitarsi; S. Male hind tibia.

and dense punctures. Hypopygium weakly ridged but nor carinate. Hind tibia
shining but microscopically alutaceous except marginally and basally. Mid
basitarsus shagreened but shining, posterior margin nearly straight. Hind
basitarsus shagreened even basally, though less conspicuously than mid basi-
tarsus, posterior margin gently curved.

Worker. — Body 11 — 1 13 mm, forewing 9.5 ~ 10.5 mm. Structure. — Similar
to queen except certain caste-linked reductions of sculpture. Hair color.-— Head
as in queen but short hairs paler, occiput and gena behind with more ochraceous
or yellowish long hairs. Other body parts ochraceous to yellow, with admixture
of dark brown hairs in variable degrees on mesosomal disc and metasomal terga
as shown later ; terga IV ^ V often more or less with dull orange tone in parallel
with increased admixture of dark hairs on tergum III. Tibiae and tarsi dark
brown; corbicular fringes ochraceous with admixture of dark brown hairs,
especially posteriorly (Fig. lOd) .

Male. — Body 11 ' 13 mm, forewing 9.5 ~ 10.5 mm. Hair color. — Uniformly

pale ochraceous to citron yellow. Mandibular beard orange yellow. Sparse dark
brown hairs on vertex, gena, face along inner orbit and mesosomal disc, slightly
denser centrally but rather uniformly distributed (Fig. lOd, III) . Terga apically
slightly paler, with no admixture of dark hairs.

Structure. — Upper, maximum and lower interorbital distances, 30:36:32.
Supraorbital area essentially as in females, punctures slightly larger than facets,
rather ill-defined though slightly more distinct than in B. beaticola ; parafacettal
corridor only sparsely punctured. Relative measurements on vertex : Ocellocular
distance 18, postocellar distance 20, distance between mid and lateral ocelli
5, diameters of mid and lateral ocelli 10, 9. Malar space as in females but
punctures finer, length : width as 25:21. Length : width ratios of flagellar
segments :I (17:11, basal width 6), II (12:11), III (15:11) (Fig. 90). Median
impunctate area of mesoscutum smooth and very narrow, with sparse punctures
along median line. Terga as in B. beaticola. Hind tibia well corbiculated (Fig.

july 1972]

SAKAGAMI — CHINESE BUMBLE BEES

169

9S), fringes longer than width of segment, upper surface slightly convex basally,
flat apically, smooth and shining, slightly shagreened at base and posterior
margin; apical bristles exceeding apex. Mid basitarsus (Fig. 9R) shining but
shagreened, with hind margin slightly tapering apically. Hind basitarsus (Fig.
9Q) finely shagreened but more shining than mid basitarsus; posterior margin
slightly tapering apically. Stipes wider than in B. beaticola, curvature of inner
margin as in B. modestus, not B. beaticola , outer apical angle more distinctly
curved than in B. beaticola (Fig. 8G, H). Penis valve rather parallel sided,
apical hook with outer margin distinctly rounded and basally rather widened,
tapering apically (Fig. 8H, K, L) .

The identification of this species was possible by the courtesy of
Dr. B. Tkalcu. The original description is very short, only, “Asia
centr., einfarbig gelb behaart.” The males are structurally identical
to the type as defined by a note and pencil sketches which Dr. Tkalcu
made from the type, and sent to me years ago for a different purpose,
and by another more detailed note sent by him according to my
request. The unique difference is the presence of sparse dark hairs
in my specimens, which could be interpreted as an extreme of variation
range. According to Dr. Tkalcu, the specimen which he labelled as
lectotype as preserved in the collection of the Zoologisches Museum
der Humboldt Universitat Berlins, with the following four labels :
1) Turkestan Kashgar 22. 7. 1900. 2) Bombus pratorum v. flavus
Fr. 8 1904 Friese det. 3) Bombus pratorum v. flavus Fr. 4) Red
label printed “type.” The genitalia is separately pinned with a label
as in 2) shown above. Dr. Tkalcu informed me that the locality was
false, the real locality being probably Szechuan or adjacent territories
(cf. Tkalcu, 1968a, p. 49, footnote) . He also examined another nearly
identical specimen from Tien-mu-shan, China. Meanwhile, a single
queen examined by me agrees well with the description of B. klap-
perichi Pittioni described from Fukien Province both in coloration
and structure, except for; 1) Smaller size (17 ^ 18 mm in B.
klapperichi) . 2) Inner orbits slightly convergent below (parallel

in B. klapperichi). 3) Distance between mid and lateral ocelli slightly
longer (ratio to diameter of lateral ocellus 12:6 in B. flavus , 12: > 6
in B. klapperichi ). 4) Flagellum I 2w > 1 in B. flavus 2w = ca.l in
B. klapperichi. These differences are regarded as lying within the
variation range and 2) and 3) could be interpreted as an outcome
of allometric change as shown in B. trifasciatus. Further the worker
coloration is variable but approximately corresponds to B. f. flavescenti-
formis. Consequently it is concluded that B. flavus, so far known from
the male alone, is conspecific with B. klapperichi, known only from

170

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Fig. 10. Color variation in B. flavus. a. Mesosomal dorsum; b,c. Metasomal
terga III and IV ; d. Selected color patterns. I. Queen, II. Male type specimens
(from note by Dr. Tkalcu), III. Male recorded in text; Other showing
representative patterns in worker.

the female, and the species has a wide distribution from Kwangtung to
Szechuan, probably throughout Central and Southern parts of China,
forming a southern and continental vicariant of B. modestus in Siberia
and B. beaticola in Japan. Comparing with the synoptic table of the
latter two species by Tkalcu (1968a, p. 33), B. flavus is closer to B.
modestus in relative lengths of flagellar segments and shape of stipes,
but closer to B. beaticola in the less shagreened hind basitarsus.

In coloration the queen is comparable to B. beaticola moshkara-
reppus Sakagami et Ishikawa from Hokkaido, Northern Japan, by the

July 1972]

SAKAGAMI — CHINESE BUMBLE BEES

171

prevalence of dark hairs on the face and mesosomal disc, but the
pleura are broadly pale as in B. b. beaticola (Tkalcu) from Central
Japan (Sakagami and Ishikawa, 1969). The variation in workers is
mainly caused by the degree of admixture of dark hairs on the meso-
somal disc and metasomal terga III and IV, which positively cor-
relates with an increase of dark hairs on head and corbicular fringes,
as well as intensified orange tone on apical terga. The variability
in mesosomal disc and metasomal terga III and IV, shown in Fig.
10a c, results in various types of combinations, the frequency
distribution of which in the specimens in good condition is: AAA
(6 specimens), ABA (2), BAA (1), AAB (2), ABC (1), ABD (1),
BCD (4), CCD (5), showing the presence of two dominant types,
a paler type (AAA) and a queen like type (BCD, CCD). Thus the
variation trend is precisely similar to that of B. beaticola (Sakagami
and Ishikawa, 1969).

Psithyrus (Metapsithyrus) pieli Maa

Psithyrus ( Metapsithyrus ) pieli Maa, 1948: 29, $, $.

Specimens examined. — LK, 9 September, 1 $ , 1 $ .

Dr. B. Tkalcu suggested this possible identification. Both specimens
were in poor condition.

Discussion

Excluding Psithyrus and the specimens collected in Luichow Penin-
sula, Kwangtung Province, all specimens examined are tabulated
according to relative abundance and seasonal distribution as follows:

Number of specimens ($-£-$)

July August September Total

late early mid late early mid late (444)

B. trijasciatus
B. imitator
B. flavus
B. pyrosoma
B. ignitus
B. atripes
B. breviceps
B. opulentus
B. kulingensis
B. flavescens

0-25-0 0-27-0 0-77-1 0-83-0 0-88-3 0-23-3 0-7-1 0-330-7 337

0—2-0 0-1-0 0-1-1 0-11-3 0-1-1 0-8-3 0-27-8 35

0-2-0 0-6-0 0-7-2 1—5-2 0-1-0 0-1-0 1-24-4 29

0-2-0 0—1-0 0—9-1 0—1-0 0-1-0 0 — 14-1 15

0—5-0 0-3-0 0-1-0 0-3-0 0-12-0 12

0—2-1 0-3-0 0 5-1 6

0-2-1 0 2-1 3

0-0-1 0-1-1 0 1-2 3

1-2-0 0 2-0 3

0 - 0-1 0 — 0-1 1

The relative abundance is strongly influenced by the overwhelming
predominance of B. trijasciatus (75.5 % of all specimens), followed

172

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

by B. imitator (7.8 %) and B. flavus (6.5 %). Although the collecting
was limited to late summer to early fall, the relative abundance may
reflect the quantitative makeup of the local fauna because of the long
annual cycle of most bumble bees. Exceptions are expected in Pyro-
bombus. It is likely that some short-cycle species of this subgenus were
not collected. The capture of males of B. flavus in late August and
early September suggests that this species possesses a life cycle relatively
longer than other Pyrobombus. But the capture records of males
of an allied species, B. beaticola in Japan (Honshu, records at
altitudes less than 2,000 m) show a remarkable variation: late May
(1), late June (1), early July (1), mid July (12), late July (18),
early August (18), late August (6), early September (17), mid
September (2). Recently Meidell (1968) asserts that Bombus ( Pyro-
bombus ) jonellus (Kirby) possesses two generations per year in
Norway. Naturally such opinion cannot uncritically be generalized
to other species showing a wide variation of male appearance. But
it is interesting to pursue this problem in various species of Pyro-
bombus, many of which are characterized by an ephemeral colony
life which can be studied in part by checking the records of preserved
specimens.

It is still premature to discuss the zoogeographical position of the
bumble bee fauna of the area considered, which belongs, in the
system by Panfilov (1957), to the Central Chinese Province in the
Japan-China-Himalayan Subregion. He comments that the few studies
in this province indicate only B. atripes, B. ( Adventribombus ) unicolor
Friese and B. ( Hortobombus ) supremus F. Morawitz as typical of the
area. Most species examined by me are those endemic to China
Proper including: B. trifasciatus, B. atripes, B. imitator, B. opulentus,
B. flavus, B. pyrosoma, B. kulingensis and P. pieli. One species, B.
ignitus, reaches Japan and two, B. breviceps and B. flavescens South
East Asia. On the other hand, no Euro-Siberian elements in the
broadest sense are recognized, though some species are regarded
“secondarily” as belonging to northern groups (B. flavus and B.
opulentus ) .

Among the bumble bees studied, there is no instance of strict
sympatric isochromy as recorded in South East Asia (e.g. Tkalcu
1968c) . The patterns recognized in the area studied are divided as
follows :

1) Pale mesosoma with dark interalaris and trichromatic (pale,
dark, reddish) metasoma. To this pattern belong the following

july 1972]

SAKAGAMI — CHINESE BUMBLE BEES

173

species with deviations from the model as given parenthetically: B.
trifasciatus (2, 2, $. Interalaris often expanded), B. kulingensis
(2, 9, S unknown. Interalaris expanded), B. flavus (2 and some
2 ) , B. imitator (2, 2, S. Interalaris often weakened). B. ignitus
( $ , though not collected), B. breviceps (2, 2, $ . Pale hairs distinctly
darker), B. pyrosoma (2, $. Considerably deviated from the model
by weakening of interalaris, absence of dark metasomal band and
in workers pale hairs nearly whitish).

2) Black with red tail. Widespread in the Palaearctics, isolately
appearing in some Andean species. B. ignitus ( 2 , 2 ) , B. flavescens
(partly) .

3) Predominantly pale. B. flavus (S, some 2), B. flavescens
(partly, though not captured from the area) .

4) Bright orange fore body with dark venter, with or without
dark apical terga. B. opulentus , B. atripes.

From this tabulation, it is easily recognized that the first mentioned
pattern prevails in the area studied.

Literature Cited

Bischoff, H. 1936. Schwedisch-chinesische Wissenschaftliche Expedition nach
den nordwestlichen Provinzen Chinas, unter der Leitung von Dr. Sven
Hedin und Prof. Sii Ping-Chang. Insekten gesammelt von schwedischen
Arzt der Expedition Dr. David Hummel 1927-1930. 56. Hymenoptera.
10. Bombinae. Ark. Zool., 27A(38) : 1-27.

Cockerell, T. D. A. 1917. Two new humble-bees from China. Entomologist,
50: 265-266.

Friese, H. 1904. Neue oder wenig bekannte Hummeln des Russischen Reiches
(Hymenoptera). Ann. Mus. Zool. Acad. Sci. Petersbourg, 9: 507-523.
1909. Neue Varietaten von Bombus (Hymenoptera). Deut. Entomol. Z.
1909: 673-676.

Friese, H. and F. v. Wagner. 1909. Zoologische Studien an Hummels. I. Die
Hummeln der deutschen Fauna. Zool. Jahrb. Abt. Syst. Geogr. Biol.
Tiere, 29: 1-104.

Frison, T. H. 1935. Records, notes and descriptions of Bremus from Asia
(Bremidae: Hymenoptera). Rec. Indian Mus. Calcutta, 37: 339-363.
Gressitt, J. L. 1953. The California Academy-Lingnan Dawn-redwood Expedition.

Proc. Calif. Acad. Sci., 4th Ser., 28: 25-58.

Gribodo, G. 1891. Contribuzioni imenotterologiche sopra alcune species nouve
o poco conosciente di Imenotteri Antofili. Boll. Soc. Ent. Ital., 23:
103-119.

Maa, T. C. 1948. On some Asiatic species of the genus Psithyrus Lepel.

(Hymenoptera: Bombidae) . Notes Entomol. Chin., 12: 17-37.

Meidell, 0. 1968. Bombus jonellus (Kirby) (Hym. Apidae) has two generations
in a season. Norsk Entomol. Tidsskr., 14: 31-32.

Morawitz, F. 1890. Insecta a Cl. G. N. Potanin in China et in Mongolia
novissime lecta. II. Horae Soc. Entomol. Ross., 24: 349-385.

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48 , NO. 3

Moure, J. S. and S. F. Sakagami. 1962. As mamangabas sociais do Brasil
( Bombus Latr.) (Hym. Apoidea) . Stud. Entomol., 5: 65-194.

Panfilov, D. V. 1957. 0 geograficheskom rasprostranenii shchmelej ( Bombus )
v Kitae. Acta Geogr. Sinica, 23: 221-239 (Chinese with Russian
summary) .

Pittioni, B. 1949. Beitrage zur Kenntnis der Bienenfauna SO-Chinas. Die
Hummeln und Schmarotzerhummeln der Ausbeute J. Klapperich ( 1937—
38) (Hymenoptera, Apoidea, Bombini). Eos, 25: 241-284.

Richards, 0. W. 1934. Some new species and varieties of the oriental bumble-
bees (Hym. Bombidae). Stylops, 3: 87-90.

1968. The subgeneric division of the genus Bombus Latreille (Hymenoptera;
Apidae) . Brit. Mus. (Natur. Hist.) Bull. Entomol., 22: 211-276.
Sakagami, S. F. and R. Ishikawa. 1969. Note preliminaire sur la repartition
geographique des bourdons japonais, avec descriptions et remarques
sur quelques formes nouvelles ou peu connues. J. Fac. Sci. Hokkaido
Univ. Ser. VI, Zool., 17 : 152-296.

Sakagami, S. F. and J. S. Moure. 1965. Cephalic polymorphism in some Neo-
tropical halictine bees (Hymenoptera — Apoidea). An. Acad. Brasil.
Cienc., 37: 303-313.

Skorikov, A. S. 1914. K faun’ shchmelej yzhnoi Tsast’ Primorskoj oblasti.
Russk. Entomol. Obozrenie, 14: 398-407.

1933. Zur Hummelfauna Japans und seiner Nachbarlander. Mushi, 6:
53-65.

Smith, F. 1852. Descriptions of some new and apparently undescribed species
of hymenopterous insects from North China, collected by Robert
Fortune, Esq. Trans. Roy. Entomol. Soc. London, (N. S.) 2: 33-45.
1861. Description of new genera and species of exotic Hymenoptera. J.
Entomol., 1: 146-155.

1869. Descriptions of Hymenoptera from Japan. Entomologist, 62: 205-208.
Tkalcu, B. 1960. Remarques sur quelques especes de bourdons de Chine.
Bull. Soc. Entomol. Mulhouse, Sept.-Oct. : 66-71.

1961a. Zwei chinesische Hummel- Arten (Hymenoptera, Bombinae) . Acta Soc.

Entomol. Cechoslov., 58: 45-59, 2. Taf.

1961b. Zur Hummelfauna der Umgebung Kuku-Nors (Hymenoptera, Bom-
binae) . Ibid. 58: 344-379, 2. Taf.

1962. Contribution a 1’ etude des bourdons du Japan I. Bull. Soc. Entomol.
Mulhouse, Nov.-Dee.: 81-100.

1968a. Neue Arten der Unterfamilie Bombinae der palaarktischen Region
(Hymenoptera, Apoidea). Acta Soc. Entomol. Cechoslov., 65: 21-51.
1968b. Revision der Arten der Untergattung Tricornibombus Skorikov (Hymen-
optera, Apoidea). Acta Rer. Natur. Mus. Nat. Slov. Bratislava, 14:
79-94-.

1968c. Revision der vier sympatrischen, homochrome geographischen Rassen
bildenden Hummelarten SO-Asiens. Annot. Zool. Bot. Bratislava,
52: 1-31.

Wu, C. F. 1941. Catalogues insectorum Sinensium. VI, Hymenoptera, 333pp.

July 1972]

SCHLISING — BEHAVIOR OF DIADASIA

175

Foraging and Nest Provisioning Behavior of the Oligolectic

Bee, Diadasia bituberculata

(Hymenoptera: Anthophoridae)

Robert A. Schlising

Pasadena, California 91101

Among bees, oligolecty — a type of flower constancy in which the
females of a species restrict their pollen-food foraging to a narrow
range of plant types — is receiving considerable attention and is be-
coming fairly well known (Linsley and MacSwain, 1958; Stephen,
Bohart and Torchio, 1969). Bees in the anthophorid genus Diadasia
Patton are oligolectic (Linsley and MacSwain, 1957, 1958), most
species restricting their pollen foraging to plants in the Malvaceae
( Sphaeralcea , Callirhoe, Sidalcea, or Sida ) . Other species in this
genus are oligolectic on Opuntia (Cactaceae), Helianthus (Compositae) ,
Clarkia (Onagraceae) , and Convolvulus (Convolvulaceae) . One species,
Diadasia bituberculata (Cresson), is known to be oligolectic on
Convolvulus Linnaeus (many species of which have recently been
transferred to the genus Calystegia Robert Brown) . The purpose of
the present study was to observe details of the pollen foraging and
nest provisioning of this one bee species on one species of Calystegia
(formerly Convolvulus ) .

Oligolectic bees commonly show inherited morphological and be-
havioral features which adapt these species for existence on a restricted
number of food plants (Linsley, 1958). In Diadasia bituberculata
these include elongated, specialized mouthparts that permit extraction
of nectar from flowers with deep tubular corollas (Linsley and Mac-
Swain, 1958), as well as seasonal and diurnal synchronization with
flowering of its host plants. Michener (1951) lists the range of D.
bituberculata as “California,” an area in which several species of
Convolvulus and Calystegia occur (Munz and Keck, 1959; Munz,
1968) . Although different populations of D. bituberculata throughout
its range utilize different species of Convolvulus or Calystegia as their
pollen source, this report is limited to observations of these bees at
flowers of Calystegia fulcra ta (Gray) Brummitt at one locality.

This study was conducted in 1971, along Mineral King Road, 3.9
miles southeast of the junction with State Highway 198, near Three
Rivers, Tulare County, California. This locality, in the hot dry
foothills of the Sierra Nevada, at an elevation of 610 m, is covered

The Pan-Pacific Entomologist 48: 175-188. July 1972

176

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

by chaparral with dense shrubs often up to 3 or 4 m tall and trees
considerably taller. The commonest woody plants on the steep slopes
and in the draws here include: Adenostoma fasciculatum Hooker and
Arnott, Aesculus calijornica (Spach) Nuttall, Ceanothus cuneatus
(Hooker) Nuttall, Eriodictyon californicum (Hooker and Arnott)
Torrey, Fremontodendron californicum Coville, Mimulus longiflorus
(Nuttall) Grant, Quercus chrysolepis Liebmann, Rhus diver siloha Torrey
and Gray, and Umbellularia calijornica (Hooker and Arnott) Nuttall.
Both the bees and their “wild morning glory” food plants occur
throughout the area — usually along rock outcroppings or other open
and grassy spots in the chaparral, or along roadsides and fire trails.

The main pollen source of the bees here was a Calystegia population
that occupied nearly 50 m of a narrow strip of east-facing roadbank,
and grew with very few annual grasses and native herbs (Fig. 1).
The creamy-white, nearly scentless flowers of C. fulcrata are tubular
at the base, and flared or trumpet shaped at the top (Fig. 2).
Nectar is secreted at the very base of the corolla tube in five nectary
areas; pollen is located on anthers near the top of the tube. These
two insect foods are available to foragers only from the early morning
opening of the flowers to their wilting and closing by afternoon or
earlier. The flowers last but one day (new flowers open each morning),
so the total sequence of flower visitors during an individual flower’s
life can be rather easily studied. The flowering peak was in late May
(Table 1), and flowering lasted about seven weeks in 1971, with a
few flowers open on 6 May and none remaining on 5 July. Flowers
were rather uniformly spread out over the population, but in several
areas where the vines were particularly dense, there were as many as
25 to 30 flowers per m 2 .

The nesting site of the bees was located about 200 m northwestward,
around a corner and out of sight of the Calystegia food plants. Nests

->

Fig. 1 . Location of Calystegia fulcrata population, on lower chaparral-facing
slopes of roadside bank, from foreground to curve of road in distance. Fig. 2.
Stems and flowers of C. fulcrata (X 0.25). Fig. 3. Diadasia bituberculata nest
site, with most burrows in three areas to left of arrows. Fig. 4. D. bituberculata
female, with full scopal loads of C. fulcrata pollen (X 3). Fig. 5. D. bituberculata
female from rear, showing dense plumose hairs of scopae, with very few pollen
grains to left of arrow ( X 4) . Fig. 6. Two turrets, amid dried annuals at top
of bank in Fig. 3, with turret entrance and top of tumulus to left of arrows
(X 0.25). Fig. 7. Typical position (posed specimen), of D. bituberculata male
in C. fulcrata flower, witth tongue reaching nectaries and dorsum of abdomen
brushing stamens and (outlined) stigma (X 2).

July 1972]

SCHLISING — BEHAVIOR OF DIADASIA

177

178

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Table 1 . Dates and number of hours of observation in 1971, noon
air temperatures (°C in shade 0.5 m above flowers and nests), and
number of open Calystegia flowers in the main population.

Date

Total

hours

Temperature

Open

flowers

May

12

2.5

—

121

13

5.0

22.5

235

15

7.0

22.0

360

16

7.25

17.0

375

24

7.5

27.5

392

25

6.0

28.5

373

June

4

6.25

24.0

285

5

8.75

27.5

279

10

2.0

-

-

11

8.75

28.5

226

12

6.0

32.5

223

were located in an exposed southeast-facing bank of the roadcut that
was 3 m from the road and nearly devoid of vegetation (Fig. 3).
A few plants of C. fulcrata grew on this bank, and several more
occurred 50 m away on a fire trail, but it is believed that the nesting
bees observed here primarily utilized the large population 200 m
to the southeast.

The dates and numbers of observation hours here are listed in
Table 1, along with noon air temperatures. No measurable precipitation
occurred during the hours of observation, but weather records (U. S.
Dep. Commer., 1971) for Three Rivers (elevation 290 m) show that
the 7.37 cm of precipitation that occurred in May was 5.13 cm
above the normal (based on a 30-year average) for the month; June
precipitation was zero, or 0.53 cm below normal. Total annual
precipitation averages 53.90 cm here. Most study days were sunny,
with moderate breezes in the afternoons. The bees, particularly at
the nest site, were very docile and could be watched at very close
range. They appeared to be totally unaffected by observers sitting
only decimeters away.

Female Foraging Behavior

Females of Diadasia bituber culata are completely reliant on Calystegia
fulcrata for nest provisioning at this site, and emergence from nests
and the beginning of foraging activity is synchronized with the frist
availability of pollen. Anther dehiscence, which is regulated in some

july 1972]

SCHLISING — BEHAVIOR OF DIADASIA

179

species by air temperature and relative humidity (Percival, 1965),
was usually very early near Three Rivers. On 5 June the first female
was seen at 5:10 a.m. (Pacific Standard Time), when some powdery,
freshly dehisced pollen was available on the anthers, and in the next
ten minutes visits by females became abundant. An overcast and
humid morning inhibits both anther dehiscence and foraging activity
of the bees. On 16 May, for example, flowers were fully open by
5:15 a.m. when it was light, but there was a heavy cloud cover and
the air, at 10° C, was very moist with a gentle mist; female bees
did not come to flowers that morning until 10:40, when the first
anther dehiscence occurred. Nectar presentation seems to occur as or
before the anthers dehisce.

A foraging female typically flies directly to, and “dives headlong”
into a Calystegia flower, where she spends one to eight (rarely up to
30) seconds taking nectar by inserting her tongue into one to five
of the nectaries at the base. Early in the morning the females may
first probe the nectaries for a second or more in nearly each flower
visited, but later in the day they may take nectar from fewer of the
flowers visited (Table 2). Early in the morning each female spends
an average of at least 2.5 times longer collecting pollen in individual
flowers than she does later in the day. Mean pollen-collecting time
for 6 bees was 23 seconds per flower at 5:30-6:30 a.m. and 9 seconds
at 9:30-10:30 on 5 June (Table 2). Although Linsley and MacSwain
(1957) noted that the elongated, specialized mouthparts of this species
(Fig. 4) permit simultaneous gathering of pollen and nectar from
Convolvulus flowers, females at this site very rarely took pollen from
the anthers while also taking nectar. They consistently climb onto
the anthers (which are well elevated above the nectar bearing areas)
for pollen gathering after taking nectar — or they settle immediately
on the anthers without taking nectar.

When collecting pollen, females usually position themselves on the
outer surfaces of the anthers or slightly below them on the filaments,
where they can rotate their bodies (sometimes buzzing loudly) inside
the flower, one to three times around the cluster of five anthers.
Pollen is removed with the forelegs and transferred to the scopae — the
dense, plumose hairs on the hind legs (Figs. 4, 5). Occasionally a
female sits on top of the five anthers a few seconds, grooming pollen
from her face and thorax, but usually she flies from the flower imme-
diately after taking pollen.

A female does not complete her pollen load with pollen from one
flower only, but leaves a flower still bearing abundant pollen for other

180

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Table 2. Representative timings (in seconds) of foraging by 4
females of Diadasia bituberculata in flowers of Calystegia fulcrata
on the morning of 5 June.

Bee

Flowers

visited

Nectar 1

Pollen

Resting

l b

1

8

61

0

2

0

20

0

3

4

41

0

4

0

47

0

5

0

18

0

6

0

28

8

0

7°

0 d

2 b

1

7

45

4

2

0

20

0

3

3

10

0

4

2

20

6

5

1

19

0

6

3

20

0

7

3

43

0

8

0

3

0

9

3

4

0

10

5

36

3

0

6°

0

11

2

32

9 d

3 e

1

0

8

0

2

2

65

0

3

0

9

0

4

0

8

0

5

0

6

0

6

0

6

0

7

0

5

0

8

0

10

0

9

0

2

0

10

0

10

2

11

0

8

0

4 e

1

0

3

0

2

0

3

0

3

0

9

0

4

0

5

0

5

0

5

0

6

0

13

0

7

0

3

0

8

0

3

0

a Nectar collection always preceded pollen collection in the same flower.

15 Observed between 5:30-6:30. Bee started foraging with little or no pollen on scopae.
c Bee continued pollen collection after rest in same flower.
d Bee left flower (probably for nest) with full scopae.

e Observed between 9:30-10:30. It was possible to record only short series of uninterrupted
flower-visit sequences at this time of day, due to the very fast and erratic flight patterns of the bees.

JULY 1972]

SCHLISING — BEHAVIOR OF DIADASIA

131

Calystegia flowers. To obtain “full” scopal loads of pollen (Fig. 4)
a female may visit from five or six to perhaps 15 flowers early in
the morning. When flowers are nearly depleted of pollen later in the
day up to 30 or more flowers are visited to obtain loads, and some
loads brought to nests later are much smaller. During the main pollen
collecting period females may fly to, but immediately desert, flowers
that still have pollen in them. Most flowers are frequently visited and
foraged in — often by one bee a few seconds after another.

The times at which pollen foraging cease vary, apparently with
weather conditions (and with times at which foraging starts) and
with the number of females of Diadasia collecting from the limited
number of flowers at one locality. In mid May, with lower air
temperatures (Table 1), and possibly fewer females present, re-
maining pollen was still being collected at 3 p.m. — even if the flowers
were already partially folded shut and wilted. But later in the season
pollen supply was depleted and foraging was completed by about
12:30 p.m. on 24 May or even by 9:30 a.m. on 12 June. When the
females cease pollen foraging (even as early as 9:30) they start nest
construction, although visits to Calystegia flowers for nectar may
continue through the afternoon. After pollen foraging was over for
the day females were also seen utilizing three other herbaceous
species for nectar (but not pollen) near the nest site: they were
occasional in flowers of Penstemon laetus Gray and rare in flowers
of Clarkia speciosa ssp. polyantha Lewis and Lewis and Calochortus
superbus Purdy ex J. T. Howell.

All Calystegia fulcrata flowers examined at this site had the pollen
removed by bees — probably by D. bituberculata in all cases — during
study days, and there is no doubt that bees of both sexes, and partic-
ularly the females, bring about pollination of the flowers. During
pollen collection, the style and stigma project above the parts grasped
by the bees. But stigmas are frequently brushed as females enter,
leave, or forage in flowers, and pollen is found lodged between the
weakly appressed stigmatic lobes after Diadasia visits. These flowers,
during their one-day life, commonly receive as many as 50 individual
visits by the bees. Similar high numbers of bee visits per flower have
been reported for tropical oligolectic bees closely related to Diadasia ,
to flowers related to Calystegia (Schlising, 1970). The situation at
Three Rivers is remarkable however, because it is nearly individuals
of one species alone — D. bituberculata — that exploit and pollinate the
plants.

