Vol. 45 JANUARY 1969 No. 1
THE
Pan-Pacific Entomologist
HYNES — The immature stages of Arctoconopa carbonipes (Alex.) (Diptera:
Tipulidae) — 1
TINKHAM AND RENTZ — Notes on the bionomics and distribution of the
genus Stenopelmatus in central California with the description of a new
species (Orthoptera: Gryllacrididae) 4
JENSEN — A new species of Pseudocloeon from Idaho (Ephemeroptera:
Baetidae) — 14
GAGNE — A review of the genus W alshomyia including a new species reared
from Cupressus galls in California (Diptera: Cecidomyiidae) 16
ELLIS AND BORDEN — Effects of temperature and other environmental
factors on Notonecta undidata Say (Hemiptera: Notonectidae) 20
NELSON AND HANSON — The genus Utaperla (Pleeoptera: Chloroperlidae) 26
BRIGGS — A new holarctic family of Laniatorid phalangids (Opiliones) 35
ZOOLOGICAL NOMENCLATURE 56
ALEXANDER — A new subfamily of winter crane flies (Diptera: Tricho-
ceridae) 51
JOHNSON — Horn’s Bruchidae type-material in the Ulke Collection (Coleop-
tera) : 54
BOHART — A new species of Dufourea from California (Hymenoptera:
Halictidae) 57
BUGBEE — New species of the genus Eurytoma from California (Hymenop-
tera: Eurytomidae) 60
SCIENTIFIC NOTE 64
PROCEEDINGS 65
CORRECTION AND ADDITION 80
SAN FRANCISCO, CALIFORNIA • 1969
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. 45
January 1969
No. 1
The Immature Stages of Arctoconopa carbonipes (Alex.)
(Diptera : Tipulidae)^
C. Dennis Hynes
California State Polytechnic College, San Luis Obispo
Recent collections of the immature stages of certain species of crane
flies along the Nooksack River in northern Washington, resulted in the
discovery of the larval and pupal stages of Arctoconopa carbonipes
(Alex.) .
The immature stages were found in a seepage area with a substrate
of fine sand mixed with leaf debris and silt. Of six larvae collected,
two were preserved, three died in rearing cages and were then preserved,
and one was reared through the pupal stage. Just prior to emergence,
the pupa died. However, the genitalia of the developing adult within
the pupa had sufficiently matured so that Dr. C. P. Alexander could
give a specific identification to the specimens concerned. Because of
the scarcity of specimens, the following descriptions are based on five
larvae and two pupae. The description of the head capsule is based
on one specimen.
Arctoconopa carbonipes (Alexander)
Last Instar Larva . — Length 14.0-14.5 mm; dorsoventral and dextrosinistral diam-
eters both 0.6-0.7 mm. Body elongate, cylindrical, with numerous golden setae
closely appressed to integument. Chaetotaxy reduced or absent in many segments.
Spiracular disk with five fleshy lobes; dorsal lobe with heavily sclerotized, thick,
and elongate spine which turns ventrally, ending parallel to surface of disk,
length spine subequal to length of dorsal lobe; darkened area, with four to six
spines at outer margin between each fleshy lobe, central spine larger and darker
than those on either side. Spiracles with reddish-brown centers, outer rims whitish-
yellow. Remainder of markings on disk black (fig. 1). Anal gills expanded at
bases, extending laterally to blunt, fleshy point. Head capsule narrow anteriorly,
wider posteriorly, greatest width occurring at about three-quarters of head capsule
length, length 0.708 mm; dextrosinistral diameter at mandible 0.187 mm, dorso-
ventral diameter at mandible 0.126 mm; clypeal area slightly sclerotized with two
small indentations on caudal margin; frontal area membranous; antennal buttress
sclerotized, curves medially becoming less sclerotized at antennal bases; post-
antennal buttress sclerotized continuing back to junction of dorsal and dorsolateral
^ This investigation is supported by National Science Foundation Grant GB-4532.
The Pan-Pacific Entomologist 45: 1-3. January 1969
2
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Fig. 1. Spiracular disk of A. carbonipes (three-quarter view). Fig. 2. Lateral
view of pupal head region of A. carbonipes. Fig. 3. Lateral view of male cauda
of A. carbonipes.
bars, which are much darker and more heavily sclerotized; dorsal bars are longest
posterior extensions of head capsule, dorsolateral bars spatulate caudally, ending
just before caudal tips of dorsal bars, caudal margins of ventral (maxillary) bars
ending short distance anterior to those of the dorsolateral bars, ventral bars greatly
spatulate at posterior ends; lateral area between posterior mandibular articulation
and ventral bar heavily sclerotized; ocular plate membranous except for sclerotized
area just anterior to premaxillary suture; maxillary plate not toothed.
Pupa . — Length 6.8-8.3 mm; dextrosinistral diameters at base of wing pads both
0.9 mm. Sheaths of labial palps not curved, armed with small spine at tips,
haustellar sheaths forming small triangular area on either side of facial point,
eye sheath with spine at middle of cephalic margin, antennal sheath with blunt,
laterally directed spine both basally and distally on scape, pedicle and flagellar
JANUARY 1969] HYNES — IMMATURE STAGES OF ARCTOCONOPA
3
segments with dorsolateral spines between anterior and posterior margins of
segments, with spines becoming progressively smaller to flagellar tip; cephalic
crest with two blunt nodes occurring at either side of midline. Central anterior
area of mesonotum surrounded by raised, rugose ridge forming just below meso-
notal breathing horns and sweeping cephalad to base of pronotal spines (fig. 2) ;
median carina blunt and slightly sclerotized to level of the pseudosuture, medial
area above the pseudosuture abruptly and broadly white with white area narrowing
and continuing to posterior margin of mesonotum as a thin line; lateral areas of
dorsal crest heavily ridged with patch of short spines arising on either side of
midline; heavily sclerotized ridges formed from dorsal crest to about midlength
of mesonotal surface; lateral carina slightly raised from surface of pupal case.
Mesonotal breathing horns with globular, black bases, entire horn short and
stubby. Leg sheaths end near posterior margins of third abdominal segment;
male and female pupal cauda with dorsal area having five heavily sclerotized
tubercles, which are homologous with the larval spiracular lobes, and bearing
small sclerotized spine, directly caudally at each tip. Dorsal tubercle in addition
with small caudal point (fig. 3).
The body form and structures described above for the genus Arc-
toconopa illustrate a close affinity to Psiloconopa. The closeness of this
relationship is also indicated by the taxonomic history of the group.
Arctoconopa carbonipes was originally described from the adult stage
as a member of the genus Erioptera (Alexander, 1929) and later placed
in Psiloconopa, considered by Alexander (1949) as a subgenus. This
evidence supports Alexander in his regarding Arctoconopa as a valid
genus (Alexander, 1965), since it shows a distinct morphology not
only in the adult stage (Alexander, 1955), but also in the larval and
pupal stages.
Literature Cited
Alexander, C. P. 1929. Undescribed species of Eriopterine crane-flies from the
United States and Canada (Tipulidae, Diptera). Part I. J. N. Y.