132

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 43, NO. 3

Female Nest Provisioning Behavior

At Three Rivers these bees utilized three areas of an eroded bank
(Fig. 3) for their nests: 1) the nearly horizontal surface at the top,
covered with short grasses and other low annuals (Fig. 6), 2) the
exposed, nearly vertical, mineral soil surface, and 3) the nearly level
deposits of soil washed down, sparsely covered with annual grasses.
The densest aggregation of nests — 27 per m 2 — was on this flatter
area only 2 m from the road edge. Most burrows had curving or
straight horizontal turrets made of dried mud pellets covering the
burrow entrances (Fig. 6) ; turrets varied from about 1.0 to 2.5
cm long, and entrances faced every direction.

Morning emergence of females from nests was observed on 5 June
at 4:50 a.m., when the air temperature 20 cm above the nests was
10.5° C. The first bee seen to emerge flew in a small circle over the
nests, then slowly to a Calystegia flower on a bank 2 m away, where
she rested in the sun, took nectar, and finally pollen. During the next
15 minutes other females emerged, and by 5:15 were abundant at
the main Calystegia population, collecting both nectar and pollen.
Observations made during the night and also as these beees emerged
indicate that surface plugs are not constructed to close burrow entrances
when occupied by females at night.

When returning to her nest with a pollen load in the morning,
a female typically enters her turret (or her burrow, if it has no
turret), until she is barely but entirely within it (out of the sun),
with her hind pollen-laden legs resting on the dorsum of her abdomen.
Then, with a twitching of the hind tarsi the only visible motion of
her body, she rests here for 0.25 to 7 (occasionally up to 15 or 17)
minutes. Then she crawls down into the nest, and 4 to 8 squeaky
buzzes, each about one second long, are heard as she removes the pollen
from her scopae. After these buzzes, a female usually appears at the
burrow entrance in a few seconds, and flies away immediately.

Such provisioning behavior is repeated throughout the foraging
period. Variation in timing among different foragers is of interest,
since all females were probably collecting pollen from the Calystegia
200 m from the nests. Representative provisioning timings are shown
in Table 3 for four females followed over a 2-hour period on 5 June.
Bee #1 averaged 8.0 minute trips in collecting apparently “full” loads
of pollen (Fig. 4), while bees #2, #3 and #4 averaged 9.6, 14.2
and 21.0 minute trips in collecting similar loads. These same four
bees averaged (respectively) periods of 5.3, 3.6, 2.2 and 1.0 minutes
spent in the nest between foraging trips, so it appears that the faster

July 1972]

SCHLISING BEHAVIOR OF DIADASIA

183

Table 3. Times of nest provisioning activities by 4 females of
Diadasia bituberculata on 5 June, from 6:40-8:40 a.m.

Bee #1

Bee #2

Bee #3

Bee #4

Return

Leaving

Return

Leaving

Return

Leaving

Return

Leaving

6:43

6:48

6:44

6:46

6:57

6:59

7:25

7:26

6:57

? a

6:59

7:00

7:14

7:16

8:04 b

8:05

7:02

7:05

7:14

7:16

7:35

7:37

8:15

8:16

7:15

7:17

7:23

7:25

7:48

7:51

8:31

8:32

7:19

7:30

7:31

7:33

8:05

8:07

7:35

7:37

7:41

7:47

8:25

8:27

7:48

7:51

7:58

8:08

8:35

8:02

8:11

8:16

8:19

8:28

a Flight from nest not seen.

b Longest (and highly atypical) foraging flight recorded during early morning.

foragers may spend longer periods in the nest. Foraging time periods
later in the day became less uniform, for all individuals timed, as
pollen became scarcer, and it commonly took 2 to 4 times longer
for a forager to return with a pollen load. In-burrow periods also
became longer and fluctuated more among individuals and within the
population as a whole later in the day, until all bees ceased foraging
and began new cell construction in the nests.

Later in the season, at least, foraging and cell provisioning are
not alternated with new cell contruction while Calystegia pollen is
still plentiful. On days when Calystegia pollen was being used up rather
early, some bees were still collecting at 9:30-10:00 a.m., but others
had turned to further cell excavation in their burrrows — not to return
to foraging and provisioning of these cells until the next morning.
Females of D. consociata Timberlake, on the other hand, may collect
pollen throughout the day from Sida hederacea (Douglas) Torrey,
and will make three or four pollen trips in succession (Linsley et al.,
1952a), but apparently do not restrict their cell excavation activities
only to times after pollen foraging is completed for the day.

During nest construction a female may alternate cell excavation
with turret construction or repair. A few seconds after entering her
burrow for excavation, a bee backs out, kicking pebbles of soil removed
from below. Although the ground is very hard and dry where the
nests are dug, these pebbles are moist; some are used to fashion
the turret and the rest accumulate as a tumulus. Frequent flights
away from the nest during construction serve in obtaining nectar as

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a source of energy and also in obtaining moisture used in wetting
the hard soil. Bees in the genera Ptilothrix Smith ( —Emiphor Patton)
and Melitoma Lepeletier and Serville, in the same tribe as Diadasia,
are among the bees known to collect water used in moistening soil
in nest construction (Grossbeck, 1911; Rau, 1930; Linsley et al.,
1952a, 1956; Stephen et al., 1969). No species of Diadasia has been
recorded as collecting water for nest construction, according to Linsley
and MacSwain (1957), but these authors suggested the possibility of
nectar being used in soil softening. Roy Snelling, of the Natural
History Museum of Los Angeles County, Los Angeles, California, has
observed females of D. bituberculata collecting nectar in Penstemon
flowers in the San Gabriel Mountains of Los Angeles County, and feels
that such nectar may the the liquid used to soften soil in nest
construction (personal communication) . The very short periods ex-
cavating bees spent away from their nests at Three Rivers suggest a
source of liquid close by. Close freewater sources were examined
during excavation periods, but no Diadasia individuals were seen
taking water. Females were collected nearby taking nectar from the
four species already listed; the use of nectar from these or other
flowers in nest construction is very probable but has yet to be
demonstrated with certainty here.

While observations were not made on below ground aspects of
the nests, information is available for several species of Diadasia ,
including D. bituberculata, on various aspects of burrow and cell
structure, nest contents, parasites, larval overwintering, and adult emer-
gences (Linsley and MacSwain, 1952, 1957; Linsley et al., 1952a,
1952b). Although there is no published record on number of cells
per nest for this species, Linsley and MacSwain (1957) listed anywhere
from 1 to 42 cells per nest for other species of Diadasia, and noted
that nesting habits for D. bituberculata are similar to those of D.
consociata, which produces 1 to 15 (commonly 5 to 10) cells per
burrow. Of the 10 females observed provisioning and then excavating
on 5 June here, only one was still working at the same nest 6 days
later. It is not known how long a female takes to completely provision
one nest, but Linsley et al. (1952a) have recorded females of D.
consociata taking as long as 5 or 6 weeks to construct and provision
a burrow with 10 or more cells. They also noted that females of D.
consociata may construct more than one burrow; this may be true
of D. bituberculata as well, for between 25 May and 12 June at
Three Rivers new nests were continually being initiated. After nests
were provisioned, some females completely removed their turrets and

July 1972]

SCHLISING BEHAVIOR OF DIADASIA

185

filled the burrows to the surface with pieces of turret and soil from
the tumulus.

Male Behavior and Mating

At Three Rivers males of D. bituberculata are found from about
6:30 a.m. to 3:30 p.m. or later visiting flowers of C. fulcrata during
a characteristic cruising flight over the plants. They follow a low,
zig-zag flight 10 to 30 cm above the flowers, as if searching for
females, and enter only 5% of the open flowers. A male may fly
directly into a flower, and while rotating his body in the corolla,
take nectar from three to five of the nectaries. Nectar visits last
from one or two (usually four or five) to 25 seconds, and while the
tongue is inserted into the nectar, the abdomen and hind legs extend
into the area of anthers and stigma (Fig. 7). Males often rest for
up to 2.5 minutes, with legs on one side of the body on the anthers
and stigma and legs of the other side on petals, or with the body
curled around the stamens at the level of the anthers or lower. In
either position, a male may sit quietly, or — often with considerable
squirming — groom head and thorax with the front legs. These males
usually have abundant pollen on abdomen and legs; they frequently
brush the stigma when entering or leaving, or while in a flower, and
undoubtedly promote pollination.

While most commonly observed cruising Calystegia flowers, males
can also be found flying over the nest site. At Three Rivers males
were also found in flowers of the three species of plants utilized for
nectar by females. They were common in the deep tubular flowers of
Penstemon laetus and in the cup-shaped flowers of Clarkia speciosa
ssp. polyantha , and were rare in the cup-shaped flowers of Calochortus
superbus. These flowers are utilized by males as nectar sources and
as resting places, mostly, but not exclusively, later in the day when
Calystegia flowers are partially or totally closed. Penstemon flowers
as well as closed Calystegia flowers had sluggishly active males in
them late in the afternoon, but numerous flowers pinched at night
contained no sleeping males, and it is not known where males spend
the night.

Mating of D. bituberculata bees was reported by Linsley et al.
(1952a) as occurring in flowers of Convolvulus ; at Three Rivers
mating may occur in flowers of three species. Bee encounters thought
to be matings were observed once only in flowers of both Clarkia and
Calochortus. Mating was more clearly seen on two occasions (13
May at 10:00 a.m. and 5 June at 6:40 a.m.), commencing in flowers

186

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

of Calystegia fulcrata. In each case a cruising male pounced on a
pollen-collecting female, and with a loud buzzing fell from the flower
to the ground with her. In the first encounter the bees remained
in contact on the ground for eight or ten seconds before separating
and flying off. The other encounter lasted about five seconds; the
male then flew off, and the female flew to another flower and continued
pollen collecting. The very few matings seen during the 67 hours
of observations suggest that mating may mainly occur at the begin-
ning of the foraging season here (which probably began one week
or more before observations started on 12 May) . Observations
here do show agreement with the suggestion of Linsley et al. (1952a),
that males persist throughout the nesting season.

Dependence of Diadasia bituberculata on Calystegia

Linsley and MacSwain (1958) and Stephen et al. (1969) emphasized
that the pollen food-plant restriction called oligolecty must be con-
sidered a relative phenomenon, with some species more highly oligo-
lectic than others. Some bees that are typically highly oligolectic
may turn to other plants in times of shortage of their “preferred”
pollen sources. For example, Diadasia australis calif ornica Timberlake,
a desert cactus oligolege, has been found collecting pollen from nearby
Phacelia flowers when the local cactus pollen supply was exhausted
(Linsley and MacSwain, 1957, 1958). All females of D. bituberculata
examined at Three Rivers had only C. fulcrata pollen on their scopae.
Yet, the very beginning or end of the foraging season was not studied
here and it may be interesting to see if these bees are less faithful
in pollen collecting early in the season or during the final days of
foraging when the preferred pollen sources are very scarce. Through-
out its entire geographic range however, Linsley and MacSwain (1957)
believe that none of the oligolectic species of Diadasia is restricted
to a single species of a plant genus. They encountered D. bituberculata
in several localities, but always taking pollen only from the introduced
and weedy Convolvulus arvensis Linnaeus; they suspected that this
species also visited native and less widely distributed species of plants
in this group. This bee is able to utilize at least four plant species,
for during the spring of 1971 pollen-taking females of D. bituberculata
were collected on three native species of Calystegia in different
localities in California: C. fulcrata, in Tulare, Kern and Los Angeles
Counties; C. longipes (Watson) Brummitt, in Tulare and Kern
Counties; and C. purpurata (Greene) Brummitt, in Ventura County.

Stephen et al. (1969) have noted that some bees that are highly

July 1972]

SCHLISING BEHAVIOR OF DIADASIA

187

oligolectic in collecting pollen also take nectar from a very limited
number of species, and Linsley (1958) noted that in some cases
male flower response may be as specific as that of the female. At
Three Rivers in 1971 both sexes of D. bituberculata were found taking
nectar from no more than the four species listed earlier ( Calystegia ,
Penstemon , Clarkia and Calochortus) , although flowers of many species
were searched for the bees. The perennial Calystegia fulcrata , the
preferred pollen source, probably also serves as the preferred nectar
source each year; the same plants of Penstemon and Calochortus , also
perennials, may be utilized each season as well. Compared with nectar
sources listed for some other species of Diadasia (Linsley and Mac-
Swain, 1957; Linsley et al., 1952a), the nectar plants known for
D. bituberculata at Three Rivers are a restricted set.

Since these bees restrict their foraging to few plants it was of
interest to record the possible competitors they have, and the total
range of C. fulcrata flower visitors was sampled at Three Rivers.
Other than a very occasional ant or clerid beetle, bees were the only
flower visitors. Only a few instances of pollen collection were noted
during the study period, by females of Halictus farinosus Smith and
Lasioglossum punctatoventre (Crawford), so females of D. bituberculata
had essentially no competition for pollen here. (Linsley et al., 1952a,
similarly noted that the oligolectic D. consociata had no competition
for pollen on Sida hederacea .) Calystegia nectar was taken by
Emphoropsis rugosissima Cockerell, Osmia gabrielis Cockerell, 0.
nemoris Sandhouse, 0. subaustralis Cockerell, and Synhalonia stretchii
(Cresson), but it seems very unlikely that these nectar collectors were
ever common enough to diminish the supply of nectar needed by
Diadasia bees. Thus, in 1971 this population of D. bituberculata had
very little competition for food — in spite of their flower visits being
restricted to only one plant species for pollen and to only four
species for nectar.

Acknowledgments

Appreciation is expressed to P. H. Timberlake, University of Cali-
fornia, Riverside, and to R. R. Snelling, Natural History Museum
of Los Angeles County, Los Angeles, for identifying or verifying
identifications of bees. Thanks are also extended to C. 0. Balogh
and L. A. Bellue for help in collecting field data.

Literature Cited

Grossbeck, J. A. 1911. A contribution toward the life history of Emphor
bombiformis Cress. J. N. Y. Entomol. Soc., 19: 238-244.

188

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Linsley, E. G. 1958. The ecology of solitary bees. Hilgardia, 27: 543-599.

Linsley, E. G., and J. W. MacSwain. 1952. Notes on some effects of parasitism
upon a small population of Diadasia bituberculata (Cresson) (Hyme-
noptera: Anthophoridae). Pan-Pac. Entomol., 28: 131-135.

1957. The nesting habits, flower relationships, and parasites of some North
American species of Diadasia (Hymenoptera: Anthophoridae). Was-
mann J. Biol., 15: 199-235.

1958. The significance of floral constancy among bees of the genus Diadasia

(Hymenoptera, Anthophoridae). Evolution, 12: 219-223.

Linsley, E. G., J. W. MacSwain, and R. F. Smith. 1952a. The bionomics of
Diadasia consociata Timberlake and some biological relationships of
emphorine and anthophorine bees (Hymenoptera, Anthophoridae).
Univ. Calif. Publ. Entomol., 9: 267-290.

1952b. The life history and development of Rhipiphorus smithi with notes
on their phylogenetic significance (Coleoptera, Rhipiphoridae) . Univ.
Calif. Publ. Entomol., 9: 291-314.

1956. Biological observations on Ptilothrix sumichrasti (Cresson) and some
related groups of emphorine bees (Hymenoptera, Anthophoridae) .
S. Calif. Acad. Sci. Bull., 55: 83-101.

Michener, C. D. 1951. in Muesebeck, C. F. W., K. V. Krombein, H. K.

Townes, et al. Hymenoptera of America north of Mexico — synoptic
catalog. U. S. Dep. Agr. Monogr., No. 2. 1420 pp.

Munz, P. A. 1968. Supplement to a California flora. Univ. Calif. Press, 224 pp.
Munz, P. A., and D. D. Keck. 1959. A California flora. Univ. Calif. Press,

1681 pp.

Percival, M. 1965. Floral biology. Pergamon Press, London, 243 pp.

Rau, P. 1930. The nesting habits of Emphor bombiformis Cresson. Bull.
Brooklyn Entomol. Soc., 25: 28-35.

Schlising, R. A. 1970. Sequence and timing of bee foraging in flowers of
Ipomoea and Aniseia (Convolvulaceae) . Ecology, 51: 1061-1067.
Stephen, W. P., G. E. Boitart, and P. F. Torchio. 1969. The biology and
external morphology of bees. Agr. Exp. Sta., Oreg. State Univ., 140 pp.
U. S. Dep. Commer. 1971. Climatological data — California, 75: 145-220.

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july 1972]

SHIELDS BUTTERFLIES AND FLOWERS

139

Flower Visitation Records for Butterflies

(Lepidoptera)

Oakley Shields

Department of Entomology, University of California, Davis, California 95616

Interrelationships of plants and insects are of great evolutionary
importance. Recent evidence suggests that pollinators and flowers
have largely coevolved through time. This paper presents some flower
visitation records for butterflies and briefly discusses the interaction
of adult butterflies and flower food sources.

Butterflies are an extremely well-known group of organisms. The
sheer volume of literature on them staggers the imagination. To
my knowledge, no one has collected together the flower-feeding records.
No attempt is made here to undertake more than a modest overview,
being a survey of sources readily available to me. However, it is
apparent that strongly suggestive correlations emerge which are
worth considering.

Recently three sources dealing with “intrafloral ecology” have
reviewed much of the accumulated knowledge: Baker and Hurd
(1968), Stebbins (1970), and Faegri and Pijl (1971). Grant and
Grant (1965) give extensive records for insect pollinators of the
Polemoniaceae, including 45 butterfly species; these were not dupli-
cated in the records here. An important old paper that has apparently
escaped the attention of recent reviewers is that of Robertson (1895).
He says, “There are few evident butterfly-flowers. The best of them
are commonly visited by long-tongued bees and flies.” The field
is young and open to speculation.

Other work of note includes Knuth (1906), Robertson (1928),
Hingston (1930), Carpenter (1946), Hamm (1948)* Clench (1955),
Dronamraju (1960), Dronamraju and Spurway (1960), and Emmel
( 1971 ) . This latter gives a detailed report on the symbiotic relation-
ship of a large hesperine skipper and a Maxillaria orchid; it possesses
a 43 mm proboscis flexed at several points to probe the coiled corolla
tube of the orchid!

Since my knowledge of botany is nil, little attempt is made to
interpret the data but rather it is presented in accessible form for
botanists. The most striking fact, it seems to me, is that in four
separate lines (Ranunculales, Theales, Violales, and Saxifragales) ,
series of rather closely related groups are visited, indicating perhaps

The Pan-Pacific Entomologist 48: 189-203. July 1972

190

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

base

Fic. 1. Plant relationships for butterfly visitations, based on the classification
of Takhtajan (1969: fig. 31) (number of plant families used given for each
order ) \

that those groups are interrelated, as the massive botanic evidence
strongly suggests (Fig. 1). One would not expect such close similarities
if the flower-preference habit developed at random. But 29 of the
possible 94 orders are fairly linearly related (in four groups) and
only 10 orders more “distantly” related are used. The most primitive
dicot orders and the line from Magnoliales through Hamamelidales
appear not to be used for the most part, and the monocots are only
spottily visited (except Liliales). The flowers chosen by the widest
variety of butterfly subfamilies (five or more) include Apocynaceae,
Asclepiadaceae, Boraginaceae, Compositae, Cruciferae, Hippocas-
tanaceae, Hydrophyllaceae, Labiatae, Leguminosae, Polygonaceae,
Rubiaceae, and Verbenaceae. The commonest family used is Com-
positae, with 29 genera. The highest similarity in plant orders utilized
by two butterfly subfamilies (44%) were the Lycaeninae-Nymphalinae
and Hesperiinae-Pyrginae lines. More definitive conclusions must
await further data. It has been my experience that butterflies do
indeed have “preferred” flowers in a given habitat and usually fly
past many flower species.

1 Those families reported in Robertson (1928; records therein are not listed here) that are
in addition to those already recorded are Acanthaceae, Acearaceae, Alismaceae, Balsaminaceae,
Campanulaceae, Caryophyllaceae, Convolvulaceae, Dipsaceae, Ebenaceae, Fumariaceae, Gentia-
naceae, Lobeliaceae, Lythraceae, Oxalidaceae, Polemoniaceae, Pontederiaceae. Primulaceae,
Santalaceae, Staphyleaceae. and Valerianaceae.

July 1972]

SHIELDS BUTTERFLIES AND FLOWERS

191

An intriguing possibility exists here. The butterfly and flower
may be evolving within the same habitat, so that a particular flower
array may accompany a constellation of butterfly forms through time.
Butterflies appear for the most part to be promiscuous in their feeding
habits, visiting a variety of the “preferred” types.

According to Faegri and Pijl (1971), butterflies alight on a blossom
margin to feed. They characterize the preferred butterfly flowers
(psychophily) as being diurnal; no closing at night; weak, pleasant
odor; vividly colored; blossom rim not very dissected; erect, radial
blossom with flat, often narrow rim; ample nectar hidden in narrow
tubes or spurs; and nectar or tongue guide present.

Butterflies often walk over a cluster of blossoms while feeding;
thus, pollen could be carried by their legs (e.g., the pollinia of
Asclepias are pulled out by the monarch butterfly’s legs) and bodies.
Many butterflies have hair trailing along the inner surface of the
hind wing upperside, on the upper and undersurface of the thorax,
and the face, which may be important in picking up pollen, as well
as the proboscis itself, leg setae, and leg hooks. Hingston (1930)
reports an instance where pollen from a Gloriosa lily is shaken on the
visiting Papilio by the action of its wingbeat when feeding. However,
the rather casual visitation of flowers by some species may not effect
pollen transfer (Percival, 1965: 187). It remains a problem to dis-
criminate between pollen transfer and casual visitation.

Flower feeding by butterflies is often in company with Hymenoptera
(especially), Coleoptera, Diptera, and others, e.g. on Chrysothamnus ,
Cirsium , Eriodictyon , Eriogonum , Melilotus , Monardella , Prunus, Rhus ,
and Solidago. In the case of Mimulus , the pollinators are humming
birds, bumble bees, sphingids, and Papilio butterflies. In a few
instances, butterfly adults feed on the same plant species as their
larvae do ( Colias eurytheme on Medicago sativa, Danaus plexippus on
Asclepias, and Apodemia mormo and Philotes rita, hattoides , and
enoptes on Eriogonum ), but more usually the butterfly adult feeds
on plants other than its larval foodplant (Faegri and Pijl, 1971: 133).
Often the larval foodplant is not in bloom when the butterfly adult is
on wing; or if it is, its flowers are usually not attractive to that
species.

Many of the flower records given, as far as I can tell, constitute
regular rather than casual visitation, but no attempt has been made to
distinguish these from rare visits. I once noticed a worn male
Euphydryas chalcedona avidly feeding on Sambucus mexicana blooms,
16 June 1971, Thompson Canyon, just NE of Monticello Dam, Yolo

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Co., Calif., on a hot, humid day. Sambucus is almost never visited by
butterflies, in my experience, but there were few other flowers in
bloom on this date.

It may be of evolutionary significance that butterflies of the sub-
family Megathyminae (larvae in Agavaceae) , generally considered
one of the most primitive groups of skippers, do not flower feed but
do take moisture along creeks, damp sand, etc. Certain butterfly
species with fresh (usually) to worn winged males and worn females
will visit moisture. Other non-flower sources for butterflies include
carrion, honey dew, rotting fruit, tree sap from wounds, dung, and
urine. Thus, perhaps it would have been possible for butterflies to
survive at a time when no flowers were available for food, i.e. before
the advent of angiosperm flowers (e.g. some lycaenids in Africa feed
on lichens as larvae) .

Nocturnal feeding at flowers by moths must be extensive since
there are about ten times as many moths as butterflies. Though
certainly of great interest to evolutionists, this will be difficult data
to acquire.

The butterfly Eumaeus atala in Florida feeds as an adult on Serenoa
(Palmae) and Bidens (Compositae) , while its larva eats Zamea
integrifolia (Cycadaceae) (Klots, 1951). Serenoa being a small
palm, and the relationship of Lycaeninae to the other butterflies at
present being obscure and ancient, one wonders about a possible
early Mesozoic association between Serenoa , Zamea , and Eumaeus.

I wish to thank Grady Webster for originally suggesting this topic
to me. Robbin W. Thorp reviewed the ms. and assisted in the literature
search. The following people furnished some records by correspon-
dence: C. Durden, D. Eff, L. P. Grey, R. Heitzman, C. Henne,
W. H. Howe, P. S. Remington, F. T. Thorne, and J. W. Tilden.
This work was supported in part by an N.S.F. graduate trainee-
ship.

Butterfly Records For Flower Visitation
AGAVACEAE

Nolina microcarpa : Erora quaderna 45 2
AMARYLLIDACEAE
Allium douglasi : Mitoura spinetorum 54

Brodiaea sp.: Battus philenor hirsuta, Erynnis tristis, Papilio zelicaon (all 69)
Brodiaea pulchra: Battus philenor hirsuta 69
ANACARDIACEAE
Rhus glabra: Chlosyne nycteis 25

2 Numbers refer to literature cited at the end of this article.

July 1972]

SHIELDS — BUTTERFLIES AND FLOWERS

193

Rhus trilobata : Callophrys apama homoperplexa 71, C. comstocki, Incisalia
iroides, Mitoura johnsoni (all 69) , M. spinetorum 54
APOCYNACEAE

Apocynum sp.: Apodemia nais 3, Chlosyne leanira ssp., C. palla, Emesis zela
(all 69) , Epargyreus darns 62, Euphydryas anicia 3, Hesperia harpalus
oregonia 53, Limenitis lorquini 69, Lycaena editha 69, Phyciodes campestris
53, Piruna pirus 3, Satyrium liparops 62, S. sylvinus 69, Speyeria egleis ssp.
69, S. hydaspe purpurascens 53, S. zerene conchyliatus 53
Apocynum androsaemifolium: Speyeria callippe juba 71

Apocynum cannabinum : Chlosyne nycteis 25, Lephelisca borealis 57, Limenitis
archippus 71, Satyrium alcestis 41, S. falacer 7 & 8, S. liparops strigosum 8,
S. Ontario 8

Trachelospermum sp.: Atlides halesus 29, Papilio polydamas 29
ARALIACEAE

Aralia hispida : Colias interior 29
ASCLEPIADACEAE

Asclepias incarnata : Atrytone conspicua 29, Chlosyne nycteis 25, Danaus
plexippus 6, Poanes massasoit 29, Speyeria cybele 6, S. idalia 6
Asclepias syriaca : Danaus plexippus 6, Epargyreus clarus 8, Satyrium acadica
52, S. falacer 7, S. liparops 62, S. titus mopsus 6, Speyeria cybele 6 & 8
Asclepias tuberosa: Chlosyne ismeria 41, C. nycteis 25, Colias philodice,
Danaus plexippus, Everes comyntas (all 6), Nathalis iole 41, Phyciodes
tharos 6, Satyrium falacer, S. Ontario autolycus, S. titus mopsus (all 41),
Speyeria cybele 8, Vanessa cardui 41
BERBERIDACEAE

Berberis sp.: Incisalia eryphon 71, Mitoura spinetorum 54
BORAGINACEAE

Echium vulgar e: Achalarus lyciades, Thorybes bathyllus, T. pylades (all 8)
Heliotropium sp.: Cercyonis silvestris 69, Lycorea ceres atergatis (to dried
plant) 34, Pseudocopaeodes eunus 69

Heliotropium indicum (dried plants) : Danaus plexippus, Hymenitis andromica,
Hypothyris euclea, Ithomia drymo, Lycorea ceres, Mechanitis isthmia,
Melinaea lilis, Tithorea harmonia (all 21)

Tournefortia argentia (dead twigs) : Danaus melissa 21, Euploea ssp. 21
GAPPARIDACEAE

Cleome serrulata: Papilio bairdii brucei 69
CAPRIFOLIACEAE

Sambucus coerulea : Mitoura spinetorum 54
Triosteum perfoliatum : Speyeria cybele 6
COMPOSITAE

Achillea sp.: Euphydryas editha baroni, Lycaena editha, Mitoura nelsoni (all
69) , M. spinetorum 54, Satyrium calif ornica 53, S. dryope 69, Speyeria
atlantis nausicaa 69

Achillea millefolium: Lycaena nivalis 35, Satyrium falacer 7
Ageratum sp.: Papilio polydorus queenslandicus 2
Anaphalis sp. : Mitoura spinetorum 54

Anaphalis margaritacea: Callophrys apama homoperplexa, Mitoura siva, M.
spinetorum (all 71)

Antennaria sp.: Incisalia augustinus 6, /. niphon 6

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Antennaria parvifolia : Mitoura spinetorum 54
Arnica alpina : Colias nastes 66
Arnica cordifolia: Lycaena rubidus 9

Aster sp. : Colias croceus, Gonepteryx rhamni, Lycaena phlaeas, Maniola
jurtina, Nymphalis io, N. urticae, Pieris brassicae, P. rapae, Polygonia
c-album, Polyommatus icarus , Vanessa atalanta, V. cardui (all 24)

Aster canescens : Lycaena mariposa 9
Baccharis sp.: Libytheana bachmanii 70
Baccharis sarothroides : Hylephila phylaeus 69

Bidens sp. : Atlides halesus, Eumaeus atala, Hemiargus thomasi, Strymon
maesites (all 29)

Carduus calif ornicus : Speyeria nokomis apacheana 9
Chrysanthemum leucanthemum var. pinnatifidum : Satyrium falacer 7
Chrysothamnus sp. : Hesperia harpalus 69, H. h. leussleri 3, H. juba 69,
Libytheana bachmanii 71, Satyrium acadica coolinensis 50, Speyeria callippe
ssp. 71

Chrysothamnus nauseosus: Speyeria egleis tehachapina 15

Cirsium sp. : Hesperia Columbia, H. harpalus dodgei, H. lindseyi (all 33),
Satyrium falacer 7

Cirsium arvense: Danaus plexippus 22
Cirsium vulgare: Danaus plexippus 22
Coreopsis sp.: Zerene cesonia 58

Coreopsis lanceolata : Chlosyne gorgone 32, Lephelisca virginiensis 32
Echinacea pallida : Chlosyne nycteis 25

Erigeron sp. : Chlosyne damoetas malcolmi 69, Mitoura spinetorum 54

Erigeron philadelphicus : Boloria selene 36

Erigeron ?strigosus: Satyrium falacer 7

Eriophyllum lanatum: Lycaena nivalis 35

Eupatorium sp. : Limenitis archippus 71

Eupatorium coelestinum : Ancyloxypha numitor, Hylephila phylaeus, Precis
lavinia coenia (all 6)

Eupatorium purpureum : Danaus plexippus 6, Hesperia leonardus 6
Gaillardia sp.: Mitoura spinetorum 71
Gnaphalium sp.: Incisalia niphon 6, Polygonia gracilis 29
Grindelia sp., probably: Ochlodes yuma 71

Haplopappus sp. : Mitoura siva 71, M. spinetorum 54, Speyeria callippe 69
Haplopappus bloomeri : Hesperia harpalus yosemite 65
Haplopappus linearifolius : Mitoura loki 69