Entomol. Soc., 37: 49-50.
1949. Records and descriptions of North American crane-flies (Diptera) Pt.
VIII. The Tipuloidea of Washington, I. Amer. Midland Natur., 42:
257-333.
1955. The crane flies of Alaska and the Canadian Northwest (Tipulidae,
Diptera). The genus Erioptera Meigen. Univ. Mich. Mus. Zool. Misc.
Publ., 90: 1-33, figures.
1965. Family Tipulidae. In, Stone, A., et al. (ed.) A Catalog of Diptera of
America North of Mexico. U. S. Dep. Agr. Handb. No. 276, pp. 16-90.
4
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Notes on the Bionomics and Distribution of the
Genus Stenopelmatus in Central California
with the Description of A New Species
(Orthoptera : Gryllacrididae)
Ernest R. Tinkham and David C. Rentz
Indio, California and Berkeley, California
During the course of the first author’s studies in Nearctic Desert
Sand Dune Orthoptera series, the discovery of a new genus and several
new species of arenicolous stenopelmatine crickets necessitated a study
of the genus Stenopelmatus Burmeister. These studies, begun in 1952,
were greatly aided by four National Science Foundation grants during
the years 1957—1960. To date ten parts have been published in this
series including the description and bionomics of the above mentioned
new genus.
The new species discussed in this paper was first recognized by the
second author while curating Orthoptera in the California Academy of
Sciences. Three specimens collected by E. C. Van Dyke at Coalinga,
Fresno County, California provided the impetus for further investiga-
tion. Mr. Lee D. Wilson, then a California Department of Agriculture
entomologist living in the area, made several trips with the second
author and aided in locating the habitat of the species.
Life History and Distribution of Stenopelmatus
Hebard (1916) was the first to suggest that the life cycle of species
of Stenopelmatus extended over a period of several years. As a result
of numerous roarings, we can now definitely confirm Hebard’s early
observation.
If second or third instar nymphs are collected in the field and
brought into the laboratory they take up to four or five additional years
to mature. Immature specimens are placed in individual jars and fed
lettuce and oatmeal. All species studied, including the new species, took
similarly long periods of time to mature. It is believed that six or more
larval instars occur in the species of this genus. The exuvium may be
either eaten or not eaten after molting.
Frequent confusion in the past, and the resulting large number of
synonyms, has existed due to the failure of students of the group to
recognize adult specimens of Stenopelmatus species. The multi-annual
life history and wingless condition of the adults further adds to the
confusion. There is still no known certain way to determine from
The Pan-Pacific Entomologist 45: 4-14. January 1969
JANUARY 1969] TINKHAM & RENTZ STENOPELMATUS SYNOPSIS
5
Table 1. Seasonal occurrence of species of Stenopelmatus based on
adult and near adult specimens, mostly males. I-indicates initial record
of adult activity, F-indicates final record. See discussion.
(S. interraedius)
(S. intermedius)
F
I
(S. pictus)
(S. longispina)
I
--F
I F
(S. fuscus)
I
--F
Jan. Feb. Mar. Apr. May
Jun.
Jul. Aug. Sep. Oct. Nov. Dec.
external features whether a specimen is an adult. Hebard (1916) was
the first to discover that adult and last instar males possess hooks on
the dorsolateral margins of the supra-anal plate. These hooks appear
as swellings or ridges in earlier instars. Adultness of females cannot
be determined with certainty. Only an estimation of overall size and
development of the ovipositor can help to determine the maturity of
females. Investigation of the development of the internal reproductive
organs (testes, ovaries) might be of great use here.
An attempt was made to determine if geographical separation of
species existed. This was accomplished by examining specimens in the
California Academy of Sciences, California Insect Survey, and the
authors’ personal collections, and then plotting the distributions. At
once tendency towards sympatry became apparent, especially in the
San Francisco Bay Region. Figure 1 shows the distributions of the
species. It must be noted that this map represents the specimens
available to us at the time of the study and must not be taken to
represent the complete range of the species involved or the distribution
of the species in California. Other specimens, which were not seen,
exist in various collections and this group of insects is under study
at The University of Michigan. The genus has a broad distribution in
the state. Our map only represents what we feel is a likely trend towards
sympatry in the distribution of tbe species in central California.
With such sympatry indicated, one immediately seeks to discover
isolating mechanisms. Some ecological separation seems to exist.
S. longispina seems to be limited to coastal areas and is frequently
associated with chaparral; S. intermedins, apparently closely related
to S. longispina, is found at lower elevations in sandy areas but in
relatively the same geographic area. The large species, S. fuscus, occurs
more inland and seems to inhabit a wide variety of habitats. Although
described from San Francisco, S. pictus is likely found only inland
6
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 1
Fig. 1. Distribution of Stenopelmatus species in central California. (See text.)
JANUARY 1969] TINKHAM & RENTZ STENOPELMATUS SYNOPSIS
7
and appears to be most common in the northern portion of the San
Joaquin Valley and western foothills of the Coast Ranges. It is possible
that the type of the latter species is mislabelled since the second author
has lived his entire life in the San Francisco Bay Region and has
collected species of Jerusalem crickets since boyhood. At no time was
S. pictus taken and there are no records of it occurring there in any
of the collections studied.
When the seasonal distribution of mature or nearly mature specimens
is plotted, seasonal isolation is also indicated. Table 1 shows the
resulting chart when adult or subadult males and certain females are
used to plot the seasonal distribution. It is interesting to note that
when the relatively extensive collections listed above were studied it
was found that more than half the specimens examined were immature.
The table is therefore composed from a rather small number of speci-
mens. Mature females may live considerably longer than males if for
no other reason than the cannibalistic habits of certain females (to be
discussed under mating habits) . Table 1 is based largely on males
since this likely gives a better indication of the breeding period.
Mating Behavior in Species of Stenopelmatus
Although Davis and Smith (1926) observed mating of both S. inter-
medins and S. fuscus, they gave no detailed description of their differ-
ences and similarities. They did note that in S. intermedins the male
pursues the female prior to mating and in S. fnscns the reverse was
true. The specimens they referred to as S. fnscns may in fact have
been S. longispina.
The second author twice observed mating of Jerusalem crickets, once
involving S. intermedins, the other occasion involving S. nigrocapitatns
Tinkham and Rentz, new species. Although no comparisons have been
made and it is not known how mating differs specifically from one
species to the other, a pattern was seen in the two matings and several
phases of the act occurred in both observations. A composite descrip-
tion is given below of events seen to be common to both.
A male and female were placed in a wooden observation “arena.”
On several occasions cannibalism occurred promptly. This was prob-
ably due to one or both of the partners not being mature. Immediately
upon detecting the presence of an acceptable mate, the male drummed
his abdomen against the substrate creating an audible sound. This
abdominal drumming can be noted when adult males are kept in jars
or cages with plenty of soil. They form burrows and drum against
the sides, apparently to attract females. This habit is also observed in
8
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Fig. 2. Stenopelmatus pictus. Fig. 3. S. nigrocapitatus Tinkham and Rentz,
new species.