Helenium sp.: Mitoura spinetorum 54, Speyeria atlantis nikias 71

Helenium autumnale: Euptoieta claudia 40, Phyciodes tharos 40

Helianthus annuus : Satyrium calanus 32

Pluchea camphorata: Chlosyne nycteis 25

Rudbeckia sp.: Atrytone arogos iowa 3

Rudbeckia laciniata: Speyeria zerene platina 3

Senecio sp.: Parnassius clodius baldur 20, Speyeria callippe ssp. 3 & 71
Senecio douglasii monoensis : Mitoura spinetorum 54
Senecio longilobus: Sandia macfarlandi 28

Solidago sp.: Cercyonis boopis, Danaus plexippus, Mitoura nelsoni, Neophasia

July 1972]

SHIELDS — BUTTERFLIES AND FLOWERS

195

menapia, Pholisora libya, Satyrium dryope (all 69), S. saepium 3, Speyeria
nokomis apacheana 69
Solidago canadensis : Danaus plexippus 6

Solidago gigantea, probably: Callipsyche behrii 12, Mitoura nelsoni 12, M.

spinetorum 12 & 54, Satyrium calif ornica 12
Vernonia sp.: Autochton cellus 29

Wyethia , or Balsamorhiza sp.: Parnassius clodius sol 69, Speyeria callippe juba
69, S. c. nevadensis 20, S. coronis ssp., S. cybele leto, S. hydaspe, S. zerene
(all 53 & 69)

Zinnia sp. : Chlosyne nycteis 25
CORNACEAE

Cornus canadensis : Mitoura johnsoni 54
CRASSULACEAE
Dudleya cymosa : Papilio indra 69
Sedum sp.: Speyeria callippe ssp. 3
CRUCIFERAE

Arabis blepharophylla: Incisalia fotis bayensis 69
Barbarea vulgaris : Strymon melinus 6

Brassica sp.: Anthocaris cethura, Colias eurytheme, Erynnis tristis (all 69),
Incisalia niphon 6, Leptotes marina 69, Speyeria callippe comstocki 71,
Strymon melinus 69, Vanessa carye 69
Brassica nigra: Pieris rapae 69
Cardamine bulbosa: Incisalia niphon 47
Erysimum sp.: Speyeria callippe macaria 51

Erysimum asperum: Parnassius clodius baldur 19, Papilio zelicaon 20
Rorippa sp. : Emesis zela 69, Mitoura johnsoni 54

Sisymbrium loeseli : Lycaena rubidus, Polygonia zephyrus, Speyeria zerene
cynna (all 69)

ERICACEAE

Arctostaphylos sp.: Mitoura spinetorum 54
Arctostaphylos nevadensis: Mitoura johnsoni 54
Gaylussacia sp.: Incisalia henrici 29
Rhododendron sp.: Tros philoxenus 68

Vaccinium sp.: Graphium marcellus 6, Incisalia henrici 29, I. niphon 6,
Thorybes pylades 6
EUPHORBIACEAE

Croton sp.: Libytheana bachmanii 71
Croton linearis : Lerodea eufala 48
Poinsettia sp.: Pratapa cleobis 68
FAGACEAE

Castanopsis sp.: Satyrium kingi 11
GERANIACEAE

Erodium cicutarium: Vanessa cardui 30
GRAMINEAE

Sorghum sp. : Libytheana bachmanii larvata 70

HIPPOCASTANACEAE

Aesculus sp.: Aporia leucodyce , Delias belladonna, Tros philoxenus, Zetides
cloanthus, Z. sarpedon (all 68)

Aesculus calif ornica: Chlosyne leanira 69, Epargyreus clarus 20, Euphydryas

196

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

chalcedona 20, Incisalia iroides, Mitoura spinetorum, Ochlodes agricola (all
69), Satyrium adenostomatis 53, S. auretorum 20 & 53, S. saepium 20 & 53,
Speyeria callippe inornata 69, S. egleis atossa 9
HYDROPHYLLACEAE

Eriodictyon sp. : Atlides halesus, Celastrina argiolus echo, Cercyonis silvestris
paulus, Euphydryas editha baroni, E. chalcedona, Heliopetes ericetorum,
Hemiargus isola, Ministrymon leda, Mitoura johnsoni, M. nelsoni (all 69),
M. spinetorum 54, Ochlodes agricola, Papilio eurymedon, P. indra pergamus,
Satyrium auretorum, S. calif ornica (all 69), Speyeria callippe macaria 51
Eriodictyon angustifolium : Mitoura spinetorum 54

Eriodictyon calif ornicum: Hesperia Columbia 33, Satyrium calif ornica 20,
Speyeria callippe macaria 71
IRIDACEAE

Iris sp. : Carterocephalus palaemon 64, Mitoura spinetorum 54, Papilio rutulus
arizonensis 71, Parnassius clodius 69
Iris missouriensis : Coenonympha tullia mono 69, Papilio rutulus 69
LABIATAE

Marrubium vulgar e: Erynnis funeralis 9, Speyeria callippe macaria 51 & 71
Monarda sp.: Erora quaderna 45, Speyeria aphrodite ethne 3 & 69, S. coronis,
S. c. halcyone, S. cybele leto, S. hydaspe, S. zerene garretti (all 69)
Monarda menthaefolia : Ochlodes snowi 3

Monardella sp.: Cercyonis silvestris 69, Chlosyne hoffmanni 20, Oeneis
nevadensis 69, Speyeria atlantis dodgei 69, S. callippe inornata 53, S. c.
juba 69, S. hydaspe 20, S. zerene malcolmi 20
Monardella odoratissima: Chlosyne hoffmanni 19, Papilio zelicaon 17, Parnas-
sius clodius baldur 19, Plebejus shasta, Polygonia zephyrus, Speyeria egleis
(all 17)

LAURACEAE

Umbellularia calif ornica: Lycaena arota 20, Incisalia iroides 69, Mitoura
johnsoni 69
LEGUMINOSAE
Astragalus sp. : Yvretta rhesus 3
Bauhinia sp. : Trogonoptera brookiana albescens 10
Cercis sp. : Incisalia henrici 29
Cercis canadensis: Incisalia henrici 6
Cercis occidentalis : Incisalia iroides 69
Crotalaria sp.: J amides celeno aelianus 10

Lupinus sp. : Incisalia eryphon 3, I. niphon 6 & 29, Mitoura nelsoni 69, M.
spinetorum 54

Medicago sativa : Cercyonis silvestris paulus 9 & 69, Chlosyne minuta 71, C.
nycteis 71, Colias eurytheme, Heliopetes ericetorum, Hemiargus gyas,
Hesperia harpalus, Leptotes marina, Pholisora catullus (all 69), Phyciodes
tharos 71, Polites themistocles 69, Speyeria edwardsii 71, Zerene eurydice 9
Melilotus sp.: Cercyonis silvestris paulus 9

Melilotus albus: Chlosyne nycteis 25, Everes comyntas 6, Satyrium falacer 7,
S. liparops 62

Melilotus officinalis : Satyrium falacer 7

Trifolium pratense : Colias eurytheme, Danaus plexippus, Hylephila phylaeus
(all 6) , Satyrium falacer 7

july 1972]

SHIELDS BUTTERFLIES AND FLOWERS

197

LILTACEAE

Camassia scilloides : Hesperia metea 26
Gloriosa superba : Papilio demoleus 27, P. polytes 27
LOGANIACEAE

Buddleia sp. : Badamia exclamationis 68, Chlosyne nycteis 25, Delias belladonna
68, Dilipa morgiana 68, Limenitis archippus 71, Papilio polyctor, Zetides
cloanthus, Z. sarpedon (all 68)

MALVACEAE

Hibiscus sp.: Catopsilia crocale, C. florella gnoma , C. pomona, C. pyranthe
minna (all 56)

Sida hederacea: Strymon columella 71
MYRTACEAE

Psidium guajava: Papilio aristodemus ponceanus 49
ONAGRACEAE

Epilobium sp. : Mitoura spinetorum 71
ORCHIDACEAE

Bonatea darwinii : Pyrgus elmo 61
Maxillaria ontoglossom : Perichares philetes dolor es 16
Orchis pyramidalis : Syricthus alveolus 61
Platanthera hookeri: Nisoniades sp. 61
Pogonia ? ophioglossides : Poanes hobomok 37
PALMAE

Serenoa sp.: Eumaeus atala 29
POLYGONACEAE

Eriogonum sp. : Libytheana bachmanii 71, Speyeria egleis tehachapina 15
Eriogonum deserticola: Eurema nicippe 69, Microtia dymas imperialis 69
Eriogonum fasciculatum : Cercyonis silvestris 69, Hemiargus isola 69, Mitoura
spinetorum 54, Philotes battoides bernardino, Satyrium sylvinus, Speyeria
callippe corns tocki (all 69)

Eriogonum giganteum : Strymon avalona 9

Eriogonum umbellatum : Mitoura spinetorum 54, Satyrium saepium 3
Eriogonum wrightii: Apodemia mormo tuolumnensis 69
Polygonum sp. : Limenitis archippus 71, Mitoura spinetorum 54
PORTULACACEAE

Calyptridium sp.: Lycaena cupreus 69, Mitoura nelsoni 20, M. spinetorum 54,
Nymphalis milberti 69, Philotes enoptes 69, Speyeria callippe elaine 53, S. c.
juba 69, S. egleis 71

Calyptridium umbellatum: Mitoura johnsoni 54, Satyrium calif arnica 20
PYROLACEAE

Monotropa uniflora: Epargyreus clarus 6
RANUNCULACEAE
Clematis sp.: Tros philoxenus 68
Delphinium carolinianum: Hesperia metea 26
Ranunculus sp.: Hesperia Columbia 33
Ranunculus calif amicus : Incisalia fotis bayensis 69
RHAMNACEAE

Ceanothus sp.: Apodemia nais 3, Incisalia iroides 69, Mitoura nelsoni muiri
69

193

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 43 , NO. 3

Ceanothus americanus : Satyrium edwardsii 42, S. Ontario 29, S. titus 42, S. t.
mopsus 6

Ceanothus cordulatus : Mitoura johnsoni 38 & 54, M. spinetorum 38 & 54
Ceanothus fendleri : Erora quaderna 45, Mitoura spinetorum 54
Ceanothus ovatus: Mitoura spinetorum 54
ROSACEAE

Chamaebatia foliolosa: Mitoura nelsoni 53 & 69
Dryas octopetala : Boloria alberta 67, B. astarte 67
Fragaria sp.: Hesperia metea 29, Mitoura johnsoni 54
Fragaria virginiana var. illinoensis: Hesperia metea 26
Potentilla sp. : Hesperia metea 29, Mitoura nelsoni 69
Potentilla fruticosa : Lycaena rubidus sirius 3, Mitoura spinetorum 54
Prunus sp.: Callophrys dumetorum 71, Glaucopsyche lygdamus ssp. 69,
Incisalia eryphon 3, 7. niphon, Limenitis weidemeyerii, Speyeria aphrodite
ethne, S. coronis ssp., S. edwardsii (all 71)

Prunus americana: Incisalia niphon 47, Mitoura spinetorum 54
Prunus fasciculata: Callophrys comstocki 69, C. dumetorum 69, Mitoura
spinetorum 54

Prunus emarginata: Incisalia eryphon 69, Mitoura nelsoni 69
Prunus melanocarpa: Limenitis weidemeyerii 71, Mitoura spinetorum 54,
Speyeria coronis halcyone 71
Prunus virens: Erora quaderna 45
Rubus sp. : Libytheana bachmanii 70
Spiraea latifolia: Celastrina argiolus pseudargiolus 6

RUBIACEAE

Cephalanthus sp.: Atrytone conspicua 29, Autochton cellus 29
Cephalanthus occidentalism. Atalopedes campestris, Atrytone pontiac, Colias
eurytheme, Epargyreus clarus, Speyeria cybele (all 6)

Ixora sp.: Papilio demolion 10
Morinda roioc : Papilio aristodemus ponceanus 49
Mussaenda sp.: Trogonoptera brookiana albescens 10
RUTACEAE

Thamnosmci montana: Papilio indra fordi 69
SALICACEAE

Salix sp. : Callipsyche behrii, Cercyonis oetus, Chlosyne acastus, C. leanira
alma (all 69), Mitoura spinetorum 54, Satyrium calif ornica 69
SAXIFRAGACEAE
Hydrangea sp.: Autochton cellus 29

Philadelphus lewisii var. calif ornicus: Satyrium auretorum spadix 20
Ribes sp. : Mitoura spinetorum 71
SCROPHULARIACEAE
Castilleja sp. : Papilio indra pergamus 69
Mimulus sp.: Battus philenor hirsuta 69
Penstemon montanus : Papilio eurymedon albanus 3

Penstemon tolmiei ssp. formosus: Speyeria callippe semivirida 53 & 69, S.

coronis, S. egleis ssp., S. zerene ssp. (all 53)

Veronica sp.: Chlosyne nycteis 25
SOLANACEAE

Lycium andersonii: Papilio indra fordi 69

july 1972]

SHIELDS BUTTERFLIES AND FLOWERS

199

THYMELEACEAE

Primelea sp. : Anisynta tillycirdi, Exometoeca nycteris, Ogyris idmo (all 4),
Papilio macleayanus 2
UMBELLIFERAE

Cymopterus sp., probably: Mitoura spinetorum 54
Daucus carota : Satyrium falacer 7
Eryngium yuccaefolium: Phyciodes tharos 6

Lomatium dasycarpum ; Callophrys viridis, Incisalia fotis bayensis, Phyciodes
mylitta (all 69)

Lomatium utriculatum: Incisalia fotis bayensis 69
VERBENACEAE
Duranta sp.: Atella phalantha 68

Lantana sp.: Atella phalantha 68, Atrophaenura coon doubledayi 10, A. nox
erebus 10, Cethosia chrysippe cydippe 2, Cirrochroa regina sophene 2, Cynthia
erota 68, Euripus consimilis 68, Graphium agamemnon 10, Idea leuconoe
60, Ismene jaina 68, Jamides celeno aelianus 10, Lebadea martha malayana
10, Papilio liomedon 68, P. polydorus queenslandicus 2, Parathyma nefte
subrata, Parthenos sylvia lilacinus, Phalanta alcippe alcesta (all 10), Prioneris
sita, Tros aidoneus, T. hector , T. jophon, Troides helena (all 68), Vanessa
carye 9, Vindula arsinoe erotella 10, Zetides agamemnon 68, Z. sarpedon 68
Lantana camara: Baoris mathias, Catopsilia pyranthe, Danaus chrysippus,
Papilio demoleus, P. polytes, Precis almana (all 13)

Verbena sp.: Appias lyncida 68, Hesperia metea 26
VIOLACEAE

Viola sp. : Erynnis brizo 29
Viola pedata: Hesperia metea 26

Subfamily Relationships of the Butterfly Genera Mentioned
PAPILIONIDAE

Papilioninae: Atrophaenura, Battus, Graphium, Papilio, Trogonoptera, Troides,
Tros, Zetides

Parnassiinae: Parnassius
PIERIDAE

Pierinae: Anthocaris, Aporia, Appias, Delias, Neophasia, Pieris, Prioneris
Coliadinae: Catopsilia, Colias, Eurema, Gonepteryx, Nathalis, Zerene
LIBYTHEIDAE
Libytheana
NYMPHALIDAE

Satyrinae: Coenonympha, Maniola, Oeneis
Danainae: Danaus, Euploea, Idea, Lycorea

Ithomiinae: Hymenitis, Hypothyris, Ithomia, Mechanitis, Melinaea, Tithorea
Nymphalinae: Atella, Boloria, Cethosia, Chlosyne, Cirrochroa, Cynthia, Dilipa,
Euphydryas, Euptoieta, Euripus, Lebadea, Limenitis, Microtia, Nymphalis,
Phalanta, Parathyma, Parthenos, Phyciodes, Polygonia, Precis, Speyeria,
Vanessa, Vindula
LYCAENIDAE

Riodininae: Apodemia, Emesis, Lephelisca

Lycaeninae: Atlides, Callipsyche, Callophrys, Celastrina, Erora, Eumaeus,

200

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Everes, Glaucopsyche, Hemiargus, Incisalia, J amides, Leptotes, Lycaena,
Ministrymon, Mitoura, Ogyris, Philotes, Plebejus, Polyommatus, Pratapa,
Sandia, Satyrium, Strymon
HESPERIIDAE
Trapezitinae: Anisynta

Hesperiinae: Ancyloxipha, Atalopedes, Atrytone, Baoris, Carter ocephalus,

Hesperia, Hylephila, Lerodea, Nisoniades, Ochlodes, Perichares, Piruna,
Poanes, Polites, Pseudocopaeodes, Yvretta
Coeliadinae: Badamia, Ismene

Pyrginae: Achalarus, Autochton, Epargyreus, Erynnis, Exometoeca, Heliopetes,
Pholisora, Pyrgus, Syricthus, Thorybes

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known range, habits, variations, and history. J. Res. Lepid., 4:
233-250 [and my field notes].

55. Shields, 0. and J. C. Montgomery. 1966. The distribution and bionomics

of arctic-alpine Lycaena phlaeas subspecies in North America. J. Res.
Lepid., 5: 231-242.

56. Shull, E. M. 1952. Migration of Catopsilia butterflies in India. Lepid.

News, 6: 68-69.

57. Simmons, R. S. 1956. Notes on ten new butterfly records for the state of

Maryland. Lepid. News, 10: 157-159.

58. Simmons, R. S. and W. A. Andersen. 1961. Notes on five new butterfly

records for the state of Maryland. J. Lepid. Soc., 15: 99-101.

59. Stebbins, G. L. 1970. Adaptive radiation of reproductive characteristics

in angiosperms, I: Pollination mechanisms. Annu. Rev. Ecol. Syst.
1: 307-326.

july 1972]

SHIELDS BUTTERFLIES AND FLOWERS

203

60. Straatman, R. 1955. Notes on methods of collecting Indo-Australian

Lepidoptera. Lepid. News, 9: 74-76.

61. Swezey, 0. H. 1945. Insects associated with orchids. Proc. Hawaii Entomol.

Soc., 12: 343-403.

62. Syme, P. D. 1961. Observations on Strymon liparops (Lycaenidae) . J.

Lepid. Soc., 15: 108.

63. Takhtajan, A. 1969. Flowering plants: origin and dispersal. Smithsonian

Institution Press, Washington, D.C. 310 pp.

64. Tilden, J. W. 1957. Carterocephalus palaemon in California (Hesperiidae) .

Lepid. News, 11: 43.

65. 1959. The butterfly associations of Tioga Pass. Wasmann J. Biol.,

17: 249-271.

66. Wyatt, C. 1957a. Collecting on the Mackenzie and in the western Arctic.

Lepid. News, 11: 47-53.

67. 1957b. Observations on Boloria distincta (Nymphalidae) . Lepid. News,

11: 142-146.

68. Wynter-Blyth, M. A. 1957. Butterflies of the Indian region. The Bombay

Natural History Society, Bombay. 523 pp., 72 pis.

69. From my field notes and those of J. F. Emmel.

70. From my unpublished ms. on the Libytheidae.

71. Records from correspondence (see acknowledgments).

SCIENTIFIC NOTE

Notes on the Feeding Habits of Haemolaelaps glasgowi (Acarina:
Laelaptidae) . — Specimens of the mite Haemolaelaps glasgowi (Ewing), utilized
in this study, were recovered from the round-tailed ground squirrel, Spermophilus
tereticaudus neglectus Merriam, near Tucson, Arizona, or from its nests.
Feeding studies were conducted at a mean room temperature of 84° F and at
a mean relative humidity of 20%. A juvenile mouse, Mus musculus Lin., was
thoroughly examined, before placing a mite on it, to ascertain that it was
ectoparasite free. All mites placed on the mouse went to the rump. Starved
mites were deprived of blood or its constitutes for 10 days. Starved adult females
remained attached to the dermis from one to two hours after which they
appeared engorged. Most of these were on the mouse from 12 to 15 hours. All
adult females fed one to three days previously on moist raw beef did ilot
feed on the mouse. Most starved adult females were attracted to the beef.
Approximately half of these wandered over it for a minute, probed it with their
mouthparts, then fed. They would then go to a different location and repeat
this probing and feeding. The time spent on the beef was four to five minutes.
One starved adult male fed 20 to 30 seconds on it while one recently fed adult
male ignored it. All protonymphs and deutonymphs ignored the raw beef. —
James D. Lang, Department of Entomology, University of Arizona, Tucson,
85721.

204

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Larval Distribution of Paraclunio alaskensis at Point Pinos
Sewage Outfall, Monterey County, California

(Diptera: Chironomidae)

Douglas T. Cheeseman, Jr. and Paul Preissler

De Anza College, Cupertino, California 95014, and Stanford University,

Stanford, California 94305

The midge Paraclunio alaskensis Coquillett, an intertidal insect,
was observed in large numbers in the area of the Point Pinos sewage
outfall of Pacific Grove, California. A five week study during the
months of July and August 1971 was undertaken because P. alaskensis
represented an unusually abundant macroscopic organism close to a
point source of pollution. The Pacific Grove Primary Sewage Treat-
ment Plant releases into the outfall an average of 1.6 million gallons
per day of effluent containing chlorine to lower coliform counts.
Chlorine concentrations of 10 ppm are common near the outfall.

The larvae of P. alaskensis were chosen for study because they
provided a better index of population distribution than the adults
which were too mobile and completed their life functions within the
time of one low tide (Saunders, 1928).

The larvae in the area of the Pt. Pinos outfall fed on organic
detritus or “slime” found in greatest concentrations on the rocks
within the first 14 meters from the outfall. The slime consisted of
organic sewage, a diatom slick, occasional stunted specimens of the
alga Gigartina papillata Agardh, and other unidentified components.
Each P. alaskensis larva spins a silken tube which attaches it to the
rocky outcrops where the slime is abundant. During the two daily
high tides the tube provides protection from heavy wave action on the
vertical rocky surfaces. Saunders (1928) noted that preference is
shown for the vertical sides of large boulders and rocks which is
also the case at Pt. Pinos. When the larvae leave the tubes to feed
during low tides they are able to cling to their rocky habitat by means
of prothoracic and abdominal pseudopods.

Methods. — Data were collected on the distribution of the larvae
in the vicinity of the outfall and correlated with distance from the
outfall. Thirty-six samples were taken at random at Point Pinos
ranging from 0.0 tidal height to approximately 1.5 m above mean
low water. Six samples were taken at two control areas within
Monterey County: rocky outcrops near Asilomar Beach and Malpaso

The Pan-Pacific Entomologist 48: 204-207. July 1972

JULY 1972] CHEESEMAN & PREISSLER — PARACLUNIO LARVAE

205

800 -

750

700 -

20cm X 20cm

Mean Population Curve

10 15 20 25 30

35 40 45 50 55 60 65 70 75 80 85

Distance from Outfall in Meters

90

-i- — r r-

100 105 110

Fig. 1. Distribution of P. alaskensis larvae as a function of distance from
sewage outfall.

Creek tidepools. Each sample covered a quadrant 20 cm X 20 cm.

The 36 samples taken in the area of the outfall were divided into
three groups according to distance from the outfall: 0-14 m, 15-28
m and 29-160 m. Attention was given to the type of substrate and
the surrounding vegetation. Ten quadrants were selected from 0—14
m, the area of granitic substrate covered with organic slime. The
substrate of the second sample area included eight quadrants with
assorted algae, one quadrant from a Mytilus bed, and only one quadrant
found with organic slime. The quadrants in the third sample area
contained slime in one and rocks with a large variety of matted
algae and invertebrates in the remaining fifteen. The samples col-
lected in the control areas were from rocks densely covered with
macroscopic algae.

Large algae samples were put directly into plastic bags after they
were removed. The exposed rock surface was then sprayed with a
50% solution of sodium hypochlorite and all remaining organisms
collected.

A collecting tray was constructed to remove larvae from the slime
covered vertical rock surfaces surrounding the outfall. This consisted
of a sheet metal box measuring 40 cm X 10 cm X 5 cm from which

206

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 8

one of the long sides had been removed. Sponges were wrapped in a
plastic bag and compressed into the tray. A piece of cheesecloth was
then folded and placed over the outer, top edge of the bag covered
sponges. This was to provide a surface to which the larvae or any
falling particulate matter could cling. The apparatus was firmly
held against the lower edge of the 20 X 20 cm quadrants so that
the sponges were compressed and took the shape of the rock. The
entire quadrant was subsequently scraped and washed with a 50%
sodium hypochlorite solution. Both particulate matter and organisms
fell onto the cheesecloth. When the quadrant was completely bare
the sponges, cheesecloth, and bag were carefully removed from the
tray. The plastic bag was turned inside out enveloping the cheesecloth
and sample and freeing the sponges. The sample was then taken to
the laboratory for counting.

Results. — In the 36 samples taken at Point Pinos (Fig. 1) the
mean population for sample group 1 (0-14 m) was 518.6 larvae per
quadrant, for sample group 2 (15-28 m) 67.1 larvae, and for sample
group 8 (29—160 m) 5.1 larvae. The t and Mann Whitney U tests,
performed to compare the three groups, passed the 95% confidence
limits. There was a significant difference in numbers of larvae as
the distance from the outfall increased. No larvae were found beyond
35 m from the outfall. In 22 quadrants containing algae, few larvae
were found, except for one sample at 23 m which contained 114
larvae. Very few larvae were in habitats other than organic slime.
For example, the sample from the Mytilus bed at 25 m from the
outfall contained no larvae, whereas at 35 m the sample, the only
one of slime in the third area, had 79 larvae. In a quadrant at 9
m, 488 larval tubes were attached to dead Balanus; much slime
was also present on the rock. No larvae of P. alaskensis were found
in the samples from the control areas.

At increasing distances from the outfall the number of attached
algae and sessile invertebrates increased and P. alaskensis larvae de-
creased. At increasing distances the slime on rocky outcrops, which
extended at least 160 m beyond the outfall, also decreased. The
larvae seem to thrive when they are not in direct competition with
other organisms for substrate, although Saunders (1928) stated that
“in spring and early summer on the Pacific Coast of Canada they
may be found in almost any matted growth of filamentous algae.”
Saunders also indicated that sufficient algae were neccessary to harbor
larvae. This was not true at Pt. Pinos where larvae subsisted on
organic detritus.

JULY 1972] CHEESEMAN & PREISSLER- — PARACLUNIO LARVAE

207

The large volume of fresh water effluent may be a significant factor
for the abundant P. alaskensis larvae and sparse intertidal inverte-
brates and algae. The effluent is quickly diluted beyond 14 m from
the outfall by sea water and this dilution may also lead to declining
numbers of P. alaskensis larvae. An unpublished study done at
Hopkins Marine Station showed that the salinity is approximately
1,000 ppt within the first 14 m beyond the outfall at high tide
instead of the normal 3,300-3,500 ppt. Chironomidae are generally
found in fresh water and P. alaskensis may do best in areas of fresh
water flow into the intertidal. Studies will be conducted to sample
areas for P. alaskensis where fresh water streams containing no sewage
empty into the rocky intertidal.

The success of P. alaskensis at Pt. Pinos may also be partially
attributed to its reproduction by copulation. The female lays her
fertile eggs in rocky crevices, whereas most intertidal invertebrates
and algae shed their gametes directly into sea water. In an environ-
ment polluted with effluence and chlorine, the fertilization of gametes
may be adversely affected.

Conclusions. — P. alaskensis larvae are abundant in the area of the
sewage outfall at Pt. Pinos where food in the form of organic slime
is plentiful on boulders and rocky vertical outcrops. A possible attri-
buting factor for the success of P. alaskensis is little or no competition
for space from other organisms not physiologically suited to polluted
water or to diluted salinity. It is possible that P. alaskensis larvae
could be used as an index of human sewage pollution in intertidal
areas where there is a primary sewage outfall and a rocky substrate.

Acknowledgment. — We express our gratitude to Hopkins Marine
Station of Stanford University which provided facilities for this study
and to Dr. W. Lee and J. Rote of Hopkins Marine Station who
offered many useful suggestions.

Literature Cited

Saunders, L. G. 1928. Some marine insects of the Pacific Coast of Canada.

Ann. Entomol. Soc. Amerlc], 21(4) : 521-545.

208

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Biology of the Zebra Caterpillar, Ceramica picta 1

(Lepidoptera: Noctuidae)

George Tamaki, R. E. Weeks, and B. J. Landis
Entomology Research Division, Agr. Res. Serv., USDA, Yakima, W ashington 98902

In the Yakima valley, several noctuid species, the bertha army worm,
Mamestra configurata Walker, the beet armyworm, Spodoptera exigua
(Hiibner), the yellowstriped armyworm, Spodoptera ornithogalli
(Guenee), and the zebra caterpillar, Ceramica picta (Harris), spo-
radically attack sugarbeets. Of these species, the zebra caterpillar is
generally the most damaging, and in late autumn in some years, it
so severely defoliates sugarbeets that only the leaf petioles are left
above ground. Therefore, from 1967 to 1971, the zebra caterpillar
was reared and studied at the Potato, Pea, and Sugarbeet Insects
Investigations laboratory at Yakima, Wash., and tests of the sex
pheromone of the female were made in cooperation with the Pesticide
Chemicals Research Branch (Wallis et al. 1972). In addition, Tamaki
et al. (1972) reported life tables which can be used to evaluate the
rearing procedure of the zebra caterpillar. The present paper reports
details of the life stages to supplement the description of Payne (1918) .

Materials and Methods

A series of tests was made to determine the reproductive potential,
optimum incubation temperature, the number and size of the larval
instars, and the duration of instars.

The reproductive potential of the adult female zebra caterpillar was
studied by counting the number of oocytes and matured eggs in 25
one-day to six-day-old females. The actual reproduction of the species
was estimated by placing 12 pairs in individual oviposition cages
and counting the numbers of eggs laid. The cages were 473-ml ice
cream cartons lined with paper toweling for ovipositional sites and
with wet vermiculite in the bottoms to maintain a high humidity and
a cotton wick saturated with 10% sucrose solution for food.

The duration of the incubation of the egg was investigated by
taking an egg cluster about 12 hours old from each of four females (laid
on paper toweling), separating each into six parts, and placing each
part in a ventilated plastic jelly cup. Then four jelly cups (one from

1 In cooperation with the College of Agriculture, Research Center, Washington State University,
Pullman 99163. Received for publication.

The Pan-Pacific Entomologist 48: 208-214. July 1972

JULY 1972] TAMAKI ET AL. — CERAMICA PICTA BIOLOGY

209

each cluster) were placed in a glass gallon-size jar and exposed in a
temperature control cabinet maintained at 10, 15.6, 21.1, 26.7, 32.2,
or 37.8° C. A wire screen stage held the cups above a saturated solution
of calcium nitrate salt placed in the bottom of each jar to maintain
the relative humidity within a range of 50-70%.

To determine the number of larval instars and the differences in
size between instars, we placed several hundred neonate larvae in a
tray with sugarbeet leaves and in another tray on Shorey’s (1963)
bean-based artificial diet (the primary diet used to rear zebra cater-
pillars in the laboratory). Each treatment was replicated three times.
Then 10 randomly selected larvae from each treatment were removed
every 12 or 24 hr (rotated between trays) , preserved in 70% alcohol,
and measured to determine the greatest width of the head capsule
and length of body. The head capsules of field-collected zebra cater-
pillar larvae found on sugarbeets in the autumn were also measured.

The developmental rate of the life stages of the zebra caterpillar
was determined by rearing a total of 600 caterpillars (200/treatment)
in the laboratory with three procedures. Procedures one and two
differed only in that the diet was sugarbeet leaves or Shorey’s (1963)
artificial diet. Otherwise, 100 neonate larvae per tray were reared
with a technique similar to that used for the tobacco hornworm,
Manduca sexta (L.) (Yamamoto 1969). Thus, the larvae were checked
daily for stage of growth, and the trays were cleaned; however,
fresh leaves were provided two to three times per week when the larvae
were small (changed daily in later instars) . Procedure three involved
rearing 200 caterpillars singly on about 15 ml of Shorey’s diet in
30-ml jelly cups capped with tight lids (no food was added to the
cups). All 600 larvae were from one egg cluster from one female
and were held in the same room at an average rearing temperature
of 25 °C (18-28°) with a 16-hr photophase. Then as the last-instar
larvae changed to the prepupal stage, the insects were transferred to
vermiculite for pupation. Pupae were easily sexed using the characters
illustrated by Butt and Cantu (1962) for sexing pupae of other
Lepidoptera.

Results

The reproductive system of the female zebra caterpillar contains a
pair of ovaries with a total of eight ovarioles. In a newly emerged
female, an ovariole is 10-12 cm long and filled with a single strand
of oocytes. Egglaying (reproductive) potential determined by dis-
secting nonparous virgin females showed that the zero- to one-day-old

210

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Table 1. Mean hatching time of eggs of Cer arnica picta held at
various constant temperatures (from adults reared on artificial diet),
December 1970.

Temperature

°C

No. of eggs
incubated

No. of eggs
hatched

Mean hatching time
(days)

38.0

512

0

—

32.2

356

0

—

26.7

294

294

5.0

21.1

279

275

6.2

15.6

388

290

12.3

10.0

365

0

—

females contained no mature eggs (eggs with ribbed rows of chorion
were indexed as mature) ; the two-day-old females had 40% mature
eggs; and the three- to six-day-old females had 50-58% mature eggs.
In these older females, egg laying had been delayed because fertilized
females normally lay their eggs two to three days after emergence.
The total number of oocytes plus mature eggs per female in 25
females averaged 1,236 (range 884-1,884). The average number of
eggs actually deposited by the 12 mated females in the oviposition
cages was 986 (range 59-1,998) . Therefore, many females were
apparently able to lay their full complement of eggs, but a few laid
only a small proportion.

The effects of temperature on the rate of development of eggs of
the zebra caterpillar are shown in Table 1. Eggs did not hatch at
high temperatures (32.2°C and 38°C), but at 26.7°C, 100% egg
hatch occurred within five days, the shortest hatching time; this
temperature was therefore designated as optimum. At 21.1 °C, 99%
hatch occurred, but the incubation time was about 20% longer than
at the optimum temperature. At 15.6°C, hatch was still relatively
high (75%), but the incubation time was increased by 140% over the
optimum. No egg hatch occurred at 10°C.

Payne (1918) reported that the zebra caterpillar had five larval
instars in Nova Scotia. However, he presented only a range of
measurement for use in separating the instars and gave no average
measurements for the width of the head capsules and the length of
the bodies. In our studies in eastern Washington, both the laboratory-
reared and the field-collected caterpillars had six larval instars (Table
2). However, the range of width of the head capsules of the second
instar given by Payne (1918) fell between our measurements for the
second and third instars.

JULY 1972] TAMAKI ET AL. — CERAMICA PICTA BIOLOGY

211

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 43, NO. 3

Table 3. Pupal weight of male and female Ceramica picta reared
by three procedures.

Rearing methods

Weight of pupa

(mg ± SE) a

$

Artificial diet — in cups

496.5 ± 9.08

600.8 dr 15.53

Artificial diet — in tray

481.9 ± 14.68

556.4 ± 25.93

Sugarbeet leaves — in tray

388.3 ± 3.13

434.0 dz 4.15

a Standard error (SE).

We found no significant differences in the size of the capsules of
the same instar among laboratory-reared larvae fed artificial diet,
laboratory-reared larvae fed sugarbeet leaves, or field-collected larvae.
However, the measurements for instars did differ significantly for
all larvae (Table 2) . Also, diet did not cause any significant differences
in the length of a given instar, and the differences in length between
succeeding larval instars were not statistically significant.

The developmental periods for each life stage of the zebra cater-
pillar reared in the laboratory on sugarbeets and on artificial diet
are illustrated in Fig. 1. At a relatively high rearing temperature
(average 25°C), the length of the first to fifth larval stadia reared
on sugarbeets ranged from two to five per stage; the sixth stadium
averaged about five days though some individual insects took as much
as 10 days. The length of the larval stadia for larvae reared on
artificial diet in cups was similar to that of larvae reared on sugar-
beets, but when the larvae were reared in trays, a few in the later
instars had an extended developmental period. Most larval develop-
ment was completed after 27 days on all diets. Also, except for the
decreasing number of insects surviving on artificial diet, no striking
differences were evident in the developmental periods of the larval
instars when the larvae were reared by the three procedures. In
contrast, Tamaki and Weeks (1972) reported that the Geocoris ssp.
fed an incomplete diet had a prolonged rate of development in the
later nymphal instars which was associated with high mortality.

The duration of the prepupal stage was difficult to assess because
the cocoons had to be disturbed to check the developmental stage;
therefore, checks were infrequent. The average prepupal period was
estimated at five days, but a few insects reared on artificial diet in
trays had an extended prepupal period (Fig. 1).

The average pupal periods for zebra caterpillars reared on sugar-
beet leaves in trays and on artificial diet in trays and in cups were

JULY 1972] TAMAKI ET AL. — CERAMIC A PICTA BIOLOGY

213

DAYS

Fig. 1 . The developmental periods of all life stages of zebra caterpillars
reared on artificial diet and on sugarbeets by three methods.

214

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

34 days (range 16-52), 30 days (range 15-40), and 27 days (range
14-41), respectively. Female pupae were heavier than male pupae
(Table 3). Pupae from larvae reared on sugarbeet leaves weighed
less than pupae from larvae reared on artificial diet, and the heaviest
pupae were obtained from larvae grown on artificial diet in cups
(these pupae also had the shortest duration of the pupal stage) .
However, as shown in Fig. 1, the number of moths was much greater
when the larvae were reared on sugarbeet leaves than on artificial
diet.

The first emergence of adults occurred at 45 days, but the mean
time for females to reach maturity by all three rearing procedures
ranged from 62 to 67 days. The preoviposition period averaged two
days; the oviposition period averaged seven days (though some
females had an extended period that lasted as much as 15 days) ;
and the postoviposition period averaged 0-1 day. When the larvae
were reared on sugarbeet leaves, the average lifespan of the female
was 10.5 days (range 2-19), and that of the male was 12 days
(range 2—21).

Literature Cited

Butt, B. A., and E. Cantu. 1962. Sex determination of lepidopterous pupae.
USDA ARS 33-75, 7 p.

Payne, H. G. 1918. The zebra caterpillar ( Ceramica picta Harris). Proc. Entomol.
Soc. Nova Scotia 1917, 3: 44-49.

Shorey, H. H. 1963. A simple artificial rearing medium for the cabbage looper.
J. Econ. Entomol., 56: 536-7.

Tamaki, G., and R. E. Weeks. 1972. Biology and ecology of two predators,
Geocoris pallens Stal and G. bullatus (Say) . USDA Tech. Bull., 1446.
46 p.

Tamaki, G., J. E. Turner, and R. L. Wallis. 1972. Life tables for evaluating
the rearing of the zebra caterpillar. J. Econ. Entomol., 65: 1024-7.
Wallis, R. L., L. I. Butler, and L. M. McDonough. 1972. Sex pheromones of
the zebra caterpillar: Bioassay and preliminary chemical investigations.
J. Econ. Entomol., 65: 945-7.

Yamamoto, R. T. 1969. Mass rearing of the tobacco hornworm. II. Larval
rearing and pupation. J. Econ. Entomol., 62: 1427-31.

July 1972]

FENDER — TWO NEW SILIS

215

Two New California Species of Silis 1

(Coleoptera: Cantharidae)

Kenneth M. Fender

Linfield Research Institute, McMinnville, Oregon 97128

The two species herein described as new to science, help confirm
Green’s hypothesis (1966) that numerous unrecognized species of Silis
await discovery, especially from western North America.

In the genus Silis, the posterior angles of the pronotum are excised,
retracting these angles and forming an anterior subangulate prominence
called the anterior process. The excised portion is occupied by a pro-
jected portion of the hypomeron, called the posterior process. Both
of these processes vary considerably in outline and are excellent for
the recognition of species groups. In some instances they appear to be
adequate for the recognition of species.

The characters for the determination of the species within the
species groups are usually found in the aedeagus of the male. As
Green (1966) has noted, a certain amount of variability in the
aedeagae resulted in his recognition of some species complexes that
he was unable to resolve. The attemped resolution of these is not yet
possible.

Silis ( Silis ) spinigerula Fender, new species

(Figs. 1, 3-5)

Black. Mandibles testaceous, becoming infuscate towards tips. Prothorax pale
rufous. Pubescence piceous.

Male. — Length 5 mm, length-width ratio 3.25 : 1. Antennae four-fifths length
of body, moderately stout, subfiliform, becoming compressed apically, intermediate
segments about three and one-half times as long as wide. Pronotum as in Fig.
1, anterior process of basal excisure bilobed, lobes rounded, posterior process
produced as forward curving spiniform process, excision only partially closed by
dorsal surface of extended hypomeron. Pubescence erect, fine, short and in-
conspicuous. Aedeagus. Dorsal plate broadly shallowly emarginate apically,
without raised median lobe. Basophyses fused into broad subparallel-sided plate
about half as wide as tegmen and with apex shallowly emarginate. Median lobe
with two pair of apical spines; in dorsal or ventral views, lower pair abutting
with tips blunt, upper pair remote with tips acute; in lateral view, both pairs
acute.

Female. — Unknown.

1 This study was supported by National Science Foundation Grant: GB-6283X.

The Pan-Pacific Entomologist 48: 215-217. July 1972

216

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

Figs. 1, 3-5. Silis ( Silis ) spinigerula. Fig. 1. pronotum. Fig. 3. aedeagus of
male, dorsal view. Fig. 4. same, lateral view. Fig. 5. Same, ventral view.
Figs. 2, 6-8. Silis (Silis) alexanderi. Fig. 2. pronotum. Fig. 6. aedeagus of male,
dorsal view. Fig. 7. same, lateral view. Fig. 8. same, ventral view.

Holotype male , Campo Lake on Mexican Border, San Diego County,
California, 2 April 1963, 2,565 feet, collected by C. P. Alexander, in
the collection of the California Academy of Sciences, San Francisco.

This species is most closely related to S. filicornis Van Dyke. That
species differs in not having the anterior process of the basal excisure
of the pronotum bilobed, having the pubescence decumbent and the

july 1972]

FENDER — TWO NEW SILIS

217

dorsal plate of the aedeagus with an apical raised and produced
median lobe.

Silis ( Silis ) alexancleri Fender, new species

(Figs. 2, 6-8)

Black. Prothorax and head in front of antennae pale rufous; palpi black;
basal two or three antennal segments testaceous beneath; head beneath pale
rufous at gular sutures. Protibiae pale piceous to dark piceous. Apical angles
of first four visible abdominal sternites narrowly testaceous. Pubescence cinereous.

Male. — Length 4.75 to 5.25 (average 5) mm. Length-width ratio 2.75 : 1.
Antennae filiform, about three-fourths length of body, intermediate segments
three times as long as wide. Pronotum as in Fig. 2, excision of posterior angles
of pronotum deep, rather narrow; posterior margin of anterior process with two
spicules, outer spicule as produced posterior margin of posterolateral furrow,
inner spicule simple; posterior process broad, subfoliate in outline. Aedeagus.
Dorsal plate apically entire, rounded, becoming sinuate towards each side. Median
lobe with an apical pair of short acute spines. Basophyses (lateral view) stout,
evenly curved up and narrowing apically, not attaining margin of dorsal plate.
Basal plate (ventral view) short and stout, apex shallowly notched.

Female. — Normal, not recognizable unless taken in association with males.

Holotype male , allotype female and 13 male and 4 female paratypes:
Seven Oaks, San Bernardino Mountains, San Bernardino County,
California, 21 May 1963, 5,000 feet, collected by C. P. Alexander,
types in the collection of the California Academy of Sciences, San
Francisco.

This species somewhat resembles the members of the cava group
of Silis. In these, the anterior process of the basal excisure of the
pronotum is simple, the posterior margin without spicules.

Named for the collector, C. P. Alexander, in appreciation of the
many novelties that he has donated to my collection and the great
interest that Mrs. Alexander and he have continually shown in my
work.

Literature Cited

Green, J. W. 1966. Revision of the Nearctic species of Silis (Cantharidae:

Coleoptera) . Proc. Calif. Acad. Sci., 32(16): 447-513, 65 figs.

218

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

SCIENTIFIC NOTES

Notes on Diaulota harteri with new synonymy (Coleoptera: Staphy-
linidae) . — Since the description of Diaulota harteri Moore and D. megacephala
Moore (Trans. San Diego Soc. Natur. Hist., 12: 103-152, 1956) many more

specimens have been collected. In all cases specimens of the former have been
females and specimens of the latter males. I am convinced that the feeble lobe
on the posterior margin of the sixth sternite of the holotype of D. harteri is a
small variation of a female specimen. Consequently, I propose the following
synonymy :

Diaulota harteri Moore, 1956: 123

Synonym D. megacephala Moore, 1956: 124 NEW SYNONYMY
I have seen a good series of this species collected from intertidal rock crevices
at Gaviota Beach, Santa Barbara County, California, 23 March 1971, by Derham
Giuliani. This is a northern extension of range for the species. — Ian Moore,
Division of Biological Control, University of California, Riverside, 92502.

Xystridiogaster Scheerpeltz, a synonym of Paracraspedomerus Moore
(Coleoptera: Staphylinidae) . — The genus Paracraspedomerus was proposed
by Ian Moore (1960, Pan-Pac. Entomol., 36: 99-101) for Cafius speculi-
frons Fauvel. Otto Scheerpeltz described the genus Xystridiogaster (1966,
Naturhist. Mus. Wien Ann., 69: 393-418) based on the same type species.
Since Xystridiogaster is isogenotypic with Paracraspedomerus, I propose the
following synonymy:

Paracraspedomerus Moore, 1960

Xystridiogaster Scheerpeltz, 1965 NEW SYNONYMY
— Ian Moore, Division of Biological Control, Department of Entomology, University
of California, Riverside, 92502.

Notes on dry-season survival in two species of Elmidae (Coleoptera).

— During June 1969, I visited a small stream which empties into the
Navarro River near Philo, Mendocino County, California. The only water which
remained in the stream was in the form of small, widely scattered pools. I
selected a spot approximately midway between two such pools about 200 feet
apart and attempted to determine, by digging into the stream bed, whether
any subsurface flow of water connected the pools. After digging down about
16 inches, I encountered water, then dug a few more inches to form a small
basin. I watched the debris settle and was able to detect a slight flow evidently
connecting the pools. Much to my surprise, I noticed several apparently dead
Elmid beetles drifting about at the bottom of the excavation. These beetles
feign death when disturbed so I observed them carefully for a few minutes
for signs of activity. After several minutes with no response, I transferred
the beetles to a collecting pan containing about one inch of water. About ten

July 1972]

SCIENTIFIC NOTES

219

minutes later the beetles began to evidence signs of life and started crawling
about the bottom of the pan. Careful further excavation resulted in a number
of additional specimens and demonstrated clearly that these insects were actually
surviving in the sandy gravel bed and had not merely fallen into the hole during
my digging.

The beetles were subsequently identified as Zaitzevia parvula (Horn) and
Narpus angustus Casey. No other adult or larval aquatic insects were found
in the gravel beds, but the few remaining pools supported abundant insect life.
This same creek was revisited in November, after the annual winter rains had
begun, and these two species were found clinging to rocks in flowing water,
the more usual niche for these insects.

Perhaps the small size of these elmids permitted them to work their way into
the gravel bed, either from above as the stream dried up, or outward from the
pools along the flow paths. Of the two, I suspect the former to be the most
likely. This experience would suggest that some species of Elmidae routinely
aestivate, or at least survive seasonal drying, within the beds of intermittent
streams. Such behavioral adaptation has obvious survival benefit in any area
subject to such seasonal fluctuation in water levels. — L. Neil Bell, 141 Mt.
Shasta Court, Clayton, California 94517.

SOCIETY ANNOUNCEMENT

The Pacific Coast Entomological Society announces the publication
of a new book in its Memoir Series: Volume 4: Robert Leslie
Usinger: Autobiography of an Entomologist.

This unique book, written by one of the outstanding entomologists
of our time, relates personal experiences and episodes from boyhood
through his professional career. Included are accounts of research
expeditions to various countries.

Edited by Drs. E. G. Linsley and J. L. Gressitt, the cloth-bound
book consists of 343 pages, 38 illustrations, and dust jacket. The
book was published in August 1972, price $15.00 (postage extra
and 5% state sales tax to be added for California residents) . Members
of the PCES will receive a 20% discount.

Orders for this limited edition (1,000 copies) may be placed with:

Dr. Paul H. Arnaud, Jr., Treasurer
Pacific Coast Entomological Society
c/o California Academy of Sciences
Golden Gate Park
San Francisco, California 94118

Please make checks payable to the Pacific Coast Entomological Society.

220

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 3

ENTOMOLOGICAL EXCHANGES AND SALES

Space is available to members and non-members who wish to advertise entomo-
logical exchanges, sales, announcements, etc. We can offer an ad in four issues
for $1.50 per line. Ads will be arranged in order of their receipt. Contact the
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fornia, Davis, California 95616.

SHARP & MUIR: THE COMPARATIVE ANATOMY OF THE MALE GEN-
ITAL TUBE IN COLEOPTERA. The classic 1912 monograph and six other papers
by the same authors; 304 pp., 43 pis., bound. $10.00. An essential book for all
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Current developments in the sugarbeet industry in the United States are pin-
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OCTOBER 1972

Vol. 48

No. 4

THE

Pan-Pacific Entomologist

MARER — An eye deformity in a tarantula spider, Aphonopelma r ever sum

(Araneae: Theraphosidae) 221

MAYO — New species of the genus Baetodes (Ephemeroptera: Baetidae) — 226

FENDER — A new species of Rhagonycha from Alaska (Coleoptera: Can-

tharidae) 242

WIRTH AND RATANAWORABHAN — A new genus of biting midge from

California related to Neurohelea Kieffer (Diptera: Ceratopogonidae) 244

HALSTEAD — Notes and synonymy in Largus Hahn with a key to United

States species (Hemiptera: Largidae) 246

MOORE AND LEGNER — Two new species of Orus from California (Coleop-
tera: Staphylinidae) 249

PINTO — Notes on the Caviceps Group of the genus Epicauta with descrip-
tions of first instar larvae (Coleoptera: Meloidae) 253

SUMMERS AND WITT — Nesting behavior of Cheyletus eruditus (Ac.arina:

Cheyletidae) 261

MARTIN — A new species of Leptopteromyia (Diptera: Leptogastridae) 270

CLEMENT — Notes on the biology and larval morphology of Stenodynerus

canus canus (Hymenoptera: Eumenidae) 271

SCIENTIFIC NOTES 277, 278

CORRECTION 243

BOOK REVIEWS 252, 269, 276

BOOK NOTICE 276

SOCIETY ANNOUNCEMENT — Usinger Autobiography 279

INDEX TO VOLUME 48 _ 281

SAN FRANCISCO, CALIFORNIA • 1972

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

THE PAN-PACIFIC ENTOMOLOGIST

EDITORIAL BOARD
R. W. Tiiorp, Editor

E. G. Linsley R. 0. Schuster, Asst. Editor E. S. Ross

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P. H. Arnaud, Jr. Treasurer W. H. Lange, Jr. Advertising

Published quarterly in January, April, July, and October with Society Proceed-
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The Pacific Coast Entomological Society

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Title of Publication: The Pan-Pacific Entomologist.

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The Pan-Pacific Entomologist

Vol. 48

October 1972

No. 4

An Eye Deformity in a Tarantula Spider,
Aphonopelma reversum

(Araneae: Theraphosidae)

Patrick J. Marer

United States International University, Elliott Campus, San Diego, California 92124

A recent study of the fauna inhabiting Palomar Mountain in northern
San Diego County, California, produced many specimens of Mygalo-
morph spiders, including several hundred tarantulas of the species Apho-
nopelma reversum Chamberlin. This is the most common large taran-
tula in the southwestern corner of the United States, its distribution
being fairly uniform throughout the county.

During the course of examining these specimens, one specimen,
Number 51, was discovered to have a severe eye structural anomaly.
This deformity is believed to be of a genetic or developmental origin.
Figures 1 and 3 are drawings of this unique structure, while Figures 2
and 4 are drawings of a normal eye structure in a male Aphonopelma
reversum , collected on the same day.

This specimen was obtained in a pitfall trap, as described by Banta
(1957) . The trap was located at station number 55 on the Nate Harrison
Grade road, leading up the southwestern slope of Palomar Mountain
from Pauma Valley. Station number 55 is at an elevation of 1,650 feet.
The spider was a freshly molted, mature male and was collected on 22
August 1970.

The extent of this eye deformity includes an irregularly shaped eye
tubercle, malformation or dislocation of all but one eye, and coloration
and hair differences in the eye area compared to a normal specimen

(Fig. 1).

The only eye that is not malformed or dislocated is the left posterior-
lateral eye. The left anterior-lateral eye appears somewhat in its proper
place and its base is of the correct size and shape. The eye itself extends
laterally to the left about twice its normal length. The whole segment is
covered with lens tissue. There is an apparent attempt for this eye to
divide into two or possibly three segments.

The posteromedian eye on the left side is in its normal position, al-

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Fig. 1. Dorsal view of abnormal eye tubercle of Aphonopelma reversion, speci-
men number 51. Note many oval shapes and darker matrix in central eye, as well
as irregular shape of tubercle.

though its separation from the adjacent posterolateral eye is greater than
the record for this species. This distance is about the width of the
posterolateral eye, while in a normal specimen there is little or no sepa-
ration between these eyes. The eye is larger than normal and is partially
obscured by the anomalous eye anterior to it (Fig. 1).

The right lateral side of the eye bulb has no definite eyes, although
in the location where the right anterolateral eye would normally be there
is found a mass of eye tissue, including lenses, which has divided into
10 different circular shapes, resembling small eyes of various sizes.
There are four additional shapes on the anterior aspect of the bulb,
nearer the midline. These four “eyes” are greatly separated from each
other as compared to the close proximity of the eye structures located on
the right lateral aspect.

In the midplane of the eye tubercle there is one other large eye struc-
ture which is extremely complex, appearing to divide into many segments
(Fig. 1). This is a teardrop shaped structure, more than twice the size
of a normal eye. It is positioned to the left side of the eye tubercle,
rather than in the center. The large end of the teardrop is anterior.
There is an extra bulb on the right anterolateral aspect composed of
additional lens tissue, and appearing to form two oval shaped eyes. The
main body of this structure resembles a large eye with a smooth lens,

OCTOBER 1972] MAKER TARANTULA EYE DEFORMITY

223

.* .• I 1 . /

Fig. 2. Dorsal view of normal eye tubercle of Aphonopelma reversum, specimen
number 42.

except the lens contains a complex dark brown marking, separating it
into 12 oval and round shapes of various sizes. Each shape studied indi-
vidually resembles an eye. This entire eye structure has the characteristic
tissue bordering it that is found around normal eyes.

The eye tubercle is considerably asymmetrical (Fig. 1 and 3). The
left side is more oblong from its midline to its lateral terminus than the
right. The right half, divided along a mid-transverse line, is broader
from front to back than the left. Viewed anteriorly, the tubercle is higher
on the left side, corresponding to the large anomalous eye structure near
the midline. The tubercle is nearly twice as thick in this location as the
normal eye tubercle (Fig. 4) .

The eye tubercle is lighter in color on the right side. The area bounded
by the left anterior and posterolateral eyes, median eye and anomalous
central eye is black in color. The rest of the tubercle is reddish brown,
lighter on the right side near its anterior margin. All the lens areas
appear pearl colored, those of the right side with a yellowish cast. The
normal eye tubercle in this species has a black marking running between
the anterior and posterior rows of eyes in a lateral direction along the
entire width of the eye tubercle (Fig. 2) .

The cephalothorax in the area of the eye tubercle is lighter colored on
the right side, and hair in this area is more sparse than on the left. This

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

1.0 MM

Fig. 3. Anterior view of eye tubercle of Aphonopelma reversion, specimen
number 51, showing asymmetry of this structure.

hair is growing in irregular patches in many different directions, as
opposed to a normal, more symmetrical growth pattern of hair.

The rest of the body and appendages of this spider are normal in
color, structure and appearance and compare favorably with other speci-
mens of mature male Aphonopelma reversum.

Due to the complexity of this abnormality, coupled with its vital loca-
tion — the eye tubercle — it seems unlikely that the deformity could have
resulted from an injury. It is apparent that the ectodermal layer which
produced the eye structure has either been damaged or infolded during
formation or in some other way is not complete. An injury to this area
of the tarantula’s body, sufficient to damage the underlying germ layer,
would most likely result in the death of the spider due to loss of blood.
A molt associated injury would seem to produce the same results or
cause irreparable damage to underlying structures. This is a vital area
of the tarantula’s body (Firstman, 1954) .

This specimen was captured during the peak activity period for male
tarantulas of this species. According to observations of the activities
of this species, this peak activity period is preceded by the final molt of
the male (Petrunkevitch, 1934) . This particular specimen’s condition
and color indicates that it had molted about one week or so before being
caught in the trap. Any injury during this final molt would not have
healed within this period of time even if the spider could have survived
the loss of blood or damage to tissues.

The only possible remaining causes of this deformity would be of either
a congenital or a developmental nature. This could explain the many

OCTOBER 1972] MARER TARANTULA EYE DEFORMITY

225

Fig. 4. Anterior view of eye tubercle of Aphonopelma reversion, specimen
number 42, showing appearance of a normal structure.

eye shapes and attempts to form eyes. The divisions within the central
eye and the smaller lateral eye segments suggest that the structure at-
tempted to divide, but the number of irregular divisions and the mis-
placement of eyes indicates that control of the process was lost or the
process itself was interrupted by some factor.

Vision in this genus of Mygalomorph spiders is very poor, the eyes
possibly being used only as photoperiod receptors. It is doubtful, there-
fore, that this deformity had any serious effect on the life of this partic-
ular specimen, especially considering that it had reached sexual maturity,
which takes as long as 12 years for this species.

Literature Cited

Bant a, B. H. 1957. A simple trap for collecting desert reptiles. Herpetologica,
3: 174-176.

Firstman, B. 1954. The central nervous system, musculature, and segmentation
of the cephalothorax of a tarantula ( Eurypelma californicum Aus-
serer) . Microentomology, 19: 14-40.

Petrunkevitch, A. 1934. New observations on moulting and mating in Taran-
tulas. J. N. Y. Entmol. Soc., 42: 289-296.

226

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New Species of the Genus Baetodes

(Ephemeroptera: Baetidae)

Velma Knox Mayo
2702 E. Seneca St., Tucson, Arizona 85716

The genus Baetodes is known at the present time from Arizona and
Texas in the United States, and from Mexico, Central and South Amer-
ica. In Central America it has been taken from Guatemala, Honduras,
Costa Rica, El Salvador, Panama and the Canal Zone. In South Amer-
ica it has been collected in Brazil, Venezuela, Ecuador, Peru, and Bo-
livia.

The genus was established in 1924 by Needham and Murphy. A
review of the characteristics of the genus by Needham and Murphy
(1924), by Traver (1943, 1944) and by Edmunds (1950) was included
in a paper by the writer (Mayo, 1968). Traver (1943) described the
only known adult, Baetodes spiniferum. She requested that the writer
emend the spelling on her B. spiniferum to Baetodes spinifer. Cohen and
Allen (1972) have described nine new species of Baetodes from Mexico
and Central America.

Generic Diagnosis

Nymphs. — Head (Fig. 16, Mayo, 1968). Ocelli prominent, two large ones
laterad of epicranial fork, and small one, below these within fork. Mouthparts as
follows: Labrum with sclerotized band along lateral margins and thick fringe of
setae around margin, apical margin with cleft at midline, row of stout, prominent
spines of varying length and number and arranged in two groups (one near mid-
line, and one closer to lateral margin) . Behind apical margin with numerous scat-
tered, shorter setae behind these (Figs. 5, 8) . Mandibles with thumblike process at
base of molar surface of left mandible that varies in size and shape in different
species, rim of molar surface near the thumb with one tooth larger than others
(Fig. 24) , lacinia well developed on left mandible, situated close to incisors (Fig.
17), lacinia on right mandible represented by small, slender, pointed process
situated half way between molar surface and incisors. Highly sclerotized “anterior
articulatory process” (Traver and Edmunds, 1968) extends parallel and posterior
to molar surfaces on both mandibles; right mandible with short bristle extending
from rim of molar surface (Fig. 4) . Maxillae with palpi two segmented (Fig. 22) ,
and crowns of maxillae with fringe of long, stout setae (Fig. 16) . Labium with
palpi three segmented (Figs. 12, 15), apical segment of labium with spines (Figs.
12, 15), paraglossae with thick brush of curved pectinate spines of varying lengths
and curving mesally with all pinnae directed mesally (Fig. 2), glossae with setae
at tips (Fig. 12). Thorax with anterior border of pronotum concave and antero-
lateral corners acute (Fig. 9) , hind wings absent, narrow, elongated cervical
sclerite articulates with posterior border of propleuron, and extends to head on

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OCTOBER 1972]

MAYO NEOTROPICAL MAYFLIES

227

either side, high dorsal crest on each coxa, legs unusually long and slender, tibiae
slightly longer than femora and tarsi slightly less than % as long as tibiae, femora
with variable number of stiff spines along dorsal crests, tibiae distinctly marked
with narrow strip which extends over % of dorsal surface and terminates at ventral
margin, shaft of tibia beyond this point abruptly more narrow and narrow point of
segment marked with diagonal streak (Fig. 14), fringe of setae along dorsal crests
of femora and tibiae, femoral and tibial articulations heavily sclerotized, claws each
with long, stout, colorless bristle originating near base of first marginal denticle
and curving out as far as the tip of claw (Figs. 3, 6), variable number of denticles,
decreasing in size rearward. Abdomen with simple gills on abdominal terga 1-5
only, gills attach near low pleural fold. Caudal filaments without setae, terminal
filament short stub and correspondingly narrower than lateral cerci.