JANUARY 1969] TINKHAM & RENTZ STENOPELMATUS SYNOPSIS
9
Gammarotettix, an arboreal raphidophorine of the same family which
drums against branches and leaves. If receptive, the female Steiiopel-
matus responds by searching for the male. The male also actively
seeks the female using his palpi and drumming while searching. In
addition to the drumming of the male, his abdomen pulsated laterally
almost continually during this period.
Upon encountering one another what appeared to be a mortal combat
began. Clicking of the mandibles of the male and occasionally of the
female was very noticeable at a distance of up to four feet. The female’s
mandibles were agape during most of the encounter as if in defensive
position, but at no time did she attempt to attack the male. Both sexes
clasped the legs of the other with the mandibles but there was no
visible damage at all seen when incompatible pairs were placed together.
These brief “wrestling matches” occurred three to four times prior to
actual mating. They lasted from a few seconds up to a minute after
which the pair would separate, run away, and then commence searching
for one another again. Attainment of the proper mating position
seemed to be lacking in all of these instances.
Successful copulation was accomplished with male atop female, facing
her posterior, grasping the hind legs (left in cases observed) in his
mandibles. The abdomen of the male then curled to meet her genitalia.
During the entire act the hind legs of the male were placed dorsally
between the middle and hind legs of the female at their bases. At the
instant of genitalic contact, the male ejected a fleshy sac possessing
finger-like appendages. A white milky substance, the spermatophore
was then affixed to the genitalia of the female; the male pumped fluid
onto the female’s posterior. A good portion of his abdomen seemed
to be emptied during this process. Pulsations of the abdomen of the
male were most noticeable at this time. The copulation period lasted
for approximately five minutes after which the pair separated and
moved a short distance away. Shortly the female began eating the
external visible portions of the spermatophore.
After one copulation of S. nigrocapitatus the male was immediately
attacked by the female and partially devoured. We believe that this
happens frequently in nature. This would seem to explain the great
scarcity of adult males both in collections and in the field.
The evolutionary significance of a single mating per male is as yet
unclear. Were it not for the multi-annual life history, this could
geographically limit the populations. A detailed investigation of the
internal reproductive system of the male would likely determine whether
multiple matings are likely to occur.
10
Fig. 4. Front view of head of 5. nigrocapitatus Tinkham and Rentz, new species,
holotype male. Fig. 5. Ventral fore tibia allotype S. nigrocapitatus Tinkham and
Rentz, new species; note apical spurs and calcars of right leg. Fig. 6. Ventral view
fore tibia 5. pictus, from Stockton, Calif.; note one pair of apical and subapical
teeth and calcars of right leg. Fig. 7. Ventral view hind tibia S. pictus, Stockton,
JANUARY 1969] TINKHAM & RENTZ STENOPELMATUS SYNOPSIS
11
Key to Adult and Subadult Stenopelmatine
Crickets of California
1. Tibial spines vestigial or lacking on apical dorsal margins of hind tibia.
Ringlet of six apical calcars of hind tibia equal in length, broadly
spatulate, suited for arenicolous habitus. Median or presubapical spur
on ventral surface of fore tibia absent. Pronotum not expanding ante-
riorly. Coloration uniformly orange. (Kelso Dunes, San Bernardino
County, California.) Ammopelmatus kelsoensis Tinkham
Tibial spines prominently developed on apical dorsal margin of hind tibia.
Ringlet of six apical calcars of hind tibia unequal in length, conical
or subconical in form, innermost calcar longest (figs, 7, 13). Median
or presubapical spur on ventral surface of fore tibia present (except
in S. nigrocapitatus Tinkham and Rentz, new species) . Pronotum
expanding anteriorly concealing posterior portion of the large head.
Coloration usually dark, especially on abdomen. (Widely distributed
in the western United States and Mexico.) Stenopelmatus Burmeister (2)
2. Fore tibia bearing only two ventral apical spurs posterior to calcars III
and IV (fig. 5). Hind tibia usually with a single ventral apical spur
(if two, the second very minute) positioned immediately anterior to
calcars III and IV (fig. 13). Occiput uniformly dark black in adults
and late instars (figs. 3, 4). Pronotum reddish
Stenopelmatus nigrocapitatus Tinkham and Rentz, new species
Fore tibia bearing three ventral apical and subapical spurs posterior to
calcars III and IV (fig. 6). Hind tibia with two median equal, ventral
apical spurs of equal length immediately proximal to calcars HI and
IV (fig. 7) 3
3. Adult size very large (35-50 mm in body length) . Color of head and
pronotum orange red. Head often megacephalic 4
Adult size medium to small (usually less than 35 mm in body length).
Color of head and pronotum not orange red but piceous or shining
black, the black isolated into irregular areas by pale sutures 5
4. Calcars of the hind tibia forming a semi-ringlet of six long spurs, the inner-
most two longer, cylindrical in form Stenopelmatus longispina Brunner
Calcars of the hind tibia forming a semi-ringlet of six spurs, these spatulate
or trowel-shaped on inner faces, the inner three relatively equal, but
longer than three outer spurs Stenopelmatus fuscus Haldeman
5. Entire head and body uniformly dark brown with black abdominal tergites.
Hind tibia with four to five internal and two to three external apical
Calif.; note apical pair of spurs and six calcars. Fig. 8. Lateral view of pronotum
S. nigrocapitatus Tinkham and Rentz, new species. Fig. 9. Lateral view pronotum
S. pictus. Fig. 10. Lateral view hind tibia of holotype male S. nigrocapitatus
Tinkham and Rentz, new species. Fig. 11. Lateral view fore tibia holotype male
S. nigrocapitatus Tinkham and Rentz, new species. Fig. 12. Ventral view sub-
genital plate holotype male S. nigrocapitatus Tinldiam and Rentz, new species.
Fig. 13. Ventral view hind tibia holotype male S. nigrocapitatus Tinkham and
Rentz, new species.
12
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45 , NO. 1
dorsal teeth Stenopelmatus intermedius Davis and Smith
Upper part of head shining black with tan sutural areas. Pronotum dor-
sally with irregular areas of shining black. Outer face of femur with
pale brown fasciations (fig. 2). Hind tibia with three to four internal
and two external apical dorsal teeth (fig. 7)
Stenopelmatus pictus Scudder
Stenopelmatus nigrocapitatus Tinkham and Rentz, new species
(Figs. 1, 3-5, 8, 10-13, Table 1)
Diagnosis. — Hind tibia usually with single, median, ventral, apical spur im-
mediately basad or proximad to the base of calcars HI and IV. A minute second
spur may be present. In all other species of the genus there are two median
ventral spurs. The fore tibia is distinctive in having only two apical spurs unequal
in length, one slightly basad of the other. All other Californian species possess
in addition, a ventral subapical spur placed just beyond the median portion of
the ventral surface of the fore tibia.
Holotype Male. — (Measurements in millimeters made with Golgau callipers.)