Discussion

The spines behind the apical margin of the labrum are prominent,
varying in length, and are often alternated, one long, one short (Fig. 5) .
According to Traver and Edmunds (1968), Macan (1961) “shows how
the pattern of spines on the dorsal surface of the labrum serves to differ-
entiate four different species of Baetis .” The spines behind the apical
margins of the labra of Baetodes described in this paper are distinctive
for each species except for B. obesus (Fig. 32) and for B. sancticatarinae
(Fig. 45) . Those of B. sancticatarinae are much farther from the apical
margin than those of B. obesus.

In some species a hump or tubercle occurs on the posterior border of
the pronotum.

The spines along the dorsal crests of femora of most species are prom-
inent but are difficult to detect on B. obesus. On one species examined
they were invisible under the binocular microscope, but three yellow
spines were visible under high power of the compound microscope.

Discussion of the long bristle behind the claw was omitted from the
descriptions of B. spinae and B. levis and from Fig. 19 (Mayo, 1968).

There are no tubercles on the abdomen of the species herein described.
Abdominal tubercles occur on the majority of species and where there are
tubercles on the abdomen there is usually one on the metanotum. These
tubercles are always unpaired, arising from the posterior border at the
midline. Some are very small, directed rearward and some are erect and
prominent.

On many species elongated fingerlike gills occur at the bases of all
three pair of coxae. Some of these are single, and some are paired.
There are three different sizes, long, intermediate, and small. The
largest, and by far the most common, are four times the length of the
smallest size.

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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Many species have scattered setae on the dorsal surface of the body
segments.

Baetodes chilloni Mayo, new species

(Figs. 1-5)

Female nymph. — Length of body about 4 mm, cerci slightly shorter. General
color brown dorsally. Head brown; darker brown spots on either side midline
between eyes; eyes bordered with yellow; antennae pale yellow. Mouthparts as in
Figs. 1, 2, 4, 5. All intersegmental membranes pale yellow. Thorax with pronotum
brown; anterior border narrowly rimmed with dark brown; mesonotum and wing
pads brown ; numerous fine white setae dorsally on thorax ; pleural sclerites brown ;
unsclerotized areas pale yellow. Coxae and trochanters brown; femora an even
brown except for a large pale yellow spot near joining with trochanters; rimmed
with dark brown at joinings with tibiae; six or seven reddish brown spines on
dorsal crests of femora. Tibiae pale brown with usual white streak entire length.
Tarsi light brown; reddish brown at joining with claw. Claw as in Fig. 3. Sternum
pale yellow except for brown, curved sclerites at apodemes. Abdomen with all
terga an even brown, anterior borders narrowly rimmed with dark brown; without
tubercles. Abdominal gills light yellow, darker on basal half, each coxa with two
fingerlike gills of long type. Sternum 1 pale brown, lighter mesally; 2-9 brown,
6-9 with pale yellow horizontal streak laterally near anterior borders; 8-10 with
triangular yellow area with base along entire posterior borders and apex at midline
on anterior borders; pleural fold yellow. Cerci and terminal filament yellow tinged
with brown distally.

Holotype female nymph, Station V, Rio Chillon, Peru, 500 m, 3
May 1958, J. lilies. In Entomological collection University of Utah,
Salt Lake City. Paratype: 1 nymph, same data, same deposition as
holotype.

Discussion. — General color of thoracic sternum pale yellow; that of
abdominal sternum brown as on dorsal surface. Baetodes chilloni can be
distinguished from B. solus by the presence of coxal gills. The thoracic
sternum of this species is pale, whereas that of B. solus is dark brown.
The labrum differs in details of the spines behind apical margin from
those of B. solus (Fig. 5, 18) . Details of the mouthparts of this species
distinguish it from B. sp. No. 1 (Figs. 7, 8, 10, 20, 24, Mayo, 1968) .

Fig. 1 . Maxilla of B. chilloni. Fig. 2. Labium of B. chilloni. Fig. 3. Claw of
B. chilloni. Fig. 4. Right mandible of B. chilloni. Fig. 5. Labrum of B. chilloni.
Fig. 6. Claw of B. traverae. Fig. 7. Cerci and terminal filament of B. traverae.
Fig. 8. Labrum of B. traverae. Fig. 9. Dorsal view of head and thorax of B.
traverae. Fig. 10. Gill IV of B. traverae. Fig. 11. Left mandible of B. traverae.
Fig. 12. Labium of B. traverae. Fig. 13. Maxilla of B. traverae. Fig. 14. Leg II of
B. traverae.

OCTOBER 1972] MAYO NEOTROPICAL MAYFLIES 229

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Baetodes traverae Mayo, new species

(Figs. 6-14)

Female nymph. — -Length of body about 6 mm, cerci about 7 mm. General color
dark brown; unsclerotized areas yellow. Head more narrow than that of other
species ; area between eyes dark brown ; yellow anterior to eyes and around bases of
antennae; first two segments of antennae yellow, remaining segments pale brown;
mouthparts as in Figs. 8, 11-13. Spines behind apical margin of labrum grouped
together (Fig. 8) ; glossae short (Fig. 12) thumb on left mandible pointed (Fig.
11). Two long bladelike spines among setae on crowns of maxillae. Large ocelli
with black crescent on inner margins; small one rimmed with black on anterior
margin. Thorax with pronotum dark brown along anterior border; irregular yellow
areas as in Fig. 9; anterior margin concave; posterior margin convex so that medial
portion of pronotum less than half as deep as lateral borders. Mesonotum brown
along anterior border; and on either side of midline to scutellum. Two diagonal,
wide yellow areas on scutum at border of wing pads; two dark brown dots close
to midline anterior to scutum (Fig. 9) and laterally, half way between anterior
border and wing pads. Metanotum dark brown. Posterior border yellow lateral to
midline. Thoracic sclerites of pleuron dark brown; unsclerotized areas pale
yellow; femora with wide pale yellow stripe on dorsal surface; dark brown at
joinings with tibiae; tibiae brown (Fig. 14) ; tarsi brown with seven spines along
lower edge; blackish brown at joining with claw; claw with ten denticles (Fig. 6).
Sternum pale yellow except for small brown sclerites around apodemes at postero-
lateral corners of meso- and metasternum. Abdomen with terga darker brown than
thorax; midline with very faint yellow streak on all terga; more prominent on 10;
shaded with yellow at pleural fold ; posterior borders of terga 3-6 yellow to midline.
Without abdominal tubercles; abdominal gills oval, tracheae white (Fig. 10).
Sterna very pale brown; narrow brown markings horizontally on 2-9 on either side
of midline close to anterior border; vertical brown streaks % distance between
anterior and posterior borders near pleural fold on segments 2-9. Cerci light
brown ; terminal filament unusually short stub ; not as long as depth of first segment
of cerci; at base about % width of base of cerci; appears to be unsegmented; better
seen from ventral aspect (Fig. 7) .

Male nymph. — Length of body about 5 mm, cerci 5 mm.

Holotype female nymph, Station II Rio Chillon, Peru, 2500 m,
3 May 1958, J. lilies. In Entomological collection University of Utah,
Salt Lake City. Paratypes five nymphs same data, same deposition as
holotype. Twelve nymphs, station III Rio Chillon, Peru, 1900 m, 3 May
1958, J. lilies. Same deposition as holotype.

Discussion. — Baetodes traverae can be distinguished from all other
species by the terminal filament which is so small as to be hard to see
from the dorsal view (Fig. 7, ventral view) . The glossae are much
shorter than those of any other species examined (Fig. 12) ; the thumb
on left mandible is more pointed (Fig. 11) ; the spines behind apical
margin of labrum are grouped together as on no other species (Fig. 8).
Thoracic sterna of mature specimens are developed as in subimago.

OCTOBER 1972] MAYO NEOTROPICAL MAYFLIES

231

This species is named in honor of Dr. Jay R. Traver, emeritus profes-
sor of Zoology, University of Massachusetts, Amherst, Massachusetts.

Baetodes solus Mayo, new species

(Figs. 15-19)

Female nymph. — Length of body about 3 mm, cerci somewhat longer. General
color yellowish brown. Head pale brown patches either side midline between eyes;
light brown in epicranial fork; first two segments antennae pale, rest pale smoky
brown. Mouthparts as in Figs. 15-18. One wide bladelike spine distally near inner
margin of each paraglossa (Fig. 15) . Two long bladelike spines among setae on
crowns of maxillae. Thorax with nota of all segments light brown mottled with
yellow. Pleural sclerites brown. Femora light brown except for yellow area near
distal end; narrowly rimmed with reddish brown at joinings with tibiae; tibiae
pale brown. Tarsi pale brown, darker at joinings with claw. Sternum dark brown
mottled with fine yellow dots. Abdomen with terga pale brown, darker along
anterior borders; without tubercles. Abdominal gills as in Fig. 19. Without coxal
gills. Sternum 1 dark brown and mottled as on thorax ; 2-9 brown, somewhat lighter
than thoracic sternum; 10 light brown. Cerci and terminal filament pale, smoky
tinged.

Holotype female nymph , Rio Bella at jct. of Rio Monzon, Monzon
Valley, Huanuco Province, Peru, 25 July 1963, W. L. Peters. In
Entomological collection University of Utah, Salt Lake City.

Discussion. — Baetodes solus has no abdominal tubercles or coxal gills.
See discussions of B. ohesus , B. bellus, and B. traverae for distinguishing
characters between these species and B. solus. B. solus can be separated
from B. sp. No. 1 by the details of the mouthparts, particularly those of
the labrum (Figs. 7, 8, 10, 20, 24, Mayo, 1968).

Baetodes andamagensis Mayo, new species

(Figs. 20-26)

Male nymph. — Length of body 4.5 mm; cerci longer. Head with brown on
either side midline between turbinate eyes; brown in fork; pale spot over small
ocellus. Antennae very pale brown. Mouthparts as in Figs. 21-26. Each para-
glossa with long seta, not pinnate, directed mesally with base below bases of
pinnate spines. Tip of each glossa with two short blunt spines at base of one long
bladelike spine (Fig. 23). Mandibles with two small lobes of incisors at inner
margin; other incisors compacted into single lobe as in Figs. 24, 25; delicate
setae scattered on surfaces as in Figs. 24, 25. Labrum (Fig. 21) with long sclero-
tized strip behind cleft. Maxillae with two wide bladelike spines with blunt tips
among setae on crown (Fig. 22) . Thorax with pronotum dark brown with pale
yellow area at midline along posterior border. Posterior rims of pleural sclerites
visible on either side of pronotum from dorsal view. Intersegmental membrane
between pro- and mesonota pale yellow. Mesonotum dark brown; dark reddish
brown streaks parallel to midline half way between midline and borders of wing
pads; mottled with reddish brown at bases of wing pads; scutellum pale; wing

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OCTOBER 1972] MAYO NEOTROPICAL MAYFLIES

233

pads brown; pleural sclerites, coxae and trochanters brown; femora brown, same
color as thoracic notum; pale yellow spot near joining with trochanter, rimmed
with reddish brown at joining with tibiae; fringe of white setae on tarsi; tibiae
and tarsi very pale brown; tibiae narrowly rimmed with reddish brown at joining
with femora. Sterna pale yellow, except for brown curved strip on prosternum
between apodemes, and dark brown apodemes at leg bases on meso- and meta-
sternum. Abdomen with terga dark brown. Intersegmental membranes yellow.
Tergum light brown anteriorly, dark brown posteriorly along posterior border.
Dark streak on either side midline on terga 2-8, indistinct on 2-5; terga 5-7 darker,
9 an even brown, 10 pale brown. All terga narrowly rimmed with reddish brown
along anterior borders. Without tubercles. Abdominal gills oval, smoky in basal
half (Fig. 20) . Two fingerlike gills from each coxa of the intermediate type. Nu-
merous white setae clustered along midline. Sternum 1 pale yellow; 2-6 brown from
anterior to posterior borders on medial third of segments, laterally yellow, each
lateral area medially with narrow brown streak extending from anterior border half
way to posterior border; 7-9 darker brown, lateral yellow areas mere streaks,
streaked with dark brown along pleural fold; 9 yellow along posterior border; 10
yellow. Cerci and terminal filament very pale brown.

Female. — Head more pale; tibiae and tarsi very pale; cerci and terminal filament
pale yellow.

Holotype male nymph , Andamago River near Ongoro, Peru, 16
May 1958, J. lilies. In Entomological collection of University of Utah,
Salt Lake City. Six paratypes, same data, same deposition as holotype.
Two paratypes, irrigation canal near hacienda, Ongoro, Peru, 16 May
1958. J. lilies. Same deposition as holotype.

Discussion. — On the mandibles there are two small lobes on the in-
cisors at the inner margin. The rest are compacted into a single lobe as
in Figs. 24, 25. This is unique. The labrum can be distinguished from
all others by the spines behind the apical margin (Fig. 21). Maxillae
have two wide bladelike spines with blunt tips among setae on crown

(Fig. 22).

Baetodes obesus Mayo, new species

(Figs. 27-32)

Female nymph.' — Length of body 3 mm, cerci about 4 mm. Stout species. Head
widely yellow around eyes; laterally tips of large ocelli very close to eyes; light
brown either side midline between ocelli, in fork and on genae; pale yellow anterior

<-

Fig. 15. Labium of B. solus. Fig. 16. Maxilla of B. solus. Fig. 17. Left
mandible of B. solus. Fig. 18. Labrum of B. solus. Fig. 19. Gill of B. solus.
Fig. 20. Gill III of B. andamagensis. Fig. 21. Labrum of B. andamagensis. Fig.
22. Maxilla of B. andamagensis. Fig. 23. Tip of glossa of B. andamagensis.
Fig. 24. Left mandible of B. andamagensis. Fig. 25. Right mandible of B. an-
damagensis. Fig. 26. Labium of B. andamagensis.

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OCTOBER 1972] MAYO NEOTROPICAL MAYFLIES

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to small ocellus; anterior margin very narrowly rimmed with dark brown between
antennae and around bases of antennae; antennae yellow, tinged with pale brown
near tips. Clypeus pale, labrum light brown. Mouthparts as in Figs. 27-30, 32.
Labrum not sclerotized below medial cleft. Two long bladelike spines among setae
on crown of maxillae ; very faintly pinnate. Intersegmental membrane between head
and pronotum white, penciled with gray medially; gray area extends from anterior
to posterior borders half way from midline to lateral border. Thorax with anterior
border of pronotum evenly bordered with brown except at midline which is pale;
pronotum yellow with pale area along midline and posterior border near midline;
two yellowish brown stripes from anterior brown border to pale area along posterior
border, about half way between midline and lateral borders; two brown oval spots
on either side near posterior border; narrow reddish brown rims of pleural
sclerites show on either side of pronotum. Mesonotum yellow except for brown
anterior border, light brown side stripe on either side pale midline tapering to half
width anterior to scutellum; mottled with light brown laterally with two darker
brown oval spots, one either side midline anterior to scutellum, and several near
borders of wing pads yellow, with three brown oval spots, one mesally and two ante-
riorly; scutellum and borders of wing pads pale; reddish brown streak on scutum
anteriorly along border of wing pads; brown area either side of scutellum. Meta-
notum dark brown. Legs pale yellow; pleural sclerites light brown. Cervical
sclerites reddish brown. Coxae and trochanters light brown. Femora yellow, darker
in medial third; spines on dorsal crests inconspicuous. Tibiae rimmed with reddish
brown at joinings with femora, otherwise pale; several spines on ventral surface.
Tarsi yellow, washed with reddish brown in distal third at joining with claw;
narrowly rimmed with reddish brown at joining with tibiae; six to eight fairly long
spines along ventral surface of tarsi. Claw with five denticles. Sternum pale
yellow. Abdomen with tergum 1 dark brown; 2 dark brown except for yellow area
mesally on posterior margin and on either side midline % way to anterior border;
tergum 3 the same except brown area more pale and yellow area larger; terga 4^6
yellow, 7-8 light brown, 9-10 yellow; intersegmental areas pale yellow. Without
tubercles. Abdominal gills oval, white (Fig. 31). Two elongated fingerlike gills
from each coxa of the long type. Sterna 1-6 pale yellow tinged with brown along
anterior margin, 7-8 amber tinged with brown anterolaterally, 9 yellow with brown
anterolaterally, 10 yellow. Cerci yellow, stout at bases; terminal filament yellow,
much thinner.

Holotype female nymph, Tenndido River, 3 km N. El Fortin, Vera
Cruz, Mexico, 1 July 1955, R. B. and J. M. Selander. In Entomo-

<r

Fig. 27. Labium of B. obesus. Fig. 28. Maxilla of B. obesus. Fig. 29. Right
mandible of B. obesus. Fig. 30. Left mandible of B. obesus. Fig. 31. Gill of B.
obesus. Fig. 31. Gill of B. obesus. Fig. 32. Labrum of B. obesus. Fig. 33.
Labrum of B. bellus. Fig. 34. Labium of B. fortinensis. Fig. 35. Thoracic
sternum of B. fortinensis. Leg bases indicated by dotted circles. Fig. 36. Labrum
of B. fortinensis. Fig. 37. Left mandible of B. fortinensis. Fig. 38. Maxilla of
B. fortinensis.

236

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

logical collection University of Utah, Salt Lake City. Paratype, 1 female
nymph, same data, same deposition as holotype.

Discussion. — Baetodes obesus can be distinguished from B. solus , B.
traverae , and B. bellus by the coxal gills. The spines on dorsal crests of
femora are fine, inconspicuous, in contrast to the prominent spines of
the other species in this group. The labrum can be distinguished from
that of all other species in this group by the spines behind the apical
margin (Figs. 5, 8, 18, 32, 33, 36, 39) . The two bladelike spines among
setae on crowns of maxillae are faintly pinnate. This is not the case
among the other species in this group.

Baetodes bellus Mayo, new species

(Fig. 33)

Male nymph.— Length of body 4 mm, cerci about 5 mm. Head light reddish
brown on vertex, between turbinate eyes, along midline and epicranial suture;
widely rimmed with yellow around turbinate eyes; pale between large ocelli and
eyes; brown in fork and between antennae; pale around small ocellus; very nar-
rowly rimmed with reddish brown between and around bases of antennae;
antennae pale yellowish brown, darker distally; intersegmental membranes pale;
clypeus and labrum pale. Labrum slightly sclerotized below medial cleft (Fig. 33).
Each glossa with pointed bladelike spine at tip. Two long, bladelike spines among
setae on crowns of maxillae. Intersegmental membrane between head and pronotum
white, penciled with gray across medial third. Thorax with pronotum yellow, nar-
rowly bordered with light brown along anterior margin; three pale brown areas on
either side midline; dark reddish brown margin of pleural sclerites shows on either
side pronotum. Intersegmental membrane between pro- and mesonota white bor-
dered with dark gray posteriorly. Anterior border mesonotum penciled with dark
brown along fairly wide strip; mesonotum yellow faintly mottled with light brown;
midline pale with brown stripes along either side terminating in two brown spots an-
terior to scutellum; another stripe lateral to each stripe; mottled with brown at wing
bases. Metanotum yellow mottled with brown, darker brown along pale midline.
Pleural sclerites light brown. Coxae and trochanters light brown; femora light
brown with pale yellow spot near joining with trochanters; pale elongated spot
below this in medial third, pale spot distally; brown at joinings with tibiae; tibiae
rimmed with reddish brown at joinings with femora; remainder of segment light
brown; few minute spines along ventral surface. Tarsi yellow in apical half and
reddish brown at joinings with claws; darker dorsally; about six minute spines
along ventral surface of tarsi. Claws with five denticles. Pro- and mesosterna pale
yellow; metasternum slightly darker. Abdomen dark brown and yellow; tergum 1
dark brown on anterior two thirds, yellow along posterior border; terga 2-3 yellow
laterally and along posterior borders at midline; remainder dark brown; 4-6 dark
brown along anterior borders, and narrowly touched with brown along posterior
borders at midline; remainder yellow, more pale laterally; 7 with yellow point at
midline on anterior border, yellow mesally and laterally; dark brown along anterior
border half way to lateral border; touched with brown along posterior border

OCTOBER 1972] MAYO NEOTROPICAL MAYFLIES

237

Fig. 39. Labrum of B. veracrusensis. Fig. 40. Left mandible of B. veracrus-
ensis. Fig. 41. Labial palpus of B. veracrusensis. Fig. 42. Maxilla of B.
sancticatarinae. Fig. 43. Labium of B. sancticatarinae. Fig. 44. Gill I of B.
sancticatarinae. Fig. 45. Labrum of B. sancticatarinae. Fig. 46. Left mandible of
B. sancticatarinae. Fig. 47. Thoracic pleuron of B. sancticatarinae. Fig. 48.
Right mandible of B. sancticatarinae. Fig. 49. Mesonotum of B. sancticatarinae.

238

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

mesally; 8-9 with yellow triangle along midline, with apex at anterior border;
dark brown on either side from anterior to posterior border, brown area extending
along anterior border half way to lateral border ; yellow laterally and along posterior
border; 10 yellow. Intersegmental membranes pale yellow streaked with dark gray
along posterior borders. Abdominal gills oval, white. Without coxal gills. With-
out tubercles. Sternum slightly darker than that of thorax. Cerci yellow, terminal
filament more pale.

Holotype male nymph , Tenndido River, 3 km N. El Fortin, Vera
Cruz, Mexico, 1 July 1955, R. B. and J. M. Selander. In Entomological
collection University of Utah, Salt Lake City.

Discussion. — Baetodes bellus can be distinguished from B. obesus ,
B. chilloni , and B. sp. No. 1 (Mayo, 1968) by the absence of coxal gills.
It can be distinguished from B. solus by the pale yellow sternum; that of
B. solus is dark brown.

Baetodes fortinensis Mayo, new species

(Figs. 34-38)

Male nymph. — Length of body 3 mm; cerci longer. Head with turbinate eyes
rust color, black around bases, very close to midline; pale rim around area of future
top in subimago. Lower eyes black. Large ocelli touching eyes; light brown
around ocelli and on genae; darker in epicranial fork; pale yellow anterior to small
ocellus, narrowly bordered with dark brown between antennae and around bases;
deeply notched at bases of antennae; antennae pale brown, darker at tips; inter-
segmental areas pale. Clypeus pale with black articulations on either side;
labrum light brown. Mouthparts as in Figs. 34, 36-38. Labial and maxillary palpi
with numerous setae. Each glossa with bladelike spine at tip; bladelike spine with-
out pinnae on each paraglossa at medial side of tip; first segment of labial palpus
shorter than other two combined. No sclerotization below medial cleft on labrum.
Thorax with pronotum yellowish brown; anterior border rimmed with dark brown
except at midline; reddish brown area on either side midline; yellowish brown
laterally; posterior border slightly elevated at midline. Mesonotum yellowish brown
with reddish brown pattern; midline pale; anterior border dark brown; reddish
brown stripe either side midline terminating about midway to scutellum; dark dot
either side midline half way between this stripe and scutellum; wider stripe lateral
to this terminating near dot; laterally mottled with reddish brown; wing pads yel-
lowish brown; scutellum pale yellow narrowly bordered with dark brown. Meta-
notum dark brown, somewhat mottled. Pleural sclerites brown; coxae and tro-
chanters yellowish brown; femora yellow, narrowly bordered with reddish brown at
joinings with tibiae; ten to twelve spines along dorsal crests small. Tibiae yellow;
tarsi light brown dorsally, darker at joining with claw; small spines on ventral
surface of tibiae and tarsi. Claw with six denticles. Sternum pale yellow with dark
brown sclerites on furcisternum as in subimago (Fig. 35) ; sclerites around apodemes
at leg bases dark brown. Abdomen with terga 1 and 7-8 dark brown, mottled;
terga 2-6 and 9 yellowish brown; 10 lighter, more yellow. Without tubercles.
Abdominal gills oval, white. Two white, elongated fingerlike gills from each coxa
of the long type. Sterna 2-6 and 9-10 yellow, 7-8 brown. Cerci yellow at bases,
yellowish brown distally; terminal filament pale yellow.

OCTOBER 1972]

MAYO NEOTROPICAL MAYFLIES

239

Holotype mature male nymph , Tenndido River, 3 km N. El Fortin,
Vera Cruz, Mexico, 1 July 1955, R. B. and J. M. Selander. In Entomo-
logical collection University of Utah, Salt Lake City.

Discussion. — The mouthparts of Baetodes fortinensis differ from
those of the others in group without abdominal tubercles. The maxillary
and labial palpi of this species have numerous setae (Figs. 34, 38) . The
first segment of labial palpus is shorter than the other two combined.
The spines behind the apical margin of labrum differ. There is no sclero-
tization below the medial cleft of labrum (Fig. 36) .

Baetodes veracrusensis Mayo, new species

(Figs. 39-41)

Male nymph. — Length of body 4 mm; cerci about 5 mm. Head narrowly
mottled with light brown between eyes on either side midline; turbinate eyes
rimmed medially with brown; brown on either side of epicranial suture, anterior to
large ocelli and on genae anterior to compound eyes; yellow around small ocellus;
narrowly rimmed with reddish brown between and around antennae; antennae and
clypeus pale yellow; labrum light brown bordered with darker brown. Bladelike
spine on each glossa pointed. Two long bladelike spines among setae on crowns of
maxillae. Labrum very slightly sclerotized below medial cleft. Mouthparts as in
Figs. 39-41. Intersegmental membrane between head and pronotum penciled with
gray near anterior border of pronotum. Thorax with anterior border of pronotum
light brown; pronotum yellow mottled with pale brown on either side of midline.
Mesonotum reddish brown along anterior border; rest of mesonotum yellow mottled
with darker brown than that on pronotum; two pair of brown stripes on either
side yellow midline converge anterior to pale scutellum; mottled with brown spots
laterally; wing pads yellow. Metanotum yellow; pale brown anteriorly. Pleural
sclerites pale yellowish brown. Coxae pale brown, trochanters yellow with brown
articulations with femora; femora light brown except pale yellow at joining with
trochanters; large elongated yellow spot medially and yellow near tibiae; fringe
of white setae on tarsi. Tibiae yellow rimmed with reddish brown at joinings with
femora and tarsi. Tarsi yellow, brown dorsally at joinings with claws; six small
spines along ventral margin. Claw with seven denticles. Sternum pale yellow.
Abdomen with tergurn 1 dark brown anteriorly and yellow posteriorly; irregular
margin of dark brown and yellow between anterior and posterior borders; 1-10
yellow posterolaterally and along pleural fold near attachment of gills; 2-3 brown
with yellow medially along posterior borders; 4^6 bordered with brown anteriorly
with narrow strip of brown along posterior border on either side midline; 7 like
6 with wider brown area anteriorly; 8-9 with brown triangles based along anterior
border and touching at midline; rest of segments yellow; 10 yellow. Dark gray
trachea visible laterally near pleural fold on 6-9. Abdominal gills oval, white. One
small white fingerlike gill from each coxa, of intermediate type. Sterna 2-9 light
yellowish brown, progressively more pale rearward; 10 yellow. Cerci and terminal
filament pale yellow.

240

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Holotype male nymph , Tenndido River, 3 km N. El Fortin, Vera
Cruz, Mexico, 1 July 1955, R. B. and J. M. Selander. In Entomological
collection University of Utah, Salt Lake City.

Discussion. — Baetodes veracrusensis can be separated from all other
species without abdominal tubercles by the presence of small coxal gills
of the intermediate type. The spines behind apical margin of labrum
differ from those of the other species.

Baetodes sancticatarinae Mayo, new species

(Figs. 42-49)

Mature female nymph. — Length of body 4 mm. Cerci about the same. Head
with prominent ocelli; large ocelli touching eyes. Reddish brown on either side
pale midline and in fork. Pale yellow around small ocellus and elsewhere on head;
narrowly rimmed with reddish brown between antennae; antennae pale brown.
Mouthparts as in Figs. 42, 43, 45, 46, 48. Two pointed bladelike spines among setae
on crowns of maxillae. Pinnate spine resembling a brush on each glossa. A long
simple spine not pinnate on each paraglossa arising below pinnate spines, mesally.
Thorax with pronotum light brown with large yellow spots; rims of pleural sclerites
visible on either side of pronotum. Mesonotum dark reddish brown with amber;
pattern as of subimago ; reddish brown stripe on either side of pale area along
midline terminates in dark elongated dot (Fig. 49). Scutellum very pale yellow;
metanotum brown; scattered setae on dorsal surface of thorax. Pleural sclerites
amber with dark reddish brown vertical line between anterior margins and coxal
articulations. The elongated dark reddish brown streak anterior to pleural sclerite
III is typical of all mature specimens (Fig. 47). Coxae and trochanters amber;
unsclerotized areas pale yellow; femora yellow except brown at joinings with
trochanters and rimmed with reddish brown at joinings with tibiae; washed faintly
with light brown in medial third; fringe of white setae on coxae; tibiae yellow;
tarsi yellow with reddish brown dorsally at joinings with claws; claw with eight
denticles; minute spines along ventral surfaces of tibiae and tarsi. Sternum typical
of mature nymph with sclerites as of subimago, sclerites of furcisternum amber in
contrast to the narrow dark brown sclerites of basisternum on segment II. Pro-
sternum pale yellow anterior to apodemes; pale brown posterior to these. Meso-
and metasternum pale brown mottled with very fine yellow dots giving slightly
roughened appearance. Unsclerotized areas around leg bases pale yellow. Abdomen
without tubercles. Setae on abdomen clustered near midline; many setae encrusted
with silt. Terga 1-2 brown, dark brown along anterior borders; 3-8 dark reddish
brown along anterior borders, remainder of segments amber; 9 more pale, 10 yellow
touched with brown along anterior and posterior borders. Abdominal gills as in
Fig. 44; tracheae visible over entire surface; gill I large, others decrease in size
rearward; white. One fingerlike gill from each coxa of the long type. Sternum 1
pale brown mottled with very fine yellow dots; 2-6 light brown; 7-9 with brown
areas decreasing in size rearward with yellow anteriorly and posteriorly. Cerci
amber; with minute reddish brown spines on segments in basal third; terminal
filament yellow.

Mature male nymph. — Length of body 3 mm. Cuticula of head and thorax split
along midline showing subimago with fairly wide pale yellow strip along midline,
pale scutellum narrowly rimmed with reddish brown. On either side of midline is

OCTOBER 1972]

MAYO NEOTROPICAL MAYFLIES

241

a reddish brown strip terminating in two prominent reddish brown elongated dots
anterior to scutellum with narrow white line separating brown strip laterally from
rest of brown scutum. Terga brown, except 4-6 amber medially; 7-8 dark brown,

9 more pale, 10 yellow touched with brown along anterior and posterior margins as
in female. Cerci amber with brown spines as on mature female; terminal filament
yellow.

Holotype female nymph, River Pirabeiraba, 26' 15 °B. 48' 54°L. 10
m, Santa Catarina State, Brazil, November 1965, F. Plaumann. In
Entomological collection University of Utah, Salt Lake City. Five para-
types same data, same deposition as holotype.

Discussion. — The pattern of spines behind apical margin of labrum
differs from all others in the group without abdominal tubercles. More
trachea show on gills than on gills of all other species. Spines on basal
third of cerci on mature specimens are unique.

Immature specimens: don’t have spines on cerci (these occur only on
mature specimens) ; none of their terminal filaments have spines; their
thoracic sterna and abdominal sternum 1 are pale yellow; 2-5 pale
brown, remainder increasingly more yellow rearward; they have a
brown streak laterally on 6-10; their cerci are pale yellow basally, light
brown distally; with the terminal filament yellow.

Acknowledgments

The writer wishes to acknowledge the kindness of Dr. George F. Ed-
munds Jr., chairman of the Department of Environmental Biology,
University of Utah for the loan of his collection of Baetodes. She is also
indebted to Dr. Jay R. Traver, emeritus professor of Zoology, University
of Massachusetts, for her help in going over the manuscript.

Literature Cited

Cohen, S. D. and R. K. Allen. 1972. New species of Baetodes from Mexico and
Central America (Epbemeroptera: Baetidae) . Pan-Pac. Entomol., 48:
123-135.

Edmunds, G. F., Jr. 1950. New records of the mayfly Baetodes, with notes on the
genus. Entomol. News, 61: 203-205.

Macan, T. T. 1961. A key to the nymphs of the British species of Ephemeroptera.

Freshwater Biol. Assoc., Sci. Publ. 20: 1-63, 37 fig.

Mayo, V. K. 1968. Two new species of the genus Baetodes from Ecuador. Pan-
Pac. Entomol. 44: 251-257.

Needham, J. G. and H. Murphy. 1924. Neotropical mayflies. Bull. Lloyd Li-
brary, 24, Entomol. Ser., 4: 55-56.

Traver, J. R. 1943. New Venezuelan mayflies. Entomol. Venezolena, 2(2): 79-
98.

1944. Notes on Brazilian mayflies Bol. Mus. Nac. Rio de Janeiro Zool., 22-20.

Traver, J. R. and G. F. Edmunds, Jr. 1968. A revision of the Baetidae with
spatulate-clawed nymphs. Pac. Insects, 10: 629-677.

242

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

A New Species of Rhagonycha from Alaska 1

(Coleoptera: Cantharidae)

Kenneth M. Fender

Linfield Research Institute, McMinnville, Oregon 97128

The following species had been tentatively placed with Rhagonycha
mandibularis (Kirby), its nearest relative. Additional material and the
redescription of Kirby’s species by Green (1940) proved this to be
misplaced. An undescribed species was at hand.

Fall (1926) resurrected Cantharis mandibularis Kirby when he com-
pared specimens with one of Kirby’s cotypes. Prior to this it had been
placed as a synonym of Cantharis fraxini Say.

In 1971, I assigned to the genus Rhagonycha Eschscholtz, that section
of Cantharis of North America in which the species had the third tarsal
segment simple and the insertion of the fourth segment apical. Rhagon-
ycha has been recognized as a valid genus by European students for
years. Both Green and McKey-Fender (1950) had suggested such a
separation but neither followed through on it.

Rhagonycha alaskensis Fender, new species

(Figs. 1-4)

Dark brown, sides of pronotum obscurely paler; head black, antennae and
maxillary palpi piceous, palpi a little paler; mandibles and trochanters of prolegs
ferruginotestaceous. Pubescence cinereous, suberect, fine, sparse and inconspicuous.

Male. — Eyes moderately large and prominent, separated by about one and one-
third combined widths as viewed from above. Head as wide as pronotum, finely
sparsely punctured, shining in front of antennae, finely alutaceous behind; clypeal
apex oblique each side of median notch ; antennae slender, filiform extending to about
middle of elytra, third segment nearly twice as long as second, intermediate seg-
ments nearly four times as long as wide. Pronotum sub quadrate, slightly wider than
long; anterior margin evenly rounded and shallowly reflexed, anterior angles evenly
rounded into nearly straight and almost parallel lateral margins; basal angles
sharply rounded; basal margin feebly arcuate, strongly reflexed medially; sides
shallowly reflexed at hind angles, becoming feebly explanate towards anterior
angles; sub-basal convexities not prominent; surface finely sparsely punctured,
feebly shining. Combined elytra wider than pronotum, about three and one-third
times as long as width at humeri, coarsely sparsely punctured basally, rugose punc-
tate beyond basal fifth, two costae feebly elevated and feebly indicated in each.
Metasternum shining, finely sparsely punctured. Abdominal sternites dull, finely
sparsely punctured, finely transversely strigulose towards sides. Tarsal claws nar-
rowly cleft at apices. Aedeagus with apical margin of dorsal plate narrowly deeply

1 This study was supported by National Science Foundation Grant: GB-6283X.

The Pan-Pacific Entomologist 48: 242-243. October 1972

OCTOBER 1972]

FENDER NEW RHAGONYCHA

243

Figs. 1-4. Rhagonycha, alaskensis. Fig. 1. Protarsal claw of male. Fig. 2. Male
aedeagus, dorsal view. Fig. 3. Same, lateral view. Fig. 4. Same, ventral view.

incised. All tarsi with third segment simple, insertion of fourth segment apical.
Length 5.5 mm.

Female. — Unknown.

Holotype male , Matanuska, Alaska, 26 June 1914, rotary trap, J. C.
Chamberlain. In the collection of the California Academy of Sciences.

This species is nearest to Rhagonycha mandibularis (Kirby) . But the
pronotum of R. mandibularis is more transverse, being three-tenths
wider than long and narrowed in front to the obsolete anterior angles.
The tarsal claws of R. mandibularis are unusually small, and the dorsal
plate of the aedeagus narrows from base to apex with the apical margin
more broadly and less deeply incised.

Literature Cited

Fall, H. C. 1926. A list of the Coleoptera taken in Alaska and adjacent parts of
the Yukon Territory in the summer of 1924. Pan-Pac. Entomol., 2:
127-154.

Fender, K. M. 1971. The genus Rhagonycha Eschscholtz in North America
(Coleoptera: Cantharidae) . Coleopt. Bull., 25: 86-87.

Green, J. W. 1940. Taxonomic studies in Cantharis (Coleoptera: Cantharidae).
Entomol. Amer., 20: 159-217.

McKey-Fender, D. 1950. Notes on Cantharis III (Coleoptera: Cantharidae).
Pan-Pac. Entomol., 26: 25-33; 26(2): 61-79.

Correction

Forister, G. W. and C. D. Johnson. 1971. Behavior and ecology of Acantho-
scelides prosopoides (Coleoptera: Bruchidae) . Pan-Pac. Entomol.,
47(3) : 224-234.

On page 226, line 16 should be inserted between lines 12 and 13. — C. D. Johnson,
Northern Arizona University, Flagstaff, 96001.

244

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

A New Genus of Biting Midge from California
Related to Neurohelea Kieffer

(Diptera, Ceratopogonidae)

Willis W. Wirth and Niphan Chanthawanich Ratanaworabhan 1

Systematic Entomology Laboratory, V. S. D. A., cfo U. S. National Museum,
Washington, D. C. 20560, and Applied Scientific Research Corporation,

Bangkok, Thailand

Wirth (1952) described Bezzia granulosa from Monterey County,
California, pointing out that in several characters it did not fit the genus
Bezzia Kieffer, stating: “The anterior radial cell is very long, the radius
produced, and the fifth tarsal segment swollen as in Neurohelea ; the hind
tarsal claws are long and unequal, and the abdomen is marked much as
in Probezzia While presently undertaking a revisionary study of the
genera of Ceratopogonidae, we have studied additional material of B.
granulosa which indicates that this species must be placed in a new genus
of Heteromyiini (Wirth, 1962) near Neurohelea Kieffer. The wing of
Neurohelea has two radial cells, the female tarsal claws are equal on all
legs, and the female abdomen is uniformly dark.

Neurobezzia Wirth and Ratanaworabhan, new genus

Type-species, Bezzia granulosa Wirth.

Diagnosis. — Female: Small black midges with whitish wing and bicolored legs
(Fig. lg) and abdomen. Very similar to Neurohelea; differing as follows: Wing
(Fig. lb) with one radial cell. Legs (Fig. lh) with tarsal claws long and curved;
equal on fore and mid legs, unequal on hind legs, shorter claw approximately
three fourths as long as longer one; claws (Fig. le) each with small, slender,
internal barb. Abdomen milky white, third and fourth segments with variable
internal pigmentation; eighth segment (Fig. li) sclerotized, forming dark brown
ring.

Neurobezzia granulosa (Wirth), NEW COMBINATION

(Fig. 1)

Bezzia granulosa Wirth, 1952: 240 (female; California).

Type. — Holotype, female, Arroyo Seco Ranger Station, Monterey County, Cali-
fornia, 1 July 1948, W. W. Wirth (Type no. 59964, USNM) .

Distribution. — California.

New Records . — CALIFORNIA: Mono Co., Convict Creek, 7,200 ft, 17 July 1968,
H. D. Kennedy, light trap, 1 female; Sardine Creek, 8,500 ft, 6 July 1951, A. T.
McClay, 1 female. Tehama Co., Red Bluff, 11 May 1949, H. P. Chandler, light trap,
1 female. Trinity Co., Trinity River Camp, 17 July 1953, A. T. McClay, 2 females.

1 Acknowledgment is gratefully made to the Southeast Asia Treaty Organization in Bangkok for finan-
cial assistance for study at the U. S. National Museum.

The Pan-Pacific Entomologist 48: 244-245. October 1972

OCTOBER 1972] WIRTH & RATANAWORABHAN NEW BITING MIDGE 245

Fig. 1 . Neurobezzia granulosa, female: a. antenna; b. wing; c. mandible;
d. palpus; e, fifth tarsomere and claws of fore leg; f. spermathecae ; g. femora
and tibiae of fore, mid and hind legs, left to right; h. same, tarsi; i. genital
sclerotization of terminal abdominal segments.

Literature Cited

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

1962. A reclassification of the Palpomyia-Bezzia-Macropeza Groups, and a
revision of the North American Sphaeromiini (Diptera, Ceratopogon-
idae) . Ann. Entomol. Soc. Amer., 55: 272-287.

246

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Notes and Synonymy in Largus Hahn with a Key to

United States Species

(Hemiptera: Largidae)

Thomas F. Halstead

Arizona Commission of Agriculture and Horticulture
Phoenix, 85005

In 1970 I described a new species of Largus Hahn, and provided a key
to southwestern species of the genus. This material was based on data
available to me prior to 1968 and ignored several taxa described by
Bliven (1956 and 1959) and a subspecies named by Van Duzee in 1923.
Since that time I have acquired additional information relating to the
genus and have described a new species from Florida (Halstead, 1972).
This paper will present comments on distribution and synonymy for
species encountered in the United States, as well as a key for their dif-
ferentiation.

Largus Hahn of the United States

1. Rectangular hemelytral patch, pronotal posterior and abdominal venter

white sellatus (Guerin)

Rectangular white hemelytral patch absent, pronotal posterior and abdom-

inal venter not white 2

2. Membrane white or very light tan, veins concolorous or light tan 3

Membrane tan to black, veins brown to black 5

3. Two very distinct transverse black maculations at juncture of claval suture

and membrane maculatus Schmidt

Without two distinct black maculations at juncture of claval suture and
membrane 4

4. Ground color distinctly yellow; abdominal venter at least basally and usually

entirely yellow pallidus Halstead

Ground color red; venter black, very rarely with slight red suffusion of
basal abdominal segments davisi Barber

5. Thoracic sterna around base of coxae red to orange 6

Thoracic sterna around base of coxae black 7

6. Coxae often red; trochanters and femora black convivus Stal

Coxae, trochanters and proximal ends of femora all red to yellow-orange
cinctus Herrich-Schaffer

7. Trochanters and proximal ends of femora orange to orange-red

succinctus Linnaeus

Trochanters and femora all black semipunctatus Halstead

A number of species of Largus show great variation in expression of a
given color trait. The same portion of the anatomy of individuals of the

The Pan-Pacific Entomologist 48: 246-248. October 1972

OCTOBER 1972 ] HALSTEAD — LARGUS NOTES AND KEY

247

same species may vary from pale yellow or tan to deep red. It is there-
fore of questionable value to base taxa on color shade. It is a completely
different matter to base taxa on patterns of coloration. The above key is
based entirely on color expression, and is simply a guide to identification
of specimens. The various species represented differ with respect to a
number of morphological characters as well.

Largus cinctus.- — W. North America from British Columbia to south-
ern Mex., e. to central Tex. Largus cinctus californicus (Van Duzee),
1923 is a synonym based on characters of pubescence and coloration
which break down when compared with a large series of specimens.
Unpublished thesis data (Halstead, 1967) involving comparative analysis
of specimens from seven populations of Largus cinctus ranging from
Washington to southern Mexico show that while a tendency for speci-
mens from the south to be paler and less pubescent does exist, some
specimens from any population will match Van Duzee’s description.
L. cinctus californicus was designated as a geographic variation from
northern areas in California, as different from specimens originating in
Arizona and Mexico. Bliven (1959) proposed Largus semipletus from
northern California citing difference in relative antennal segment length
as the major separatory character. Comparison of Bliven’s antennal
segment measurements with measurements of specimens from four Cali-
fornia counties (Halstead, 1967) shows that data cited by Bliven fall well
within variation shown by L. cinctus populations in California. The
other major character cited by Bliven was pubescence. The description
of this characteristic is not sufficient to separate the new taxon from L.
cinctus. Careful analysis of the long description given by Bliven has
failed to show me any other criteria than the two mentioned above which
could possibly justify proposal of a new species. It is my opinion that
L. semipletus is a synonym of L. cinctus. Largus sculptilis Bliven, 1959
was separated from other Largus species by reason of differing colora-
tion, pubescence, punctation and antennal segment length. The range of
variation with respect to all of these characters shown by L. cinctus
populations is sufficient to encompass Bliven’s new species. Comparative
analysis shows that antennal segment length values given by Bliven fall
within the range for L. cinctus populations (Halstead, 1967). Largus
sculptilis Bliven is synonymous with L. cinctus. I have examined several
thousand specimens of Largus from California localities; to date all
have been L. cinctus. This includes material determined by Van Duzee
and others as L. convivus. I have examined Torre-Bueno’s “paraplesio-
types” (specimens bearing that label) through the courtesy of Dr. G. W.
Byers of the Snow Entomological Museum of the University of Kansas

248

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

and find specimens of both L. cinctus and L. convivus bearing L. cinctus
labels.

Largus convivus. — Mountainous regions of Ariz. w. to Tex., s. to
central Mex. Wupatkius semo Bliven, 1956 was proposed as a new
Largid genus and species from northern Arizona. Bliven’s description
of Wupatkius shows absolutely no variation from Hahn’s original de-
scription of Largus in 1831. Wupatkius must therefore fall to Largus.
Furthermore, Bliven’s description of W. semo is very clearly a descrip-
tion of L. convivus. Wupatkius semo Bliven must therefore fall in synon-
ymy to L. convivus.

Largus succinctus. — Fla. n. to N.Y., w. to Tex. The common Eastern
species corresponding to L. cinctus in the West.

Largus semipunctatus. — S. Ariz., s.e. N.M., with one doubtful Cali-
fornia record. A rare mountain species.

Largus maculatus. — Brownsville, Tex. and perhaps s. Ariz. s. to Co-
lombia. This is the Largus bipustulatus of my 1970 key. Examination
of Stal’s type through the courtesy of Dr. Per Inge Persson of the
Stockholm Museum proved to me that this apparently rare Mexican
species differed from the specimens from the Brownsville area.

Largus sellatus. — Extreme s. Fla. Differs from all other Largus of the
United States by the distinct white markings, as noted in the key. A
Caribbean species.

Largus davisi.- — Fla. only, not uncommon.

Largus pallidus.— Key Largo, Fla. and adjacent areas. Generally
larger and more robust than L. davisi.

Literature Cited

Bliven, B. P. 1956. New Hemiptera from the western states with illustrations
of previously described species and new synonymy in the Psyllidae.
Occidental Entomol., Now. 1956: 1-27.

1959. New Pyrrhocoridae and Miridae from the western United States. Occi-
dental Entomol., 1(3) : 25-33.

Hahn, C. W. and Herrich-Schaeffer, G. A. W. 1831-1853. Die wanzenartigen
insecten. Nurnberg. Nine volumes.

Halsted, T. F. 1967. A taxonomic investigation of the various specific com-
ponents of the genus Largus Hahn, 1831 of western North America.
M.A. Thesis. San Francisco State College. 1-69.

1970. A new species of the genus Largus Hahn with a key to the species of the
genus in the southwestern United States. Pan-Pac. Entomol. 46(1) :
45-46.

1972. A new species of Largus. Can. Entomol., 104(6) : 959.

Van Duzee, E. P. 1923. A new subspecies of Euryophthalmus cinctus (Hemiptera).
Can. Entomol., 55(11) : 270.

OCTOBER 1972] MOORE & LEGNER NEW CALIFORNIA ORUS

249

Two New Species of Orus from California

(Coleoptera: Staphylinidae)

Ian Moore and E. F. Legner 1
Division of Biological Control, University of California, Riverside, 92502

The genus Orus was completely revised by Herman (1964, 1965). In
his revision he gave detailed drawings of the male secondary sexual
characters and of the aedeagus of each species making identifications of
male specimens positive. Two new species not represented in the mate-
rial which Herman examined have come to hand from areas in Cali-
fornia. These are described and illustrated below. Both belong to the
subgenus Orus {sensu stricto ) .

Key to the Males of Orus (5. str .)

1 .

2 .

3.

4.

5.

6 .

7.

8 .

9.

Last visible abdominal sternite shallowly incised 2

Last visible abdominal sternite deeply incised 3

Fifth visible abdominal sternite emarginate; neck one-fifth the width of

head montanus Fall

Fifth visible abdominal sternite sinuotruncate ; neck one-fourth the width

of head shastanus Casey

Fifth visible abdominal sternite sinuate along the posterior margin

sinuatus Herman

Fifth visible abdominal sternite with the posterior margin lobed or emar-
ginate 4

Fifth visible abdominal sternite emarginate 5

Fifth visible abdominal sternite lobed 7

Fifth visible abdominal sternite deeply emarginate (Fig. 1)

giulianii Moore and Legner, n. sp.

Fifth visible abdominal sternite shallowly emarginate 6

Fifth visible abdominal sternite deeply impressed (Fig. 2)

frommeri Moore and Legner, n. sp.

Fifth visible abdominal sternite very shallowly impressed fraternus Fall

Fifth visible abdominal sternite with a tubercle 9

Fifth visible abdominal sternite without a tubercle 8

Paramere short, not extending beyond apex of median lobe, broad, with

lateral margins straight in dorsal aspect hemilobatus Herman

Paramere long, extending beyond apex of median lobe, slender, with lateral

margins sinuate in dorsal aspect punctatus Casey

Abdominal tubercle well developed, margined laterally by carina; meta-
femora carinate femoratus Fall

Abdominal tubercle feeble, not margined by carina; metafemora not
carinate _ ___ distinctus Casey

1 Staff Research Associate and Associate Professor of Biological Control, University of California,
Riverside, respectively.

The Pan-Pacific Entomologist 48: 249-252. October 1972

250

THE PAN-PACIFIC ENTOMOLOGIST

[VOL. 48, NO. 4

Orus giulianii Moore and Legner, new species

Holotype male. — Piceous with tarsi and mouth parts just perceptibly paler.
Head one-fourth longer than wide; above finely, densely punctured and finely,
densely microreticulate, beneath sculptured much as above except smooth and not
impressed gula. Pronotum one-tenth longer than head, one-fifth longer than wide,
surface sculptured much as head except for vague smooth central longitudinal area.
Elytra one-fourth longer than pronotum, conjointly nine- tenths as wide as long,
surface sculpture much like that of pronotum. Abdomen more finely punctured
than elytra, finely microreticulate; first four visible sternites unmodified; fifth
visible sternite with the posterior margin deeply emarginate, bottom of emargina-
tion produced as small cusp, sides of emargination slightly convergent posteriorly;
sixth sternite with posterior margin moderately deeply incised.

Holotype male, Oak Creek, Inyo County, California, 4,100 feet
elevation ultraviolet black light, June 1971, Derham Giuliani collector
[California Academy of Sciences]. Paratypes, same data as holotype
(1 male) ; Big Pine, Inyo County, California, ultraviolet black light,
July 1970 (4 males) ; March 1971 (1 male), Derham Giuliani collector
[California Academy of Science and University of California, River-
side]. We have also seen twenty -two females with the above data which
are not designated paratypes.

The modifications of the fifth and sixth sternites of this species are
similar to those of 0. ferrugineus and O. guatemalenus, both of which
belong to the sub genus Leucorus having either a bidentate or edentate

OCTOBER 1972] MOORE & LEGNER NEW CALIFORNIA ORUS

251

labrum. The labrum of 0. giulianii is distinctly and strongly quadri-
dentate and the sixth sternite is more shallowly and broadly incised than
those of the two species mentioned above.

Orus frommeri Moore and Legner, new species

Holotype male. — Castaneous with head and bases of elytra darker, legs testa-
ceous, beneath testaceous. Head one-fifth longer than wide; finely somewhat
densely punctured above with very feeble microreticulation; beneath sculptured
much as above except highly polished and not impressed gula. Pronotum one-tenth
longer than head, seven-tenths as wide as long; sculpture very similar to that of
head. Elytra one- fifth longer than pronotum, conjointly four-fifths as wide as long;
surface densely but somewhat more coarsely punctured than head and pronotum,
with more pronounced ground sculpture. Abdomen very finely punctured with fine
but dense microreticulation; with first four visible sternites unmodified; fifth
visible sternite with posterior edge emarginate, bottom of emargination shallowly
sinuate, surface of sternite deeply, broadly impressed for its entire length before
emargination; sixth visible sternite moderately incised, shallowly impressed in
middle.

Holotype male, vicinity of Santa Ynez River, Santa Barbara
County, California, 1,000 feet elevation ultraviolet and white light, 13
June 1971, Saul Frommer collector [California Academy of Sciences].
Paratype male, same data as holotype (1 male) [University of California,
Riverside].

This species is most similar to 0. sinuatus in the secondary male sex-
ual characters. It differs from that species by the more deeply emargi-

252

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

nate posterior margin of the fifth visible sternite and the more shallowly
incised posterior margin of the sixth sternite and particularly in the
wide, deep impression on the surface of the fifth sternite.

Literature Cited

Herman, Lee H., Jr. 1964. A revision of Orus Casey. I. Subgenus Leucorus
Casey and a new subgenus (Coleoptera: Staphylinidae) . Coleopt. Bull.,
18: 112-121.

1965. Revision of Orus. II. Subgenera Orus, Pycnorus and Nivorus (Cole-
optera: Staphylinidae). Coleopt. Bull., 19: 73-90.

BOOK REVIEW

The Lives of Wasps and Bees. By Sir Christopher Andrews. American Elsevier

Publishing Company, Inc., New York. 204 pages, illus. 1970. $5.75.

As a popular account of the life histories of aculeate Hymenoptera attempting
to “show that among wasps and bees in particular there are many with fascinating
and often bizarre habits” the book is generally successful. The book consists of 27
short, easy to read chapters, 16 plates of excellent photos, and 15 additional well
drawn figures. For the systematically oriented, scientific names of species men-
tioned along with phonetic guides to their pronunciation appear at the end of each
chapter, and the higher classification (superfamilies to genera) of the taxa treated
follows the last chapter. The numbers of chapters devoted to wasps versus bees or
to those with solitary versus social habits balance nicely. The author limits cover-
age principally to species of Europe and North America and to some of the more
general literature principally in English from 1905-1967.

Errors consist of several types: Typographical errors (e.g. the first initial for
K. V. Krombein on page 44; Bembix captures flies resting, not “nesting on vege-
tation” on page 55; etc.) ; Misleading statements (e.g. from the discussion on page
28 one expects to see a grasshopper, not a caterpillar in plate 2A; the title “Some
Wood-borers” for Chapter 6 which deals with species of Trypoxylon which use
pre-existing cavities or construct pipe-organ mud nests) ; and Errors of omission
(e.g. works of many European and American authors including Friese, von Frisch,
Grandi, Malyshev, Linsley & MacSwain, Plath; discussion of massarid wasps).
The errors of omission disturb only specialists and other errors are easily overlooked
by the general reader.

This popularized treatment does expose some of the marvels of the insect world
to public view. As such it provides entertaining and informative reading for those
whose knowledge of bees and wasps does not extend much beyond the habits of
honey bees and yellow jackets. — R. W. Ti-iorp, University of California, Davis,
95616.

OCTOBER 1972] PINTO CAVICEPS GROUP OF EPICAUTA

253

Notes on the Caviceps Group of the Genus Epicautci
with Descriptions of First Instar Larvae 1

(Coleoptera: Meloidae)

John D. Pinto

Department of Entomology, University of California, Riverside, 92502

Fifteen species of North American Epicauta have been placed in the
Caviceps Group (Werner, 1955; Werner, Enns and Parker, 1966). Those
included are E. aspera Werner, E. wheeleri Horn, E. rehni Maydell, E.
occipitalis Werner, E. singularis Champion, E. diver sipubescens Maydell,
E. cicatrix Werner, E. excavatifrons Maydell, E. straba Horn, E. afoveata
Werner, E. impressifrons Van Dyke, E. caviceps Horn, E. rileyi Horn,
E. insueta Werner, and E. stuarti LeConte. Adults of this group are
primarily active in autumn. Records for several species suggest that
adults feed solely on the inflorescences of fall-blooming Compositae.
Geographically, the group is almost totally confined to southwestern
North America. Exceptions are E. excavatifrons from southeastern
United States, and E. insueta from southern Mexico.

The purpose of this paper is to briefly review the adult and larval
anatomy of the Caviceps Group and to alter its limits slightly by includ-
ing two additional species. Also, the first instar larvae of three species
are described and compared with those previously studied by MacSwain
(1956). New food-plant records for adults are included.

Adults

The Caviceps Group has been defined primarily by adult anatomy
(Werner, 1955). It has contained all North American species having
males with the ventral and posterior surfaces of the meso- and meta-
femora, and the hind two pair of trochanters denuded, the two denuded
femoral areas being separated by a fringe of long hairs. Long, tapering
antennae is the only other important characteristic present in all mem-
bers of the group. Several other traits which are absent in some of the
species, however, do help delimit this taxon. These include various head
and eye modifications, black spots on the abdomen and elytral base, a
whorled setal pattern on the pronotum, and an elevation of the elytral
suture near its basal third.

Based on both adult and larval anatomy E. californica Werner and E.
alphonsii Horn, two cognate species from southern California, should, in

1 This study was supported in part by Giant GB 30907 from the National Science Foundation.

The Pan-Pacific Entomologist 48: 253-260. October 1972

254

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

my opinion, be included in the Caviceps Group. Although mentioned as
being similar to species of this group, reasons for their exclusion were
not given by Werner (1955). These two species lack any indication of
the head modifications which characterize several species of the Caviceps
Group. However, their leg modifications are pronounced. The ventral
surface of the meso- and metafemora in males of both species is not only
denuded but also distinctly widened and flattened, and this surface is
noticeably concave from base to apex. In addition, the ventral surface
of the mesofemur is fringed with long hairs anteriorly as well as poste-
riorly. These differences merely represent specialized states of morpho-
clines already found in the Caviceps Group sensu Werner. Thus, in
some species of the group (e.g. E. caviceps and E. rileyi ) the ventral
surface of the hind two pair of femora is simply denuded but in others
it is also widened (e.g. E. wheeled) , and in E. afoveata slightly concave
as well. The anterior fringe of long hairs on the mesofemur, although
absent in most members of the group, does occur in E. aspera.

Associated with these leg modifications the metasternum of E. califor-
nica and E. alphonsii is denuded and distinctly concave rather than
hirsute and convex as in most meloids. This trait also occurs to a lesser
degree in E. wheeled, E. impressifrons, and E. afoveata.

The leg and metasternal modifications of males of E. alphonsii and
E. californica are correlated with a distinctive courtship behavior. This
involves the male’s stimulation of the female by the vigorous rubbing
of his venter and hind two pair of legs over her elytra. Similar but less
elaborate behavior has been observed in E. wheeled, E. impressifrons,
and E. straba (Pinto, in preparation) .

Adults of E. californica and E. alphonsii are structurally similar to
those of other species of the Caviceps Group although, excepting the
characteristic leg and antennal structure, and a poorly developed eleva-
tion of the elytral suture, they lack all of the other specialized traits listed
by Werner (1955). However, as in E. straba, two of these attributes
(black spots and type of pronotal pubescence) do not apply to these two
species since both are almost completely black and have only sparse
pubescence on the pronotum. In any case, all traits other than leg and
antennal structure vary greatly interspecifically within the group and
most of these attributes are also absent in E. aspera and E. rehni (Wer-
ner, 1955) .

The inclusion of E. californica and E. alphonsii in no way complicates
the diagnosis of the Caviceps Group. The group now contains all species
of North American Epicauta in which the meso- and metafemora and
trochanters of the male are denuded ventrally, the denuded metafemoral

OCTOBER 1972] PINTO CAVICEPS GROUP OF EPICAUTA

255

Fig. 1. Ventral view of head of first instar larvae of Epicauta afoveata. Fig. 2.
Same, E. alphonsii. A similar illustration of the head of E. californica is given
by MacSwain (1956).

area being fringed with long hairs posteriorly, and the mesofemoral
area either fringed posteriorly only, or anteriorly as well.