Body length 28.2; width of head 8.7 X II-7 in total depth; pronotum 6.1 long X
8.8 wide; hind tibia 10.0 X 2.6 wide.
Form typical, medium in size for genus. Head typical of genus, depth of head
from occiput to upper margin of ocular breadth (4.7 mm) measured at narrowest
points. Eyes inverted, pyriform, slightly prominent, similar to S. fuscus and S.
longispina. Head in other structural features typical of other Stenopelmatus
species.
Thorax typical in form. Pronotum not ampliate forward as in S. longispina and
S. fuscus, lateral margins not at all broadened at the ventrad projecting bulge on
anterior lateral third of pronotum. In lateral profile, dorsum of pronotum arcuately
humped in posterior five-sixths, only slight evidence of median suture or groove
as seen in paratypes. Anterior margin of pronotum gently emarginate, typically
fringed with fine decurved hairs. Posterior margin slightly less emarginate than
anterior margin.
Supra-anal plate of abdomen broadly triangulate with typical uncinate hooks at
lateral bases. Cerci acuminate, erect, pubescent. Subgenital plate broadly
spatulate.
Fore legs, with femur in lateral profile, with upper margin strongly arcuate,
ventral margin very slightly arcuate, without spination. Fore tibia unspined dor-
sally with typical apical calcars, three inner long, two outer short, dorsal external
shortest. Ventral surface of fore tibia with two, unequal apical spurs, distal spur
larger.
Median femora typical. Median tibia with five calcars (smaller than those of
fore tibia) , preceded by two apical spurs of similar size, placed approximately in
apical two-thirds of dorsal margin. Ventral surface with pair of apical spurs (only
one on one tibia), smaller than calcars, placed at base of calcars III and IV.
Hind legs typical in form. Femora unarmed. Hind tibia with six apical calcars,
increasing in size gradually from outer-external to innermost calcar. Dorsal surface
with three outer apical and subapical spines, three inner ones more widely spaced,
apical internal spines with apices even with external apical spur. Number II spur
subequal to number HI. Basal spur small, slightly proximal of center. Ventral
JANUARY 1969] TINKHAM & RENTZ — STENOPELMATUS SYNOPSIS
13
surface of tibia with only one apical, distinctive spur situated at base of and
between calcars III and IV (one hind tibia with additional minute spur).
Allotype Female.^ — Total length to apex of ovipositor 31.0; length to base of
ovipositor 29.1; pronotum 4.75 long X 7.0 wide; hind femur 8.1 long X 3.4 wide;
hind tibia 12.2 X 2.0 wide.
Allotype similar to holotype except as follows: size larger, abdomen with supra-
anal plate and cerci smaller; subgenital plate very shallowly and broadly triangular
with acute apex; ovipositor typical, short, recurved, unarmed; median tibia with
pair of apical ventral spurs, hind femora similar to holotype. Hind tibia with
single apical ventral spur on each leg.
Type Data. — Holotype male and allotype, Jacalitos Canyon 6 miles
SSW OF CoALiNGA, Fresno County, CALIFORNIA, 27 January 1961,
D. C. Rentz, collector.
Type Locality. — The type locality is a portion of the Jacalitos uplift,
an area of Paleocene sediments. It is under exploitation by oil com-
panies at the present time. The species seems restricted to the areas
where sandstone occurs and is found under such rocks especially at the
bases of hillsides. The Jacalitos uplift extends to the north for some
50-75 miles and this species should be looked for in these areas. Type
deposition. The holotype and allotype are deposited in the California
Academy of Sciences, number 10,120.
Paratypes.— Same locality data as holotype but with following dates and
collectors: 4 $ $ , 18 2 ? , 27 January 1962, D. C. Rentz, J. R. Heifer, collectors;
7 2 2,1 February 1962, D. C. Rentz, R. Bandar, collectors. Additional paratypes
from Fresno County as follows: Jacalitos Creek road. Rocky Buttes, 20 February
1960, 1 2, H. L. Wilson. Near Coalinga, 31 March 1960, 2 2 2 , H. L. Wilson.
Coalinga, Oil City, 18 March 1931, 3 2 2 , E. C. Van Dyke.
Range of Variation. — Eleven females were measured by the first
author and range as follows; body length 22.6-28.6; pronotum 7.8-8.1
wide X 4.8-5.2 long; hind femur 7.5-9.2; hind tibia 7. 6-8.2 X 1.4-1. 8
mm. Some paratypes, as seen in fig. 3, have the pronotum with brownish
areas but never developed as in S. pictus (fig. 2). Paratypes are
deposited in the authors’ collections, California Academy of Sciences,
and J. R. Heifer collections.
A single female, likely adult, from Visalia, Tulare County, California,
collected on 25 November 1948, W. D. Murray collector, could possibly
represent this species. The specimen is pinned and poorly preserved.
The head and pronotum are similar to the species herein described but
the calcars of the hind tibiae are a bit shorter, perhaps due to wear.
Another specimen, a female from Hot Springs, Tulare County, Califor-
nia, 15 April 1938, collected by E. C. Van Dyke is definitely this species
and demonstrates that the species occurs on the fringes of the San
Joaquin Valley.
14
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Literature Cited
Davis, A. C., and L. M. Smith. 1926. Notes on the genus Stenopebnatus with
description of a new species (Orthoptera) . Pan-Pac. Entomoh, 2: 174-
181.
Hebard, M. 1916. A study of the species of the genus Stenopebnatus found in
the United States. J. N. Y. Entomol. Soc., 24: 70-86.
A New Species of Pseudocloeon from Idaho^
(Ephemeroptera : Baetidae)
Steven L. Jensen
University of Utah, Salt Lake City
While studying the mayflies of Idaho, I found two male images that
represent an undescribed species of Pseudocloeon. I take extreme
pleasure in naming this species for Dr. George F. Edmunds, Jr., in
recognition for his many contributions to the study of Ephemeroptera.
I express my appreciation to Mr. Arwin Provonsha for aid in prepar-
ing the illustrations.
Pseudocloeon edmundsi Jensen, new species
Male Imago (in alcohol). — Length: body 4.5-5.0 mm; forewings 4.5-5.0 mm.
General color light brown. Head light brown; antennae light brown basally, pale
apically; ocelli white; compound eyes divided, upper portion turbinate, separated
dorsally at midline, pinkish -tan, lower portion black (figs. 1-2) . Thoracic notum
reddish-brown; pleural sclerites light reddish-brown, membranes white; thoracic
sterna light reddish-brown. Forewings hyaline, stigmatic region cloudy white,
longitudinal and crossveins hyaline, stigmatic crossveins slightly anastomosed
(fig. 3) ; hind wing absent. Legs white, faintly brown at apex of tarsi. Abdominal
tergum 1 subhyaline light brown; terga 2-6 subhyaline white with very faint
black spiracular markings; terga 7-10 light tan, opaque. Abdominal sterna 1-6
same color as terga; sterna 7-9 white, opaque; subgenital plate and forceps white
(fig. 4). Caudal filaments white.
Types. — Holotype male imago. Snake River at U. S. Highway
#20-26, 8 MILES NW. Parma, Canyon County, Idaho, 29 August 1958,
G. F. Edmunds, Jr., and R. K. Allen, in collection of University of Utah,
Salt Lake City. Paratopotype : I male imago, same data and deposition
as holotype.
Remarks. — Pseudocloeon edmundsi is the fourth species in the genus
known from western North America. The other three species, P. futile
McDunnough, P. ruhrolaterale McDunnough, and P. turbidum McDun-
^ The research upon which this report is based was supported in part by a University of Utah
Graduate Research Fellowship.
The Pan-Pacific Entomologist 45: 14-15. January 1969
JANUARY 1969] JENSEN A NEW PSEUDOCLOEON FROM IDAHO
15
Figs. 1—4. Pseudocloeon edmundsi, male imago, paratype: Fig. 1. dorsal view
of head; Fig. 2. lateral view of head; Fig. 3. forewing; Fig. 4. male genitalia.
nough, were originally described from Alberta, Canada. P. edmundsi
can easily be distinguished from these species by thoracic and abdominal
color pattern. The following key, adapted from Traver (1935), serves
to distinguish the male images of the four species.
1. Abdomen mostly or completely white or yellowish-white 2
Abdomen clay-colored or dark olivaceous brown 3
2(1). Abdomen subhyaline white with very faint black spiracular markings;
thoracic notum reddish-brown P. edmundsi, new species
Abdomen yellowish-white with lateral maroon patches; thoracic notum
black P. rubrolaterale McDunnough
3(1). Thorax and abdomen dull olive brown or clay-colored, terga 7-10 pale
ochreous; forelegs greyish-white P. futile McDunnough
Thorax dark brown; abdomen dark olivaceous brown, terga 7-10 similar
to preceding segments; forelegs with femora olive brown, tibiae and
tarsi white P. turbidum McDunnough
Literature Cited
Traver, J. R. 1935. Systematic, Part II, pp. 267-739. In: J. G. Needham, J. R.
Traver, and Yin-Chi Hsu, The Biology of Mayflies. Ithaca; Comstock
Publishing Co. XVI + 759 pp.
16
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
A Review of the Genus Walshomyia Including A New
Species Reared from Cupressus Galls in California
(Diptera : Cecidomyiidae)
Raymond J. Gagne
Entomology Research Division, Agr. Res. Serv.,
USD A, W ashington, D. C.
The western North American genus Walshomyia Felt (Cecidomyi-
inae: Oligotrophidi) may he distinguished from all other North
American Oliogotrophidi by the following combination of characters:
palpus short, one or two segmented ; R 5 curved distally to j oin C behind
wing apex; claws short, each about one-half as long as empodium;
ovipositor short, barely protrusible, the upper lamella entire or bifurcate.
The species comprising Walshomyia are gallmakers on species of
Juniperus and Cupressus. The genus differs from Oligotrophus Kieffer
only in the number of palpal segments. In more important characters,
such as the genitalia, Oligotrophus hetheli Felt and Oligotrophus patter -
soni White, both reared from Juniperus, actually show a greater re-
semblance to species of Walshomyia than to the other Nearctic species
of Oligotrophus. Nevertheless, Walshomyia does key down well as a
group, and its possible synonymy with Oligotrophus should await a
generic revision of the super-tribe Oligotrophidi.
Felt assigned three species to Walshomyia: W. juniperina Felt (the
type species), W. texana Felt, and W. insignis Felt. I am here trans-
ferring Rhopalomyia sahinae Patterson to Walshomyia and describing
one new species, Walshomyia cupressi, to bring the total number of
species in the genus to five. A yet undescribed species belongs here;
it was reared from Libocedrus decurrens Torr. in Yosemite Park, Cali-
fornia and referred to by Felt (1940: 46) as Rhopalomyia sp. Felt’s
specimens of this species consist only of several pupae and several very
teneral adults that I pulled out of the pupal skins. Although it is obvious
that they are a new species of Walshomyia, they are not in good enough
condition to warrant a description and are left undescribed for the
present.
I can find no morphological differences to separate W. juniperina
from W. sahinae and would consider them synonyms except that W.
juniperina was reared from fruit of Juniperus (Felt, 1908) and W.
sahinae from apical bud galls (Felt, 1921) of Juniperus. Although
Felt stated that the palpi of W. juniperina were one segmented (Felt,
1915) and those of W. sahinae two segmented (Felt, 1921), both one
and two segmented palpi are present in both type series.
The Pan-Pacific Entomologist 45: 16-19. January 1969
JANUARY 1969] GAGNE— REVIEW OF GENUS WALSHOMYIA 17
Fig. 1. Female antennal flagellomere VIII of JF. insignis. Figs. 2-5. Antennal
flagellomere V of IF. cupressi: Figs. 2-3, female, ventral and dorsal view, respec-
tively; Figs. 4-5, male, ventral and dorsal view, respectively.
Key to Adults of Species of Walshomyia
1. Ovipositor (fig. 7) partially sclerotized, dorsal lamella entire or bifurcate,
ventral lamella small, less than half as long as dorsal; male sternum X
entire 2
Ovipositor (fig. 9) completely soft, dorsal lamella bifurcate, ventral lamella
large, almost as long as dorsal; male sternum X bifurcate; reared from
apical bud galls on Cupressus spp. JF. cupressi, new species
2. Female antennal flagellomeres (fig. 1) with interconnected striations (a
second, smaller set of circumfila?) crisscrossing each flagellomere;
setulae on flagellomeres present only laterad and proximad of basal row
of setae; dorsal lamella of ovipositor entire, similar to that of JF.
juniperina; reared from oval, apical bud gall on Juniperus scopulorum
Sarg. (only female known) JF. insignis Felt
Female flagellomeres without striations and covered with setulae; dorsal
lamella of ovipositor entire or bifurcate 3
3. Dorsal lamella of ovipositor bifurcate; aedeagus and claspettes nearly
straight, posteriorly directed (as in fig. 11) ; reared from apical bud
galls on Juniperus ashei Buckholz (= Sabina sabinoides L.)
W. texana Felt
Dorsal lamella of ovipositor entire (fig. 7) ; aedeagus and claspettes re-
curved dorsally (fig. 6) 4
4. Reared from fruit of Juniperus utahensis Engelm. JF. juniperina Felt
Reared from conical apical bud galls of J. ashei and /. monosperma
(Engelm.) Sarg. JF. sabinae (Patterson) n. comb.
18
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
8
Figs. 6-7. Lateral view of male genitalia and female ovipositor, respectively, of
flF. juniperina. Fig. 8. Dorsal view of lamellae of ovipositor of W. cupressi. Fig.
9. Lateral view of same. Fig. 10. Dorsal view of male genitalia of IF. cupressi.
Fig. 11. Lateral view of same.