First Instar Larvae

The first instar larvae of E. californica , E. caviceps , and E. impressi-
frons were described by MacSwain (1956). Those of E. alphonsii, E.
afoveata, and E. wheeleri are described and compared with the former
three species in Table 1. The terminology used in this section closely
follows that of MacSwain (1956) .

MacSwain followed Werner in placing E. californica in a group of its
own, separate from E. caviceps and E. impressifrons. Yet he considered
the three as separable from all other Epicauta by their uniformly dark
coloration and by the strongly convex lateral margin of segment III
of the maxillary palpi.

The original separation of E. californica from the two other members
of the Caviceps Group was justified by several differences. The maxil-
lary and labial palpi of E. californica are longer, the head capsule is
abruptly constricted rather than evenly convergent behind, the sensory
area of segment III of the maxillary palpi is more confined in area and
more mesal in position, and the gular setae are much shorter. With the
discovery of the three additional species, however, a continued division
based on these attributes is no longer tenable. Thus, E. alphonsii agrees
with E. caviceps and allies for some of these traits rather than with its

Table 1. Comparative anatomy of the first instar larvae of six species of the Caviceps Group a

Species

Characters caviceps impressifrons wheeleri ajoveata alphonsii californica

Color dark brown dark brown light yellow dark brown dark brown dark brown ex-

256

THE PAN-PACIFIC ENTOMOLOGIST

[VOL. 48, NO. 4

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Apical-most two or three may not be visible in outline when viewed from above or below.

OCTOBER 1972] PINTO — CAVICEPS GROUP OF EPICAUTA

257

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258

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

closest relative, E. calif ornica; and E. wheeleri and E. afoveata agree
with E. calif ornica and/or E. alphonsii for others (see Table 1) .

As with adult anatomy then, larval characteristics indicate that the
retention of E. alphonsii and E. californica apart from the Caviceps
Group is an unnatural division. Although intragroup relationships
remain obscure, I am tentatively considering E. alphonsii and E. califor-
nica closest to E. afoveata and E. straba.

Based on the six species treated here the larvae of the Caviceps Group
can now be characterized as follows :

Head capsule (Figs. 1, 2) with lateral margins abruptly constricted or evenly
convergent behind. Antennae with conical sensory organ attaining apex of segment
III. Mandibles moderately slender with 7 to 12 teeth. Maxillary palpi with lateral
margin of segment III markedly convex; sensory organ large, dorsomesal (Fig. 3)
or almost completely dorsal in position. Thorax with line of dehiscence present full
length of pro- and mesonotum, line entirely absent on metanotum. Legs with 6
or 7 lanceolate setae on femora. Abdomen uniformly colored; line of dehiscence
absent ; 10 setae present on marginal row of tergites I to VIII ; sclerous evaginations
absent at base of median transverse row of tergal setae, present at base of at least
some marginal setae on segments I to V or VII; first abdominal spiracle slightly
smaller than that of mesothorax, slightly larger than that of segment II; sternum
of segments I to VI unsclerotized (occasionally 2 or 4 minute sclerites present on
segments V and VI) , that of VIII and IX fully sclerotized, that of VII partially
sclerotized.

The uniformly colored abdomen, the posteriorly convergent sides of
the head, the partially sclerotized sternum of segment VII, and the
strongly convex lateral margin of segment III of the maxillary palpi
serve to distinguish species of this group from all other known North
American Epicauta. Based on these traits all six species will key to
couplet 10 in MacSwain’s (1956) “key to the species of Epicauta .” The
only needed modification is the deletion of the first part of couplet 9,
“abdomen yellow or yellow-brown.” Five of the six species of the group
have a dark brown abdomen but in E. wheeleri the body is uniformly
yellow.

In comparing material with MacSwain’s descriptions I disagree in

A*

Figs. 3-5. Scanning electron micrographs of the sensory area on segment III of
the maxillary palpi of species of the Caviceps Group. Fig. 3. Dorsal view of
right palpus of Epicauta alphonsii (800 X ) • Fig. 4. Side view of right palpus of
E. alphonsii showing relative length of two-segmented appendix and surrounding
papillae (1000 X)- Fig. 5. Same, of left palpus of E. impressifrons (1000 X)-
Micrographs made on a Jelco unit (JSMU-3).

OCTOBER 1972] PINTO — CAVICEPS GROUP OF EPICAUTA

259

260

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

only two instances. The terminal seta of the antenna in E. impressifrons
is only slightly longer than segment II, not twice as long; and the two-
segmented appendix on segment III of the maxillary palpi in E. caviceps
and E. impressifrons is subequal to the surrounding papillae (Fig. 5)
rather than twice as long.

Material examined. — MacSwain’s (1956) material of E. impressi-
frons , E. caviceps and E. calif ornica was available for this study. Addi-
tional material examined is as follows :

Epicauta impressifrons. — One specimen from a mass of eight eggs; adults,
Whitewater Canyon, Riverside County, California, 16 October 1970, feeding on
inflorescences of Haplopappus acradineus (Greene) Blake, and Lepidospartum
squamatum (Gray) Gray.

Epicauta wheeleri. — Larvae from a mass of 35 eggs; adults, Whitewater Canyon,
Riverside County, California, 16 October 1970, feeding on inflorescences of Haplo-
pappus acradineus and Lepidospartum squamatum.

Epicauta afoveata. — Three larvae from a mass of 10 eggs; adults, 12 mi. SSW
Borrego Springs, San Diego County, California, 15 November 1970, feeding on
inflorescences of Chrysothamnus paniculatus (Gray) Hall.

Epicauta alphonsii. — Larvae from masses of 20 and 43 eggs, respectively; adults,
4 mi. NNW Lancaster, California, 20 October 1971, feeding on inflorescences of
Chrysothamnus nauseosus (Pallas) Britton.

Epicauta calif ornica. — Larvae from a mass of 45 eggs; adults, Menifee Valley,
Riverside County, California, 5 November 1970, feeding on inflorescences of an
unidentified Compositae.

Acknowledgments

I would like to thank Mr. Steve Ward for his generous assistance in
this study. I am also obliged to Dr. J. A. Chemsak (University of
California, Berkeley) for making Dr. J. W. MacSwain’s collection of
first instar larvae available to me. Plant identifications were kindly
provided by Mr. 0. F. Clarke. Figures 1 and 2 were prepared by Mr.
W. D. Sumlin III.

Literature Cited

MacSwain, J. W. 1956. A classification of the first instar larvae of the Meloidae
(Coleoptera) . Univ. Calif. Publ. Entomol., Vol. 12.

Werner, F. G. 1955. Studies on the genus Epicauta of the North American
Continent (Coleop., Meloidae) II — The Caviceps-Group. Bull. Brook-
lyn Entomol. Soc., 50: 1-12.

Werner, F. G., W. R. Enns and F. H. Parker. 1966. The Meloidae of Arizona.
Agr. Exp. Sta. Univ. Ariz. Tech. Bull., 175.

OCTOBER 1972] SUMMERS & WITT — NESTING OF CHEYLETUS

261

Nesting Behavior of Cheyletus eruditus

(Acarina: Cheyletidae)

F. M. Summers and R. L. Witt

University of California, Davis, 95616

The nesting behavior of some species of Cheyletus is possibly one of
the key factors that severely restricts the period within which mating may
occur. Mating is most apt to occur very soon after Cheyletus females
complete their final moult. The acceptance of males by older, unmated,
gravid females appears to be unusual. Evidence from laboratory cultures
of one species, C. malaccensis Oudemans, suggests that there may be
limited times or special conditions when older virginal females will mate.
In a preliminary study of mating and oviposition in C. malaccensis , we
postulated that the mating of older virginal females may be prevented by
their belligerence after a nest is made (Summers et al ., 1972) .

Local strains of C. malaccensis have both sexes and the females are
facultatively partheno genetic and arrhenotokous. Brooding females of
this and other Cheyletus species reared in our laboratory exhibit marked
ferocity when their nests are disturbed. The place in which eggs are
laid is called a “nest” and it is here that the female exhibits an aggressive
kind of behavior, which may be interpreted as defense of territory or
protection of eggs. This kind of overt behavior has not been detected
before the nest is established.

Cheyletus eruditus (Schrank) was selected for this initial study be-
cause the race being reared has no males and parthenogenesis is the-
lytokous. It is thus possible to observe nesting and oviposition apart from
mating and other complications which the presence of males may intro-
duce. This species is also quite prolific. Our estimates of productivity
by C. eruditus fed on A carus siro Linnaeus greatly exceed the values
recorded by Beer and Dailey (1956) for C. eruditus fed on Tyrophagus
sp.

There were three reasons for setting up this study. The nesting activ-
ities of these predaceous mites are intrinsically interesting. We hope to
pursue further research on the interrelations of mating and nesting
within several of the species which normally have males and the infor-
mation developed in this study may be basic. A third reason is that our
long practice in the handling of isolation cultures of Cheyletus assures
that case histories of individual mites can be followed to completion with
expectations of a low incidence of mortality by accident.

The Pan-Pacific Entomologist 48 : 261 - 269 . October 1972

262

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Methods

Active or moulting deutonymphs of the predator were sealed into isola-
tion cells containing a moderate number of prey mites and five to seven
flakes of wheat bran. The flakes were selected so as to provide nesting
sites acceptable to the mite and convenient for an observer to manipulate.
These were usually polygonal in outline, rigid, somewhat dark in color,
without complicated curls or folds, and cupped on the rough (endosperm)
side.

The isolation cells ultimately developed (Fig. 1) proved to be very
serviceable. Each cell comprised a 10 mm length of thick-walled glass
tube sealed at one end (bottom) with brown cigarette paper and closed
at the other end (top) with thin sheet plastic (Saranwrap®) . The cells
were sawed from a glass tube approximately 32 mm O.D. and 24 mm I.D.
The cigarette paper was permanently affixed with warm, dilute gelatin
and the surplus paper margins later burned away. The circular Saran-
wrap closures were cut with scissors from a paper sandwich, plastic
placed between sheets of rough paper. The top closures were fastened
was a very thin film of vaseline. They could be peeled off and resealed
several times before replacements were required. The tissue paper bottom
was freely permeable to water vapor. Each cell was provided with a
square plastic base or holder, 50 X 50 X 6 mm, drilled with a 24 mm
center hole and a 36 mm countersunk shoulder. The holders made the
handling of the cells much easier. The cells containing mites were stored
at room temperature in a moist chamber having a wire grid shelf sus-
pended over saturated aqueous KC1 (80-85% R.H.). This fairly high
humidity was possibly more vital for Acarus siro (Solomon, 1962;
Kniille, 1965) than for the predators.

Stocks of the prey mite, A. siro , were stored under similar conditions.
The acarids were cultured in glass tubes 60 mm long, 18 mm O.D., which
were sealed on one end with cigarette paper and with a snap-cap on the
other. These mites grew very well when they were fed on wheat bran
plus quick-cooking oats, about 20:1 by volume. The transfer of prey
mites to isolation cells was accomplished easily with the aid of the snap-
caps. A cap removed from a thriving acarid culture was inverted over an
open culture cell and lightly tapped with a pencil. The striking force was
adjusted to sprinkle food mites in small quantities, about 25 to 100 mites.
In this manner there was no carry-over of additional bran or crude
debris.

The unsupported paper bottoms of these isolation cells are resilient,
like the membrane of a drum so that slips of manipulating tools some-
times catapulted both bran flakes and mites out of the cells. Two non-

OCTOBER 1972] SUMMERS & WITT NESTING OF CHEYLETUS

263

Fig. 1 . Components of the culture cells. The one in the lower center is assem-
bled but has no plastic film cover.

flexible tools were generally used: a fine-pointed jeweler’s forceps and
a rigid-shafted needle having a lancet tip. When a nest was operated
upon to remove eggs, shells or dead prey, the cell was lifted from its
plastic supporting base and placed directly on the glass stage of a micro-
scope. Closed forceps were used to press the bran flake (nest) against
the glass-supported paper bottom while the needle was used as a scoop
or pick. Active progeny of the cheyletids were routinely destroyed.

These cells were especially useful for the rearing of Cheyletus because
the species cultured are cryptic and rarely climb far up the walls of the
cells.

Observations

Individuals of C. eruditus are negatively phototaxic and positively
thigmotaxic within limits not precisely determined since the physical
conditions provided were fairly constant. Mites of this species developed
vigorous colonies in glass culture tubes partly filled with bran and
acarids, and they established nests (Fig. 2) under bran flakes dispersed
on the paper bottoms of the isolation cells.

The nesting sites provided were concave flakes of bran having slight

264

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Fig. 2. A prime nest of Cheyletus eruditus on the rough surface of a bran flake.
Females most frequently stand upon or over the egg mass. The highly reflective
spot within the opisthosoma of this female is an accumulation of crystals within the
so-called excretory organ (proctodeum) .

twists or projections to prop them slightly above the bottoms of the cells.
Appreciably elevated pieces were rarely selected by the mites for nesting-
sites if other options were available. The homesites most often occupied
were those in which the vertical dimensions of the free space were not
much greater than the thickness of the mites’ bodies. Individuals entering
nests were not deterred by limited clearance at the access point: they
merely lifted the margins by pushing under. Nesting mites most fre-
quently reposed upside down, clinging to the ceiling of the domicile.
They either crawled under in the upright position and then inverted
themselves, or they crawled in an upright position to the edge of a chip
and inverted as they turned under, as though moving from roof to
ceiling. Body orientation appeared not to be a critical factor in site
selection. The cheyletids were observed to perch in all positions with
respect to the gravity axis, usually with their bodies lightly wedged into
a restricted space. Movements of the mites on nests beneath bran flakes
were often revealed when the flakes themselves appeared to creep.

Two actions were clearly displayed by nesting females. One was

OCTOBER 1972] SUMMERS & WITT NESTING OF CHEYLETUS

265

combativeness. A nesting or brooding mother quickly attacked an in-
truding probe. This overt act involved forward body thrust and strong
clamping motion of the pincer-like pedipalps. Another aspect of the
nesting behavior was the marked tendency of the mother to remain
within the nest, on or very close to her clutch of eggs. The counting of
eggs by an observer often required that the female be pushed off the
pile of eggs. Whenever mothers were completely dispossessed and the
homesites removed, they tended to hover and probe as though trying to
return.

The actions of the predators appeared to vary when prey mites entered
their nests. If food mites were continuously abundant, the predators
either tolerated their presence or confronted and turned them away.
Less often the predaceous species retreated and avoided the intruders.
Of course, at other times, the prey mites were speedily grasped and fed
upon. Nests were frequently cluttered with dry or empty bodies of
acarids. Partly devoured bodies adhered to the nest or to the eggs
therein. The food remnants were sticky and often adhered for a time to
various parts of any of the living mites. It is believed that food mites
whose body fluids were partly consumed at one feeding were used again.
It has been noted that a nesting mother may carry or push food rem-
nants out of the nest. These housecleaning acts were possibly a conse-
quence of efforts of the occupant to feed upon too-dry food remnants.

The deposition of eggs has never been witnessed during the course of
many hours of attention to cultures. The eggs were deposited in clusters
or piles and on the smooth (integument) or rough (endosperm) surfaces
of bran flakes, on paper or on glass. Older, prime nests contained a
single mass or pile of embryonated eggs and empty shells. Common-
place also were a few collapsed eggs fed upon by the mother or her
neonatal daughters.

The eggs are not fastened to the substrate with copious amounts of
adhesive secretion. Their surfaces are tacky, however, and the eggs
remain fairly secure when hanging from above or as placed on vertical
walls. When they are scooped out of nests with a pick, they readily come
away from the surface and it then becomes apparent that they are loosely
bound together with strands of silk. The silk or silk-like strands are
otherwise rarely noticed.

In a preliminary study of reproductive capability and nest occupancy,
18 moulting deutonymphs were placed in separate cells and reared ac-
cording to the four regimes listed in Table 1. In this trial, the cells were
opened and inspected once daily except Saturdays and Sundays. The
eggs were counted individually when possible, otherwise the number was

266

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Table 1 . Summary of 18 case histories, four regimes of treatment.

Vital activities (averages)

Regimes

No. of
observa-
tions

Life

span

(days)

Total
eggs per
female
(est.)

Number
of nests
made

Rl. Normal-minimum disturbance

4

56.8

129.0

5.3

R2. Dispossessed

4

47.8

107.8

8.0

R3. Starved continuously

7

52.6

32.9

2.0

R4. Starved first 46 days

3

77.3

110.7

5.7

estimated. Whenever the mites maintained under the dispossessed regime
(R2) formed nests, they were dislodged at the time of the inspection and
the nests moved to new locations within the cells. The eggs within these
nests were not otherwise molested. The data secured in this experiment
indicate that the periodically dispossessed females (R2) lived shorter
lives, laid fewer eggs and made more new nests than less disturbed
females (Rl). However, the frequency with which displaced females
returned to their original nests was greater than anticipated. There is
possibly no individual specificity among nests because we were able to
interchange nests and mothers with 100 per cent success (n = 6) and
without obvious disruption of the laying functions.

Ten individuals isolated as moulting deutonymphs were maintained
thereafter without food for most or all of their adult lives. Each starved
female deposited a few eggs during the first 15 days of adulthood and
then entered upon a nomadic period during which no nests were estab-
lished and incidents of cannibalism were commonplace. The continuously
starved individuals (R3) did not perish prematurely but their nesting
capabilities were severely curtailed.

Three of the starved individuals (R4) were presented with food on
and after the 47th day. The starved mites had become pale, flat and
somewhat angular in outline. After three days with ample food they
plumped up and resumed laying. In respect to nesting behavior and egg
productivity, the later fed females were not clearly distinguishable from
continuously fed females but their lives appeared to have been prolonged
by the initial fasting period. The presentation of prey to the starved
predators evoked reactions not very obvious among amply fed individ-
uals. When about 50 acarids were sprinkled into the cells for the first
time, the hitherto listless predators quickly became excited or alert to
the proximity of food. Their groping gyrations and other overt motor

OCTOBER 1972] SUMMERS & WITT NESTING OF CHEYLETUS

267

functions were intensified considerably before the first physical contacts
occurred. Starved individuals displayed no arousal from lethargy when
human breath was gently pipetted into their cells. But a burst of search-
ing activity was triggered very soon after the paper bottoms of isolation
cells were placed over the open ends of vials containing cultures of acarid
mites.

The foregoing experiment showed that nesting females must feed and
are able to leave their nests to forage and then return, and that prolonged
starvation induces a state of restlessness or wandering. The estimates of
egg production and the data on duration of nest occupancy are fairly
crude, however, and the natural causes of nest abandonment were not
apparent.

A more refined experiment was then set up to pinpoint, if possible,
the duration and rhythms of oviposition and nest occupancy. In this
trial, 50 individuals were maintained under three regimes of manipula-
tion and with examinations once every day. All were amply fed on
Acarus siro but only the 40 included in R2 and R3 were isolated as
deutonymphs. Individuals in regime R1 were inspected regularly but
disturbed only to the extent necessary to ascertain that the female was
“at home.” Individuals of regime R2 were manipulated as little as
needed to provide accurate counts of eggs laid. Some eggs and all
empties were removed so that, as a rule, about five to ten of the newest
eggs were left after each inspection. Individuals in regime R3 were
dispossessed and their nests taken out whenever found during inspec-
tions. These individuals often scattered their eggs instead of depositing
them in clusters.

The second experiment (Table 2) provides somewhat better assurance
that less disturbed individuals (R2) lived longer lives and laid more
eggs than those whose nests were removed daily (R3). Differences in
degree of manipulation did not appear to affect the duration of the
laying period.

The numbers of eggs obtained under regime R2 showed that egg
production per female, as determined by actual counts, exceeded the
average production per female according to estimated numbers of eggs
(Rl, Table 1). The maximum number of eggs laid on one day was 27
and the highest productivity occurred during the first one-third of the
laying period. All active instars of cheyletids show marked cannibalism.
Accordingly, these estimates exceed all previously reported values be-
cause the active progeny were routinely destroyed and the counts include
the collapsed eggs fed upon by the mother or by recently hatched larvae.

The period during which reproducing females exhibited nesting be-

268

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Table 2. Summary of 50 case histories, three regimes of treatment.

Vital

activities

No. of

Life

Laying

Total

Number

observa-

span

period

eggs per

of nests

Regimes

tions

(days)

(days)

female

made

Rl.

Mild disturbance :
determinations of

10 Mean

—

—

—

2.5

nest occupancy

Range

—

—

- —

1-4

R2.

Moderate disturbance:

19 Mean

61.6*

44.2

169.4*

5.0

nests inverted, some
eggs removed

Range

33-83

22-59

70-317

1-12

R3.

Severe disturbance:
nests removed

21 Mean

49.8*

41.1

141.6*

—

daily

Range

20-76

14-60

77-204

- —

* Differences significant at P = 0.05 by £ test.

havior slightly exceeded the duration of laying. Ten females tested
under regime R1 (Table 2) were maintained in isolation cells only for
data on nest tenure. They were vigorous, young adults when isolated.
These individuals averaged one-half as many nests as the individuals
manipulated as described in regime R2. Three of 10 individuals in the
R1 lot and two of 19 individuals in the R2 lot never left their original
nests, except for brief excursions, until oviposition ended. Among R2
individuals, for which actual egg counts were made, approximately 48
per cent of the eggs were deposited in the first nest and an additional 28
per cent in the second nest. Thereafter the frequency of new nest forma-
tion increased while oviposition waned. Females in regime R2 averaged
3.9 days pre-oviposition, 44.2 days of oviposition and 13.0 days in the
post-ovipositional, wandering phase.

Three extrinsic factors have been observed to terminate or prevent
nesting in laboratory cultures of this species: (1) starvation, (2) over-
crowding with prey mites, and (3) the turning-over of the nests — phys-
ical disruption of the microcavern, too much light or heat. In these
trials, the changing of nests by females in the prime period of oviposition
may have been induced artificially by movements unavoidable in the
handling of cultures. The intervals between the first few nests were short
and not noteworthy. The waning of egg laying correlated with the onset
of discontinuity in nesting, and most of the new nests were established
during the attenuated portion of the oviposition period. In this male-less
race of C. eruditus , belligerent nesting behavior seems to persist through-
out most of the laying cycle.

OCTOBER 1972] SUMMERS & WITT NESTING OF CHEYLETUS

269

Literature Cited

Beer, R. E. and D. T. Dailey. 1956. Biological and systematic studies of two
species of Cheyletid mites, with a description of a new species (Acarina,
Cheyletidae) . Univ. Kans. Sci. Bull., 38, part I, (5) : 393-437.

Knulle, W. 1965. Die sorption and transpiration des wasserdampfes bei der
Mehlmilbe ( Acarus siro L.) . Z. Vergl. Physiol., 49: 586-604.

Solomon, M. E. 1962. Ecology of the flour mite, Acarus siro L. (= Tyroglyphus
farinae DeG.). Ann. Appl. Biol., 50: 178-184.

Summers, F., R. Witt and S. Regev. 1972. Evaluation of several characters by
which five species of Cheyletus are distinguished (Acarina: Cheylet-
idae). Proc. Entomol. Soc. Wash., 72: 230-252.

BOOK REVIEW

The Insect Realm. A Guide to the Hall of Insects. By Charles L. Hogue and
Fred S. Truxal. Los Angeles County Museum of Natural History. 99 pages, 106
figs. 1970. $2.00 [paperback].

Designed to complement the displays in the Hall of Insects of the Los Angeles
County Museum of Natural History, the book admirably fulfills its goal. Although
expressly not intended as a textbook in entomology, it could stand on its own merits
as an introduction to many fascinating aspects of entomology.

The book is not organized in the same sequence as the displays, but an Appendix
with map serves as a cross-reference. The book begins with considerations of the
systematic relationships of insects to other animals and the geologic history of
insects; then to discussions of structure and function, growth and development;
followed by extensive considerations of insects in relation to their environment and
insects in relation to man as medical, agricultural and house and garden pests as
well as beneficial insects; to final sections on the orders of insects and how to
collect and preserve insects.

The discussions and displays are aimed at the general public, but contain suffi-
cient diversity and information to be of interest to undergraduate entomology
majors. The major deficiencies in the book are the discrepancies between the
ordinal names and phylogenetic relationships used in the genealogical chart (page
4) and the major groups chart (page 70), and the lack of many of the display
illustrations. The latter is undoubtedly intentional, so that one must visit the
displays and use the book to provide a record of a memorable tour of “The Insect
Realm.” — R. W. Thorp, University of California, Davis, 95616.

270

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

A New Species of Leptopteromyia

(Diptera: Leptogastridae)

Charles H. Martin
Oregon State University, Corvallis, 97331

The name Leptopteromyia argentinae Martin (1971) appeared in a
key without an accompaning description, hence the species is described
below. Leptopteromyia argentinae Martin belongs to the group in the
genus with longitudinal stripes. As Martin’s (1971) key indicates, it
most closely resembles L. americana Hardy.

Leptopteromyia argentinae Martin, new species

Male. Length 8 mm. Head black, face grayish white tomentose, front and
occiput gray tomentose; mystax with four short whitish hairs, occipital bristles
white.

Thorax yellowish red, median stripe reddish brown extending over anterior de-
clivity, lateral stripes reddish brown, reaching the lateroventral margin of the
thorax, white tomentose laterally over stripes and posteriorly; bristles pale; scutel-
lum blackish, white tomentose; pleura yellowish anteriorly, brown below wing
bases, white tomentose.

Abdomen dark reddish brown, tergite 2 with median yellow band and posterior
margin yellow, tergite 3 with yellow bands on anterior and posterior margins, tergite
4 with narrower yellow bands; tergites with brown semirecumbent hair; epandria
apically tapering to point and curving upward.

Wings slightly brownish.

Hind femora reddish brown, clavus with yellow band basally and yellow spot
apicoposteriorly, hind tibia reddish brown, yellow basally, tapering gradually
from clavus to base.

Holotype male, Argentina Rd., Tucuman, Diovo Codillal, Argen-
tina, 16 January, 1929 (R. Golbach) (Universidad Nacional de Tucu-
man). Allotype, female, V. Padre, R. A. Tucuman-Burruyacu, 17
January, 1948 (R. Golbach) (Universidad Nacional de Tucuman).
Paratype male, same data as allotype (Coll. Charles H. Martin) .

Literature Cited

Martin, C. H. 1971. A review of the genus Leptopteromyia in the western
Hemisphere (Diptera: Leptogastridae). Pan-Pac. Entomol., 47: 264-270.

The Pan-Pacific Entomologist 48 : 270. October 1972

OCTOBER 1972] CLEMENT STENODYNERUS BIOLOGY & LARVA

271

Notes on the Biology and Larval Morphology of
Stenodynerus canus canus

(Hymenoptera: Eumenidae)

Stephen L. Clement
University of California, Davis, 95616

Stenodynerus canus canus R. Bohart is a small wasp about 8—9.5 mm
long and has been recorded from northwestern United States (Idaho,
Washington, Oregon, Nevada, Utah, Wyoming, Montana, Colorado)
and British Columbia (Bohart, 1966). The present paper includes some
fragmentary information on the biology of this species and a description
of the larva.

Markin and Gittins (1967) reviewed the biological information on the
North American species of Stenodynerus. Since their publication, Krom-
bein (1967) and Evans (1970) have added biological information on
the group.

Six nests of S. c. canus were discovered during summer 1971 at a
site (2,395 meters) about 0.6 km west of West Thumb, Yellowstone
National Park, Wyoming. Nests were separated from each other by at
least 0.3 m but were all found in an area of about 2.5 X 6 m.

Natural crevices within a few volcanic outcroppings housed the nests.
The narrow linear crevices ranged from 4—7 mm in width and 1-7 cm
in length. Although there appeared to be no preference for crevices
which faced a particular direction, wasps all selected crevices which
were more or less on a horizontal plane. Each nest entrance was sur-
mounted by a curved mud turret 10—12 mm long and 3.5 mm wide
(Fig. 1).

Personal communication with R. M. Bohart and a review of the liter-
ature reveal that this is the first reported instance of a species of Steno-
dynerus using natural crevices of rocks for nest sites (Fig. 1). Other
species construct turrets but all known species which do so nest in the
ground.

Three female wasps were observed in the act of constructing their
turrets, two on 3 July and one on 15 August. Each wasp spent a few
seconds searching for a dirt particle before transporting it in the mandi-
bles a few centimeters to the nest. The mandibles, legs, and the tip of
the abdomen were used to incorporate the dry particles into and around
the edge of the turret openings. The female wasps apparently mixed
small quantities of water with the dry particles as they were added. Fin-

The Pan-Pacific Entomologist 48 : 271 - 275 . October 1972

272

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Fig. 1 . Upper left. — A partially constructed turret of Stenodynerus canus canus
protruding from a narrow linear crevice. Upper right. — A section of rock showing
the placement of cells and one prepupa in situ.

ished turrets angled upwards 20-40° before curving so that the openings
faced downwards.

One of the wasps observed on 3 July took about two hours to add 5
mm to the length of her turret. The wasp observed on 15 August was
adding material to her partially constructed turret at 11:15 a.m.
(RMDT) . At this time the turret measured 4.5 mm in length. Approxi-
mately four hours later this wasp had finished the turret, which measured
10 mm. Up to two minutes was spent by each wasp adding a single
particle to the turret.

Nests were excavated on 26 June, 3 July, 10 July, and 28 August. Two
completed nests found on 26 June produced one and two cells respec-
tively. A completed nest with four cells was found on 3 July. These
three nests all yielded prepupae. Two nests excavated on 10 July each
produced a completed inner cell with a feeding larva and a partially
provisioned outer cell. The 28 August nest contained a prepupa, inner
cell; a late feeding larva; and an outer cell being actively provisioned.

The length of a crevice may influence the number of cells constructed
by a wasp within it. The completed nest with four cells was housed in a
crevice 7 cm long while the other five nests were found in shorter
crevices.

Widths of each of 11 completed cells coincided with the varying tunnel
widths of the crevices. Four cells provisioned in a crevice with a diam-
eter ranging from 5.5 mm to 7 mm had corresponding widths. The
shorter lengths of these four cells ranged from 4-5 mm. The other cells
varied from 6—7.5 mm in length and 3.5-5 mm in width.

Eggs are apparently laid before provisioning of the cell commences.
An unhatched egg was found at the base of each of the three partially

OCTOBER 1972 ] CLEMENT STENODYNERUS BIOLOGY & LARVA

273

3

Figs. 2-5. Prepupa of Stenodynerus canus canus. Fig. 2. Head capsule,
frontal view. Fig. 3. Spiracle, optical section. Fig. 4. Right mandible, ventral
view. Fig. 5. Right mandible, inner view.

provisioned cells examined. Most reports on the biology of the genus
Stenodynerus mention the attachment of eggs by slender filaments to
cellular walls. I did not observe this characteristic in this study but
future observations of S. c. canus nests may reveal such egg attachments.
My excavations through the rocks housing the nests damaged two of the
observed eggs and in so doing, destroyed any evidence of egg attach-
ments. The third egg could easily have been loosened from a suspended
position during the course of my digging. Subsequent development of
this one viable egg in a gelatin capsule failed.