JANUARY 1969] GAGNE — REVIEW OF GENUS WALSHOMYIA
19
Walshomyia cupressi Gagne, new species
Adult. — Head . — Male antenna with 17 flagellomeres, each closely girdled by
2 circumfila (figs. 4-5) ; female antenna with 17-18 flagellomeres, each with
circumfila forming a reticulate pattern ventrally (figs. 2-3) ; eyes many facets
long at vertex but separated by about diameter of one eye facet; palpus with one
short, cylindrical segment; labella small, only slightly longer than palpus, rounded
apically, covered with setae as long as labella. Chaetotaxy: dorsocentral setae,
95-125; parascutal: anterior, 23-34, posterior, 12-19; mesanepimeral, 24-36;
dark scales covering legs, wing membrane, halter, and pregenital segments of
abdomen; scales absent from thoracic sclerites. Wing length, 3.75-4.40 mm.
Proportions of segments of foreleg: femur, 1.00; tibia, 1.10-1.14; tarsomere I,
0.09; II, 0.64-0.70; III, 0.29-0.33; IV, 0.21-0.23; V, 0.08-0.09. Male genitalia
(figs.) 10-11) ; tergum X bilobed, the lobes rounded, setose; sternum X bifurcate;
basimere long, cylindrical; distimere long, flattened-cylindrical, covered dorsally
with many short setae ; aedeagus long, straight, tapering gradually to rounded apex,
many small, postero medially pointing setulae present apicodorsally; claspettes
very short, setose. Female postabdomen: tergum VIII quadrate, covered with setae
and scales; ovipositor short, unsclerotized, about 0.2 length of entire abdomen,
barely protrusible; dorsal lamella (figs. 8-9) bifurcate, setose; ventral lamella
almost as long as dorsal, setose ventrally.
Larva. — Spatula absent. Papillae: two groups of three laterals each on either
side of median line of thorax; six haired dorsals; two haired hind ventrals; four
unhaired fore ventrals on abdominal segments; and four terminals with hairs no
longer than diameter of papillar bases.
Material Examined.- — Holotype male, ex Cupressus pygmaea. Ft. Bragg,
Mendocino County, California, 21 April 1967, G. W. Frankie, U.S.N.M. Type
No. 69984. Paratypes: 4 ^ 4 2 2,1 larva, same data as holotype; 6 2 2,
ex Cupressus sargentii, 3 mi. SE Occidental, Sonoma Co., Calif., collected 1 March
1968, G. W. Frankie, emerged 10 to 18 March 1968.
Mr. Gordon W. Frankie, University of California, Berkeley, collected
and reared the specimens described and kindly submitted them to me
for description. This species causes a gall on buds of Cupressus
pygmaea (Lemmon) Sarg. and C. sargentii Jeps. The gall is a greatly
swollen bud. Mr. Frankie is presently studying the biology of this
species.
Literature Cited
Felt, E. P. 1908. Appendix D. N. Y. State Mus. Bull., 124: 286-422.
1915. Appendix: A study of gall midges. III. Ihid., 180: 127-288.
1921. New species of reared gall midges (Itonididae) . J. N. Y. Entomol.
Soc., 29: 115-118.
1940. Plant Galls and Gall Makers. Ithaca, N. Y., 364 pp.
20
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Effects of Temperature and Other Environmental Factors
on ISotonecta undulata Say
(Hemiptera: Notonectidae)
R. A. Ellis and J. H. Borden^
Simon Fraser University, Burnaby, British Columbia
Notonecta undulata Say is widely distributed throughout North
America (Brooks and Kelton 1967, Bueno 1905). Exhaustive studies
of external anatomy, life history and some briefer notes on the ecology
and behavior of this aquatic insect have been made (Hungerford 1919,
Clark 1928 and Essenburg 1915). This paper concerns the importance
of temperature as a factor in this species’ distribution as determined by
both field and laboratory measurements.
Materials and Methods
Specimens were collected mainly from Lost lake, B. C., a small bog
pond rich with aquatic insects. About 15 adults per 15 gallon aquarium
could be maintained without significant cannibalism if a surplus of
fresh food was provided daily. Flour beetles, bark beetles, ants, cori-
xids, aphids, Daphnia sp. and fry of the common guppy, Lebistes reti-
culatus Peters were supplied as prey. The aquaria were provided with
sand, filters, aerators, heaters and lighted canopies. Water temperature
was maintained at 25 ± 2'’C, pH at 6.6 and water hardness at 25 ppm.
Bunches of Anacharis canadensis Michx. were provided as sites for
oviposition and protective cover. Notonecta undulata will oviposit on
a wide variety of objects in captivity including the glass walls of aquaria,
plastic filter boxes and rubber sink matting. It appears to prefer
plants to these, however, probably because plants are easier to cling to.
When eggs were oviposited on plants they could then be transferred
without damage to an incubation aquarium.
In spring and summer, 1967, a survey was made of fresh water habi-
tats in the Lower Fraser Valley, B. C., to determine where and under
what conditions N. undulata was most abundant. Water temperature,
pH, and hardness were measured and notes on the major flora and
fauna recorded whenever this species was found.
A simple temperature preference apparatus, similar to that used by
Omardeen (1957) for Aedes aegypti (L.) was set up in the laboratory
under constant light. An aluminum trough filled with wet sand was
^ Graduate student and Assistant Professor respectively, Department of Biological Sciences.
The Pan-Pacific Entomologist 45: 20-25. January 1969
JANUARY 1969] ELLIS & BORDEN NOTONECTA ECOLOGY
21
placed with one end on a hot plate, the temperature of which could be
controlled and the other end in an ice bath. An aluminum based glass
trough (15 X 15 X 60 cm) was placed on top of the sand and filled to
1.5 cm with filtered aquarium water. This allowed free horizontal move-
ment by the backswimmers, while limiting convection currents which
occurred in deeper water. Temperature was measured by 8 thermom-
eters suspended at equal intervals throughout the length of the apparatus.
With this set-up a temperature gradient (eg. 15-20°C) could be estab-
lished in 30 minutes by adjusting the control dial of the hot-plate.
Insects tested for temperature preference were 18 first instars, 4 fifth
instars and 7 adults in 42, 33 and 65 trials respectively. Insects were
introduced into the middle of the gradient apparatus and left approxi-
mately 5 minutes before their position was recorded. The difference
in temperature between the two ends of the apparatus varied from 13.0
to 27.0°C (first instars), 7.5 to 10.0°C (fifth instars) and 4.0 to 5.0°C
(adults). Counts of individuals at each temperature were made every
5 minutes after shifting the gradient several degrees for each trial; no
2 gradients were the same.
The upper limit of water temperature in which N. undulata can sur-
vive was determined using large glass beakers (3000 ml) heated with
standard aquarium heaters and gently aerated to ensure uniform water
temperature. The number of survivors at each temperature was noted
hourly, for nine hours. Death was verified by returning an insect to
25°C to check for revival. Temperatures and number of adults tested
were 34, 35, 36, 37, 38, 39, and 40°C and 6, 22, 13, 17, 20, and 19
respectively.
Results
Temperature Preference. — The mean preferred temperatures for
first and fifth instar nymphs and adults were 27.5, 27.1 and 27.0°C
respectively (Fig. 1). None of the insects tested ventured into water
exceeding 39°C and only one was found at 16°C, the lowest temperature
recorded. As the temperature of the ‘hot’ end was slowly increased or
decreased (2-5°C per 5 minutes) the majority of the individuals (es-
pecially first instar nymphs) grouped in the optimum area would, al-
most as a unit, shift their position to the water temperature at which
they had just been.
Upper Limits of Temperature. — All six adults tested at 34°C sur-
vived over 9 hours (Fig. 2) . Mortality slowly increased above this tem-
perature, while the premortality exposure time decreased. At 9 hours,
percent mortality varied almost directly with temperature.
22
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
100
90
CD
zs
"O
80
TJ
_C
“o 70
O)
E 60 -
3
Z
50 -
40
30
20
10
First Instars
Mean = 27.5°C
(N=778)
15 20 25 30 35 40
Fifth Instars
Mean = 27.1°C
(N = 132)
15 20 25 30 35 40
Preferred Temperatures (°C)
Adults
Mean-27.0°C
(N=451)
15 20 25 30 35 40
Fig. 1. Temperature preference of N. undulata nymphs and adults in a tem-
perature gradient.
At high temperatures the backswimmers repeatedly tried to escape.
They swam excitedly below the surface, climbed up the air-line out of
the water, or swam dorsal side-up on the surface. The beakers were
covered with paper towelling to prevent any of the insects from es-
caping.
Distribution and Habitat Conditions. — The wide variety of aquatic
habitats in which N. undulata exists and reproduces is illustrated by 4
examples. In early May it was found in a swimming pool (Coquitlam,
B. C.) with a high chlorine content (approx. 40 ppm) relative to natural
waters. The water temperature was 10.0°C, water hardness 28 ppm
(very soft) and the pH 6.0. Vegetation was limited to a variety of fresh-
water algae (e.g. Nostoc and diatoms). Two species of dytiscids and
several gerrids were abundant in the same pool. All were observed feed-
ing on terrestrial insects trapped in the surface film. Mating and ovi-
position of N. undulata began in mid-May. First instars were observed
in this pool when the water was 17°-20°C in later May.
Lost Lake (B. C.), a bog pond 150 x 200 m, contained hundreds
of backswimmers in various instars throughout May, June and July. N.
undulata was the predominant backswimmer but a Buenoa sp. was rela-
tively numerous for about three weeks. All the major orders of aquatic
insects were present. The water was very soft (30 ppm) , the pH 6.0, and
JANUARY 1969] ELLIS & BORDEN NOTONECTA ECOLOGY
23
40°
39°
38°
37°
36°
35°
34°
Fig. 2. Effect of exposure to high temperatures on the survival of N. undulata
adults.
when the air temperature was 22.0°C the shallow water near the shore
inhabited by the N. undulata was 24.0°C. A varied and luxurient array
of aquatic plants grew around the shallow margins.
Deer Lake (Burnaby, B. C.) contained relatively few N. undulata.
Other aquatic insects (mainly corixids) were present but not in large
numbers. When the air temperature was 29.0°C the surface water was
21.0°C, water hardness was 34 ppm; pH was 7.1. Food was abundant
at this time as evidenced by large plankton hauls of Daphnia sp. Vegeta-
tion also was plentiful.
Several hundred N. undulata in the later instars were found in an ex-
cavation pond (10 m^ and 0-25 cm deep), and a shallow drainage ditch
leading from it on Burnaby Mountain, B. C., during late July. Also
24
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
found in the pond were several hundred tadpoles, several dytiscids and
a few gerrids. When the air temperature was 28.0°C the pond was
32.5°C and the ditch 29.0°C. Both pond and ditch had a pH of 7.0 and
water hardness of 220 ppm (very hard) . Only a few emergent plants
were present.
Discussion
Laboratory trials and field measurements indicate that N. undulata
is able to sustain itself over a wide range of water temperature (4-
32°C), water hardness (28-220 ppm) and pH (6.0— 7.1). The temper-
ature range in which mating and oviposition can occur appears nar-
rower, 10-32°C (Hungerford 1919). Four generations per year can be
reared in laboratory aquaria kept at 25°C but there are usually only
two generations per year in Southern British Columbia habitats — ^one
in late spring and the other in late summer.
Although N. undulata can withstand relatively high temperatures lor
short periods of time (Fig. 2), 34°C is the highest temperature at which
no mortality occurred in the 9 hour trials. Exposure for this period would
not normally occur in backswimmer habitats. Above 34° C the survivor-
ship decreased rapidly. These findings are consistent with the observa-
tion that at a cuticle temperature between 32-34 °C there is a sharp
increase in water evaporation rate in Notonecta sp. This is probably
because of changes in molecular orientation of the cuticular lipid mono-
layer and in nature would result in lethal water uptake (Beament 1961) .
In nature N. undulata will attempt to escape from water at 34°C. When
the water in the excavation pool, mentioned above, reached 32.5°C many
adults were observed swimming rapidly at the surface and flying away.
Habitats in S. W. British Columbia, would seldom reach 34° C.
The ability to reproduce over a fairly wide range of temperatures and
to withstand high temperatures during the reproductive season would
allow N. undulata to inhabit small, sometimes temporary, bodies of
water where the temperature may rise far above the preference level and
vary considerably in a short time.
Notonecta undulata often occurs in the same interior B. C. lakes as
Notonecta kirhyi Hungerford which lives in waters with a pH of 7.0 to
9.2 (Scudder 1965). Thus, N. undulata, can probably live and repro-
duce in water with a pH of 6.0-9.2. It would appear then that pH, as
well as water hardness, has little direct effect on the habitat preference
of N. undulata. These factors may indirectly influence prey abundance
and protective cover by determining the abundance and species com-
position of shore plants.
JANUARY 1969] ELLIS & BORDEN NOTONECTA ECOLOGY
25
Clark (1928) summarized the characteristics of an optimum habitat
for N. undulata in the Winnipeg region: a pool teeming with aquatic
life, vegetation, no current, ample sunlight and a certain amount of
water not covered by algae. Lost Lake (B. C.), which fits this descrip-
tion very well, supported the largest population of N. undulata in the
survey, yet the excavation pool had almost as dense a population but
with very little vegetation. However, the presence of N. undulata in
almost all types of relatively static waters in S. W. British Columbia is
evidence of its tolerance of a wide range of environmental character-
istics.
Acknowledgments
We are grateful to Dr. R. Sadleir for advice and assistance, to Dr.
F. S. Truxal for taxonomic information, and to Drs. N. H. Anderson,
G. G. E. Scudder and S. B. Haven for review of the manuscript.
Literature Cited
Brooks, A. R. and L. S. Kelton. 1967. Aquatic and semiaquatic Heteroptera
of Alberta. Saskatchewan and Manitoba. Mem. Entomol. Soc. Can.,
51: 38-42.
Beament, J. W. L. 1961. The waterproofing mechanism of arthropods II. The
permeability of the cuticle of some aquatic insects. J. Exp. Biol., 38:
277-290.