Prey of this wasp consisted of small caterpillars of the families Gelechi-
idae and Gracilariidae, although only one larva of the latter family was
found in a cell. The number of caterpillars found in the partially pro-
visioned cells were one, four, and eleven.

274

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

After provisioning, the cell is sealed with a mud partition about 0.5
mm thick. The cap of the outer -most cell of two of the completed nests
was flush with the base of the mud turret. The completed nest with four
cells had a vestibular cell about 3 cm long. A separate mud cap about 0.5
mm thick plugged this nest at the base of the turret. The provisioned
cells examined were flush with one another. Summer rains partially de-
stroyed completed turrets and in so doing, separately plugged completed
nests.

Postdefecating larvae line their cells with a thin layer of silk before
entering the prepupal stage. On 3 July two prepupae were transferred
from their cells to gelatin capsules. These were subsequently kept at
room temperature. Later in the laboratory, a male emerged on 23 No-
vember and a female on 3 December. The pupal stage of both lasted
about 12 days.

Grandi (1937) figured and described the larva of Ancistrocerus
parietum (Linnaeus) (as Odynerus ) , a species introduced to north-
eastern United States and Canada from the Palaearctic. It appears, how-
ever, that to date no larvae of eumenids endemic to North America have
been described.

Larvae of S. c. canus (Figs. 2—5) conform to the main characters
used by Reid (1942) to separate eumenid larvae from other vespoid
larvae. The width of the labrum in this species is as great as the width
of the clypeus where the two join, and secondly, the distance from the
antenna to the nearest mandible is less than the distance from the mid-
point on the anterior margin of the labrum to a line drawn between the
bases of the mandibles. Prepupae of S. c. canus also exhibit a median
groove dividing the labrum into two lateral lobes and a membranous area
between the anterior margin of the clypeus and the posterior margin of
the labrum (Fig. 2). Both of these characters are mentioned by Reid
as constituting additional eumenid characters but expressed in varying
degrees of constancy within the group. The following description em-
ploys, in part, the terminology and organization used by Torchio (1970)
in his larval descriptions of two species of the family Masaridae.

Prepupa. — Head-. Integument sclerotized; mandibular apices and articulation,
maxillary and labial palpi, salivary lips, and anterior tentorial pits heavily pig-
mented; antennae, posterior tentorial pits, pleurostomal and hypostomal thicken-
ings, and posterior thickening of head capsule lightly pigmented; head capsule
and mouthparts with few small setae and scattered sensoria; epicranial suture
distinct, incomplete, terminating well above clypeus; parietal bands narrow,
feebly developed; antennae located low on head, each with three sensoria, pair of
small moderately deep pits positioned above antennae; pair of larger shallow
indentations on frons above epistomal suture; epistomal suture well developed, indi-

OCTOBER 1972] CLEMENT STENODYNERUS BIOLOGY & LARVA

275

cated by slightly arched line between anterior tentorial pits; clypeus moderately
protuberant with sensoria more numerous on lower half; labroclypeal suture
distinct, indicated by narrow membranous area between anterior border of clypeus
and posterior margin of labrum; labrum divided by heavily pigmented, median
sulcus into two lateral lobes, distal margin strongly emarginate and minutely
spinulate centrally; mandibles robust, sclerotized, apices tridentate, inner apical
surface concave and limited basally by transverse carina; maxillae distinct, galea
and palpi subapical and conspicuous, palpi with four sensoria; labium with pre-
mentum and postmentum distinct, palpi subapical and conspicuous, each with three
sensoria; salivary opening transverse with sclerotized lips projecting slightly above
prementum. Body: Slightly bent anteriorly, head not touching any abdominal
sternum; 6.5-8 mm long; intersegmental lines complete, fairly conspicuous; dor-
solateral tubercles slightly elevated; spiracles not elevated above body, peritreme
present, atrium slightly sclerotized, lacking spines or ridges, primary tracheal
opening with narrow collar, without spines, subatrium expanded with diameter
equal to that of atrium, posteriorly constricted into primary trachea; anus transverse
slit approximately at midline.

No parasites or other insects were associated with the six nests, al-
though one nest was found adjacent to the nest of a bee, Dianthidium
heterulkei heterulkei Schwarz.

Acknowledgments

Dr. R. M. Bohart determined the Stenodynerus and Mr. M. R. Gardner
(California Department of Agriculture) identified the lepidopterous
prey. I am grateful to Professors R. M. Bohart, R. W. Thorp, and A. A.
Grigariclc for their advice concerning the text of the manuscript. Per-
mission to carry on this study was granted by authorities of Yellowstone
National Park.

Literature Cited

Bohart, R. M. 1966. New Stenodynerus from North America (Hymenoptera:
Eumenidae). Proc. Entomol. Soc. Wash., 79: 73-82.

Evans, H. E. 1970. Ecological-behavioral studies of the wasps of Jackson Hole,
Wyoming. Bull. Mus. Comp. Zool. Harvard Univ., 140: 451-511.

Grandi, G. 1937. Contributi alia conoscenza degli imenotteri aculeati. 16. Boll.
1st. Entomol. Univ. Bologna, 9: 253-348.

ICrombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests, and
associates. Smithsonian Press, Washington, D. C. 570 p.

Markin, G. P., and A. R. Gittins. 1967. Biology of Stenodynerus claremont ensis
(Cameron) (Hymenoptera: Vespidae) . Idaho Univ. Res. Bull., no. 74,
24 p.

Reid, J. A. 1942. On the classification of the larvae of the Vespidae (Hymenop-
tera). Trans. Roy. Entomol. Soc. London, 103: 27-84.

Torchio, P. F. 1970. The ethology of the wasp, Pseudomasaris edwardsii (Cres-
son) , and a description of its immature forms (Hymenoptera: Vespoidea,
Masaridae) . Los Angeles County Mus. Contrib. Sci., no. 202, 32 p.

276

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

BOOK REVIEW

How to Know the Spiders (2nd edition). By B. J. Kaston. Wm. C. Brown Co.,

Dubuque, Iowa, x + 289 pp. + 15 pp., 647 figs. 1972. $5.25.

Dr. Kaston’s original “How to Know the Spiders” has been widely used by both
amateur and student for twenty years. Now in its 2nd edition, this book is even
more useful and informative. As in the 1st edition, the keys are not overly difficult
and are liberally illustrated (with 95 more figures). Unfortunately, the pictures
of the webs did not reproduce satisfactorily, but, a 15-page errata sheet is included
with each book. The introduction is essentially the same, with the addition of
some information on the various numbers of eggs laid and an explanation of the
pit-fall trap method of collecting.

Additions include a short section on parasites and other enemies of spiders and
a list of the families of spiders indicating which of them occur in America north of
Mexico. The section on venoms has been expanded, including a discussion of the
effects of the venom of the brown recluse, Loxosceles reclusa.

The 2nd edition covers four more families, 40 more genera, and 134 more species
than the first, which, along with more distribution data, gives a better coverage of
the western states. Also, the number of species now known from each genus is
indicated in the keys. Finally, name changes or changes in placement have been
indicated by a list in the back of the book. All things considered, those interested
in spiders will find this book very useful and at a price they can afford. — John T.
Hjelle, California Academy of Sciences, San Francisco, 94118.

BOOK NOTICE

Manual of Medical Entomology. By Deane P. Furman and Elmer P. Catts.

National Press Books, Palo Alto, California. 163 pages, 356 figs., 3rd edition

1970, $5.45 [paperback].

This third edition of a standard medical entomology manual has been thoroughly
revised and updated. The total number of illustrations has been increased, but with
some loss in quality (e.g. Blattaria, Mallophaga, immature insects, etc.). Some
keys have been shortened (Coleoptera) and others expanded (immature insects)
and an entirely new chapter on spiders has been added. This new edition should
prove very useful in laboratory courses oriented toward the identification and
recognition of medically important groups of arthropods. — R. W. Thorp, University
of California, Davis, 95616.

OCTOBER 1972 ]

SCIENTIFIC NOTES

277

SCIENTIFIC NOTE

Additional prey records for the forest-dwelling spider wasp, Priocnemis
oregona (Hymenoptera rPompilidae) . — The range of spiders used by P. ( Prio -
cnemissus ) oregona Banks is of interest because this wasp, while widespread on the
Pacific Coast, generally lives in mesic, forested habitats. As a result its activities
take place mainly on shaded ground covered by leaf litter and have rarely been
observed. Until comparatively recently nothing was known of the nesting behavior,
but three species of prey, all primitive burrowing spiders, Actinoxia versicolor
Simon (Ctenizidae) , Atypoides riversi P. -Cambridge (Antrodiaetidae) , and Brachy-
thele sp. (Dipluridae) , were recorded by Hurd and Wasbauer (1956, J. Kans.
Entomol. Soc., 29: 169) and Wasbauer and Powell (1962, ibid., 35:394). We spec-
ulated then that a wider taxonomic spectrum of spiders might be expected for
P. oregona because the closely related P. ( Priocnemissus ) minorata Banks of
eastern North America and European species of Priocnemis use a variety of wan-
dering ground spiders. However, in recent years additional captures of the same
kind of prey previously associated with P. oregona tend to confirm the possibility
that this pompilid displays restricted hunting behavior, specializing on burrowing
Orthognatha.

On two occasions field course groups from the Department of Entomology, Uni-
versity of California, Berkeley, have discovered prey associated with P. oregona
(female wasps identified by M. S. Wasbauer, California State Department of
Agriculture, Sacramento). The habitat in both instances was quite similar: a
partially open area in mixed forest of Lithocarpus, Arbutus, Ceanothus, Pseudo-
tsuga, and Sequoia, marginal to more dense redwoods and a few meters above a
creek. The first was near Alpine Dam, Marin County, where on 10 April 1968,
H. V. Daly, members of the Immature Insects class, and I found a female P. oregona
with a paralyzed, mature female of Antrodiaetus pacificus (Simon) adjacent to a
burrow entrance which was located on a rocky, north-facing, shaded slope. The
second discovery occurred quite fortuitously, on 24 April 1970, at Bates Creek,
about three miles NNE of Soquel, Santa Cruz County, when E. I. Schlinger and
members of the Field Entomology class were excavating trapdoor spider nests at
night. A female P. oregona was unearthed several cm deep within a burrow, along
with a paralyzed spider, a female of Actinoxia versicolor. The burrow entered a
nearly vertical roadbed bank which had many nests of orthognathous spiders.
M. M. Bentzien, who identified both prey spiders and assisted us with the exca-
vation at Bates Creek, suggested that in all probability the burrow was that of this
or a similar spider.

Thus, there are conflicting possibilities as to whether P. oregona digs her burrow
or appropriates that of the spider. Moreover, it is unknown if the prey is usually
stung within its own nest or is enticed or dragged from the tunnel, but these spiders
are nocturnal and do not normally wander from the nest even at night. If the
wasp digs her own burrows and cells, the question remains as to whether the nest
consists of one or more cells, but the chance of discovery of a pompilid and its
prey in a single cell burrow is highly improbable. In New York, Priocnemis
minorata, which uses wandering ground spiders (not Orthognatha), constructs a
nest with up to seven sequentially provisioned cells (Yoshimoto, 1954, Bull. Brook-
lyn Entomol. Soc., 49: 130). Although tunnel digging was not observed by

Yoshimoto and abandoned holes could be appropriated by P. minorata, use of

278

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

primitive spiders and their burrows would be a remarkably divergent adaptation
by P. oregona.

Multi-cellular nests have been described only for the genus Priocnemis among
North American Pompilidae, and use of abandoned arthropod or vertebrate burrows
is known for this and related genera in Europe and South America (Yoshimoto,
ibid. : 138). On the basis of the description of the unnamed wasp, the locality
(Brookdale, Santa Cruz County), and the season (April), the curious account by
F. A. Leach (1921, Wild Life in California, Tribune Publ. Co., Oakland.: 99)
appears to refer to P. oregona. In this case the wasp provisioned a “bulky” spider
about twice her own size in a burrow located about 18 inches up from the base of a
nearly vertical, root-choked roadbed bank. The wasp did not fill the tunnel follow-
ing deposition of the spider (at 3:15 p.m.), and then continued to occupy the
burrow over a five day period of inclement weather. This tends to support the
suggestion that sequential cell occupancy of the nest is practiced by Priocnemis
oregona in a manner similar to its eastern congener. — J. A. Powell, University of
California, Berkeley, 94720.

SCIENTIFIC NOTE

Synonymy of the pselaphid beetles Actium retractum and A. hatchi
(Coleoptera:Pselapidae). — -In our revision of Actium Casey and Actiastes
Casey (1971, Univ. Calif. Publ. Entomol., vol. 67) we indicated that Actium re-
tractum Casey was probably the same species as A. hatchi Park and Wagner but
were unable to locate the type specimen of A. hatchi. Dr. H. S. Dybas, Field
Museum of Natural History, located the specimen and kindly brought it to our
attention. Examination of this type confirmed the suspected synonymy.

Actium retractum Casey, 1908. Can. Entomol., 40(8): 270. Holotype $, Queen
Charlotte Islands, Canada (U. S. National Museum — 38643).

Actium hatchi Park and Wagner, 1961. Univ. Wash. Publ. Biol., 16: 20. Holo-
type $, Snoqualmie Pass, Washington, U.S.A. (Field Museum of Natural
History, Chicago). NEW SYNONYMY — Albert A. Grigarick and Robert 0.
Schuster. University of California, Davis, 95616.

SCIENTIFIC NOTE

New Synonymy in the genus Meloe (Coleoptera :Meloidae) . — In their
recent revision of the North American species of Meloe, Pinto and Selander (1970,
Illinois Biol. Monogr., No. 42) tentatively treated Meloe (Meloe) quadricollis Van
Dyke as a distinct species. They recognized that individuals associated with this
name were almost identical to those of M. calif ornicus Van Dyke, differing hut
slightly in coloration and minor details of pronotal punctation. However, only three

OCTOBER 1972 ]

SCIENTIFIC NOTES

279

females, including type material, were available for study at that time. Since
females of some distinct species of Meloe are almost impossible to distinguish, M.
quadricollis was allowed to stand pending the discovery of males.

A male specimen collected at Davis, California, 9 February 1931 by C. C. Wilson,
with the characteristics of M. quadricollis was recently sent to me by Dr. Fred
Andrews of the California Department of Agriculture. As with the females previ-
ously studied, this specimen differs from typical, completely black, individuals of
M. californicus by the faint metallic green body color, and the slightly coarser pro-
notal punctae only. All other traits, including antennal and genitalic structure,
agree perfectly with those of M. californicus (see Pinto and Selander, Figs. 120
and 178). Considering these important similarities, and the fact that coloration
and punctation are highly variable within species of Meloe, M. quadricollis should
now be treated as a junior synonym of M. californicus. — John D. Pinto, University
of California, Riverside, 92502.

SOCIETY ANNOUNCEMENT

The Pacific Coast Entomological Society announces the publication
of a new book in its Memoir Series: Volume 4: Robert Leslie
U siNGER: Autobiography of an Entomologist.

This unique book, written by one of the outstanding entomologists
of our time, relates personal experiences and episodes from boyhood
through his professional career. Included are accounts of research
expeditions to various countries.

Edited by Drs. E. G. Linsley and J. L. Gressitt, the cloth-bound
book consists of 343 pages, 38 illustrations, and dust jacket. The
book was published in August 1972, price $15.00 (postage extra
and 5% state sales tax to be added for California residents) . Members
of the PCES will receive a 20% discount.

Orders for this limited edition (1,000 copies) may be placed with:

Dr. Paul H. Arnaud, Jr., Treasurer
Pacific Coast Entomological Society
c/o California Academy of Sciences
Golden Gate Park
San Francisco, California 94118

Please make checks payable to the Pacific Coast Entomological Society.

280

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

ENTOMOLOGICAL EXCHANGES AND SALES

Space is available to members and non-members who wish to advertise entomo-
logical exchanges, sales, announcements, etc. We can offer an ad in four issues
for $1.50 per line. Ads will be arranged in order of their receipt. Contact the
Advertising Manager, W. H. Lange, Department of Entomology, University of Cali-
fornia, Davis, California 95616.

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ITAL TUBE IN COLEOPTERA. The classic 1912 monograph and six other papers
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State University Press.

FOR SALE. Orders solicited for Pacific Northwest terrestrial and aquatic insects.
Queries answered promptly. S. G. Jewett, Jr., Route 1, Box 339, West Linn,
Oregon 97068.

FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca,
Department of Biology, University of Arizona, Tucson, 85721.

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Jungles, very rare moth (14" wing span), Scorpion, giant sized Indonesian flying
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Sim Yam Seng, 21, Joon Hiang Road, Singapore, 19, Singapore.

OCTOBER 1972]

INDEX TO VOLUME 48

281

INDEX TO VOLUME 48

Allen, new Baetodes, 123

Analetris eximia, 138

Arachnida

Acarina, Cheyletidae, 261
Pyemotidae, 97
Laelaptidae, 203

Araneae, Theraphosidae, 71, 72, 221
Arhaphe capitata, 5
mexicana, 3

Arnaud, Obituary Notice — G Dallas
Hanna, 59

note on T. W. Cook entomology
library, 63

note on sound production in Agrilus,
69

note on high flight of butterflies, 72
note on dragonfly mass movement, 75
note on distribution of Enlinia, 77
note on butterflies visiting Sax-
fragaceae, 78
Baetodes adustus, 123
andamagensis, 231
bellns, 236
caritus, 124
chilloni, 228
deficiens, 126
fortinensis, 238
fuscipes, 128
inermis, 129
noventus, 130
obesus, 233
pallidus, 132
pictus, 133
sancticatarinae, 240
solus, 231
traverae, 230
tritus, 133
veracrusensis, 239
Bell, note on two Elmidae, 218
Bohart, Pulverro biology, 145
note on Fedtschenkia, 149
Book notices, 85, 96, 120, 276
Book reviews, 61, 252, 269, 276
Brothers, a new Thaumalea, 121
Burke, book review, 61
Caenitoides californica, 46, 48
idahoensis, 49
mexicana, 49

Cheeseman, Paraclunio larvae, 204
Chemsak, Review of Cirrhicera, 86
new cerambycid, 150
Clement, Stenodynerus biology and
larva, 271

Cohen, new Baetodes, 123
Coleoptera

Buprestidae, 69
Cantharidae, 215, 242
Carabidae, 63, 64

Cerambycidae, 21, 86, 150
Curculionidae, 61
Elmidae, 218
Limnichidae, 108
Meloidae, 253, 278
Nitidulidae, 116
Pselaphidiae, 278
Staphylinidae, 218, 249
Correction, 243

Dailey, note on Arizona and Pacific
Slope floras, 74

Davies, note on California damsel fly,

68

Diptera

Anthomyiidae, 81
Asilidae, 51, 94
Bombyliidae, 37
Ceratopogonidae, 244
Chironomidae, 204
Dolichopodidae, 77
Leptogastridae, 270
Muscidae, 81
Sciomyzidae, 8
Stratiomyidae, 76
Thaumaleidae, 121

Edmunds, Acanthametropodinae, 136
Ephemeroptera
Baetidae, 123, 226
Siphlonuridae, 136

Erwin, note on lost Brachinus type, 63
note on egg-hatching in bombardier
beetles, 64

Ewing, Galapagos arthropods, 100
Fender, two new Silis, 215
new Rhagonycha, 242
Fisher, Sepedon systematics, 8
Ford, Galapagos arthropods, 100
Gabel, note tarantula burrows, 72
Gillogly, a new Mystrops, 116
Grigarick, note on pselaphid synonymy,
278

Grissell, Pulverro biology, 145
Hall, new heterotropine Bombyliidae,
37

Halstead, review of Arhaphe, 1
Largus notes and Key, 246
Hemiptera

Largidae, 1, 246

Hewitt, grasshopper cytology, 27
Hjelle, note on tarantula behavior, 71
book review, 276

Huckett, Bigot’s North American type-
specimens, 81
Hymenoptera
Apidae, 153
Andrenidae, 94
Anthophoridae, 175
Cynipidae, 74

232

THE PAN-PACIFIC ENTOMOLOGIST [VOL. 48, NO. 4

Eumenidae, 271
Pompilidae, 277
Sapygidae, 149
Sphecidae, 145
Johnson, correction, 243
Koss, Acanthametropodinae, 136
Landis, Ceramica picta biology, 208
Lang, note on feeding by mite, 203
Langston, note on Philotes sonorensis,
67

Leech, note on F. C. Hottes, 67
note on death of E. R. Leach, 74
note on photocopying entomological
literature, 76

note on Australian sod fly, 76
Legner, new California Orus, 249
Lepidoptera, 78, 189
Danaidae, 72
Lycaenidae, 67
Noctuidae, 208
Nymphalidae, 72, 144
Leptopteromyia argentinae, 270
Leptostylus spermovoratis, 150
Linsley, Callinicus preying on An-
drena, 94

Marer, tarantula eye deformity, 221
Martin, new Leptopteromyia, 270
Mayo, Neotropical mayflies, 226
Mexico litoralis, 113, 114
Moore, note on Diaulota synonymy, 218
note on staphylinid synonymy, 218
new California Orus, 249
Moser, Pyemotes scolyti, 97
Mystrops costaricensis, 116
Neurobezzia, 244
Notice, 107
Obituary notice, 59
Odonata, 67, 68
Libellulidae, 75
Orth, Sepedon systematics, 8
Orthoptera
Acridoidea, 27
Orus frommeri, 251
giulianii, 250

Parkin, D. T., Galapagos arthropods,

100

Parkin, P., Galapagos arthropods, 100
Philip, note on red dragonflies of Tokyo,
67

Pinto, Caviceps Group of Epicauta, 253
note on Meloe synonymy, 278
Powell, note on mass movements of
Nymphalis, 72, 144
note on spider wasp prey, 277
Preissler, Paraclunio larvae, 204
Proceedings, Pacific Coast Entomologi-
cal Society, 66
Prorates arctos, 42
boydi, 40
frommeri, 45
melanderi, 43
nigrescens, 44

Raske, biology of Anelaphus, 21
Ratanaworabhan, new biting midge, 244
Rhagonycha alaskensis, 242
Roton, Pyemotes scolyti, 97
Sakagami, Chinese bumble bees, 153
Schlising, behavior of Diadasia, 175
Schroeter, grasshopper cytology, 27
Schuster, note on Fedtschenkia, 149
note on pselaphid synonymy, 278
Shields, butterflies and flowers, 189
Silis (Silis) alexanderi, 217
spinigerula, 215
Sintoria cazieri, 52
cyanea, 53
lagunae, 55
mojavea, 56
pappi, 57
rossi, 58

Society Announcement — U singer Auto-
biography, 135, 219, 279
Spilman, jumping shore beetle, 108
Summers, nesting of Cheyletus, 261
Tamaki, Ceramica picta biology, 208
Thaumalea santaclaraensis, 121
Thorp, book reviews, 252, 269
Weeks, Ceramica picta biology, 208
Wilcox, Sintoria revision, 51
Wirth, new biting midge, 244
Witt, nesting of Cheyletus, 261
Zoological Nomenclature, 65

MAILING DATES FOR VOLUME 48

No. 1 5 May 1972
No. 2 28 September 1972
No. 3 27 December 1972
No. 4 28 February 1973

Published by the

Pacific Coast Entomological Society

in cooperation with

The California Academy of Sciences

VOLUME FORTY-EIGHT

1972

EDITORIAL BOARD

R. W. THORP, Editor
R. 0. SCHUSTER, Assistant Editor
E. G. LINSLEY
HUGH B. LEECH
E. S. ROSS
P. D. HURD, JR.

P. H. ARNAUD, JR., Treasurer
W. H. LANGE, Advertising

PUBLICATION COMMITTEE

1972

1973

1974

H. V. Daly, Chairman D. D. Linsdale

A. A. Grigarick E. I. Schlinger

J. A. Powell
M. A. Gardner

San Francisco, California

1972

11

CONTENTS FOR VOLUME 48

Arnaud, P. H., Jr.

Obituary Notice — G Dallas Hanna 59

Thomas Wrentmore Cook entomology library 63

Sound production in Agrilus pulchellus Bland 69

High flight of butterflies in San Francisco 72

Mass movement of Sympetrum corruotum (Hagen) (Odonata:

Libellulidae) in central California 75

Distribution of Enlinia calif ornica (Diptera: Dolichopodidae)

in Oregon 77

Butterflies at flowers of Escallonia rubra (Saxifragaceae) in

central California 78

Bell, L. N.

Notes on dry-season survival in two species of Elmidae 218

Bohart, R. M. and E. E. Grissell

Nesting habits and larva of Pulverro monticola 145

Bohart, R. M. and R. 0. Schuster

A host record for Fedtschenkia 149

Book Notices 85, 96, 120, 276

Book Reviews 61, 252, 269, 276

Brothers, D. R.

A new species of Thaumalea from California 121

Burke, H. R.

Book Review: Curculionidae Tribe Ophryastini of North

America (Coleoptera) 61

Cheeseman, D. T., Jr. and P. Preissler

Larval distribution of Paraclunio alaskensis at Point Pinos

sewage outfall, Monterey County, California 204

Chemsak, J. A.

Review of the genus Cirrhicera Thomson 86

A new seed inhabiting ceramycid from Costa Rica 150

Clement, S. L.

Notes on the biology and larval morphology of Stenodynerus

canus canus 271

Ill

Cohen, S. D. and R. K. Allen

New species of Baetodes from Mexico and Central America 123

Dailey, D. C.

Contiguous areas of Arizona and Pacific Slope floras in northern

Baja California, Mexico 74

Davies, T. W.

Oviposition of the California Damsel fly, Archilestes calif ornica

MacLachlan 68

Edmunds, G. F., Jr. and K. W. Koss

A review of the Aoanthametropodinae with a description of a

new genus 136

Erwin, T. L.

The lost type of Brachinus sallei Chaudoir 63

Observations on the life history of bombardier beetles: mech-
anism of egg -hatching 64

Fender, K. M.

Two new California species of Silis 215

A new species of Rhagonycha from Alaska 242

Fisher, T. W. and R. E. Orth

Resurrection of Sepedon pacifica Cresson and redescription of

Sepedon praemiosa Giglio-Tos with biological notes 8

Gabel, J. R.

Further observations of theraphosid tarantula burrows 72

Gillogly, L. R.

A new species of Mystrops from Costa Rica 116

Grigarick, A. A. and R. O. Schuster

Synonymy of the pselaphid beetles Actium retractum and A.

hatchi 278

Hall, J. C.

New North American Heterotropinae 37

Halstead, T. F.

A review of the genus Arhaphe Herrich-Schaffer 1

Notes and synonymy in Largus Hahn with a key to United
States species

246

IV

Hjelle, J. T.

Some observations on tarantula behavior 71

Book Review: How to know the spiders 276

Huckett, H. C.

Notes on Bigot’s North American type-specimens at the Uni-
versity Museum, Oxford 81

Johnson, C. D.

Correction 243

Lang, J. D.

Notes on the feeding habits of Haemolaelaps glasgowi 203

Langston, R. L.

The Sonora Blue — 1971 the earliest season for the north 67

Leech, H. B.

F. C. Hotte's, 1899-1970 67

Death of E. R. Leach 74

On the photocopying of entomological literature 76

The Australian sod fly Altermetoponia rubriceps (Macquart)

in Marin County, California 76

Linsley, E. G.

The robber fly Callinicus calcaneus (Loew) as a predator on

Andrena omninigra Yiereck 94

Marer, P. J.

An eye deformity in a tarantula spider, Aphonopelma reversum _ 221
Martin, C. H.

A new species of Leptopteromyia 270

Mayo, V. K.

New species of the genus Baetodes 226

Moore, I.

Notes on Diaulota harteri with new synonymy 218

Xystridiogaster Scheerpeltz, a synonym of Paracraspedomerus

Moore 218

Moore, I. and E. F. Legner

Two new species of Orus from California 249

V

Moser, J. C. and L. M. Roton

Reproductive compatibility between two widely separated pop-
ulations of Pyemotes scolyti 97

Notice 107

Obituary Notice 59

Parkin, P., D. T. Parkin, A. W. Ewing and H. A. Ford.

A report on the arthropods collected by the Edinburgh Uni-
versity Galapagos Islands Expedition, 1968 100

Philip, C. B.

The red dragonflies of Tokyo 67

Pinto, J. D.

Notes on the Caviceps Group of the genus Epicauta with descrip-
tions of first instar larvae 253

New synonymy in the genus Meloe 278

Powell, J. A.

Mass movements of Nymphalis californica 72

Mass movements of Nymphalis californica (Boisduval) in the

San Francisco Bay area during 1971 144

Additional prey records for the forest- dwelling spider wasp,

Priocnemis oregona 277

Proceedings, Pacific Coast Entomological Society 66

Raske, A. G.

Immature forms, genitalia, and notes on the biology of Anelaphus

albofasciatus Linnell 21

Sakagami, S. F.

Bumble bees collected by the California Academy — Lingnan

Dawn — redwood Expedition to Central West China, 1948 __ 153

Schlising, R. A.

Foraging and nest provisioning behavior of the oligolectic bee,

Diadasia bituberculata 175

Schroeter, G. L. and G. M. Hewitt

Cytology of some California grasshoppers I. Taxonomic con-
siderations 27

Shields, 0.

Flower visitation records for butterflies 189

VI

Society Announcements — Usinger Autobiography 135, 219, 279

Spilman, T. J.

A new genus and species of jumping shore beetle from Mexico ____ 108
Summers, F. M. and R. L. Witt

Nesting behavior of Cheyletus eruditus 261

Tamaki, G., R. E. Weeks and B. J. Landis

Biology of the zebra caterpilar, Ceramica picta 208

Thorp, R. W.

Book Review: The lives of wasps and bees 252

Book Review: The insect realm. A guide to the hall of insects __ 269

Wilcox, J.

The genus Sintoria Hull 51

Wirth, W. W. and N. C. Ratanaworabhan

A new genus of biting midge from California related to

Neurohelea Kieffer 244

Zoological Nomenclature 65

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THE PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

Papers on the systematic and biological phases of entomology are favored, including
articles up to ten printed pages on insect taxonomy, morphology, behavior, life history,
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Essig, E. O. 1926. A butterfly migration. Pan-Pac. Entomol.,

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1958. Insects and mites of western North America. Rev.
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