Bueno, J. T. de la Torre. 1905. The genus Notonecta in American north of
Mexico. J. N. Y. Entomol. Soc., 13: 143-167.
Clark, L. B. 1928. Seasonal distribution and life history of Notonecta undulata
in the Winnipeg region, Canada. Ecology, 9: 383-403.
Essenburg, C. 1915. Habits and natural history of the backswimmers. J. Anim.
Behav. (Cambridge), 5: 381-390.
Hungerford, H. B. 1919. The biology and ecology of aquatic and semi-aquatic
Hemiptera. Univ. Kans. Sci. Bull., 2: 1-341.
Omardeen, T. a. 1957. The behavior of larvae and pupae of Aedes aegypti
(L.) in light and temperature gradients. Bull. Entomol. Res., 48:
349-357.
Scudder, G. G. E. 1965. The Notonectidae (Hemiptera) of British Columbia.
Proc. Entomol. Soc. Brit. Columbia, 62: 38-41.
26
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
The Genus Vtaperla^
(Plecoptera : Chloroperlidae)
Charles H. Nelson^ and John F. Hanson
Department of Entomology, University of Massachusetts, Amherst
The genus Utaperla was proposed by Ricker (1952) for a single new
western North American species Utaperla sopladora Ricker. Recently,
while examining some stonefly material from China, it was our good for-
tune to encounter a species new to science that belongs in this formerly
monotypic genus. The discovery of the new species has led the authors
to review the genus Utaperla in order to more clearly characterize this
group.
Utaperla Ricker
1952, Utaperla Ricker, Indiana Univ. Publ., Sci. Ser., 18: 174 {U. sopladora type
species by original designation and by monotypy) .
1964, Gaufin, Gewasser Abwasser, 34/35: 39.
1965, lilies, Annu. Rev. EntomoL, 10 : 134.
1966, lilies Das Tierreich, 82 : 428, fig. 19 (distribution map).
1966, Gaufin, Nebeker and Sessions, Univ. Utah Biol. Ser., 14 (1) : 21, 25, 28, 72.
Ricker (1952) based on his study of U. sopladora, placed Utaperla
within the subfamily Paraperlinae of the Chloroperlidae and character-
ized it as follows, “The head capsule is slightly elongated, but much less
so than in Paraperla and there is no straight sided section behind the
eyes. The general aspect, in fact, is that of an Alloperla of the subgenus
Sweltsa. However, the basal body of the male supra-anal process is sus-
pended at the anterior edge of the 10th tergite and lies free in the deep
median cleft, instead of being attached to the sides of the cleft as in Allo-
perla. On the 7th sternite there is a raised knob, thickly covered by
spines. The venation also is much like Paraperla, though with fewer
crossveins.”
Of the preceding characters only those involving the head capsule
clearly distinguish U. sopladora from the species of other paraperline
genera, especially from those of Paraperla. The supra-anal process, con-
trary to Ricker’s observation, does not lie free in the median cleft, but
is typical of the processes of most stoneflies in that it is held to the
hemitergites of the tenth segment by the cowl which attaches to the
^ Contribution No. 1396 from the Department of Entomology (Systematics and Morphology Research
Laboratory), University of Massachusetts. Published with the aid of the Guy Chester Crampton
Research Fund of the University of Massachusetts.
^ Personal contribution No. 2 from the senior author.
The Pan-Pacific Entomologist 45: 26-34. January 1969
JANUARY 1969] NELSON & HANSON UTAPERLA REVISION 27
0.2 5 mm
Figs. 1-3. Utaperla sopladora Ricker. Fig. 1. Head, dorsal view. Fig. 2.
Thoracic sterna. Fig. 3. Thoracic terga. (eel =: ecdysial cleavage line; pocs rz post-
occipital suture; sut = mid-ventral suture of meso thoracic furcasternum; scl' =
scutellar ridge) .
28
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
basal lateral sides of the free part (basal body). The slight ventral
production of the posterior half of the seventh sternite and its covering
of long thick setae are characteristic of the males of Utaperla and of
Paraperla frontalis (Banks). The variational tendencies in the number
of wing crossveins of both Utaperla and Paraperla are great enough to
have overlapping ranges so that no crossvein character can be applied
consistently enough to distinguish these genera from each other. Gaufin,
et al. (1966) use in their key to the genera of Utah Chloroperlidae their
observation that P. frontalis possesses three or more crossveins in the
costal area beyond the subcosta, as opposed to fewer than three in U.
sopladora. However, their own figure of the wing of P. frontalis shows
only two crossveins. During this study an examination of available
specimens of P. frontalis revealed that the number of crossveins in the
apical part of the costal area may range from two to four.
In addition to the above mentioned similarities between Utaperla and
Paraperla it should be mentioned that the coloration, especially the
abdominal patterning, of both of these genera is also very closely similar.
At the present time the following unique features appear best for
distinguishing Utaperla from the other genera in the Paraperlinae.
First, the length of the stem of the cranial Y-shaped ecdysial cleavage
line (eel) from the postoccipital suture (pocs) to the arms of the cleav-
age line is greatly reduced, less than one-fifth the length of each arm
extending to the ocellus. Unfortunately, the head and prothorax of the
only specimen of the new Chinese species of Utaperla had been broken
off. Thus, it is not known how similar the head structures of the North
American and the Chinese species of Utaperla are to each other. How-
ever, since both species greatly resemble each other in all observable
details, it is fairly certain that the head of this new species will be found
to be nearly identical to that of U. sopladora. Second, the mesothoracic
furcasternum (fig. 2) is divided along the mid- ventral line by a suture
(sut) which from the spinastemum extends anteriorly to lie between
the furcal pits, but it does not reach the sternacostal suture that extends
from one furcal pit to the other. Third, the metathoracic scutellar ridge
(fig. 3 scT) on each side of the insects mid-dorsal line is produced
meso-posteriorly into a strongly sclerotized somewhat digitate protrusion
bearing a sharply pointed “finger.” Fourth, the male supra-anal process
(fig. 6, 10) is apically bifurcate.
Utaperla sopladora Ricker
(Figs. 1-8)
1952, Utaperla sopladora Ricker, Indiana Univ. Publ., Sci. Ser., 18: 174-176, figs.
125 (male terminalia, dorsal), 126 (head and prothorax, dorsal), 127
JANUARY 1969] NELSON & HANSON — UTAPERLA REVISION
29
Figs. 4f-6. Utaperla sopladora Ricker. Fig. 4. Male terminalia, ventral view.
Fig. 5. Male terminalia, dorsal view. Fig. 6. Male terminalia, lateral viet^. (ap =
apical piece; bp = basal piece; ip = inner part.)
30
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
female terminalia, ventral) , 128 (male terminalia, lateral) , 129 (wings of
male holotype) , 130 (wings of female allotype) .
1955, Gaufin, Proc. Utah Acad. Sci., 32: 120.
1959, Jewett, Oreg. State Monogr., Stud. EntomoL, 3: 77, figs. 29 e (after Ricker
1952, fig. 125), 29 f (after Ricker 1952, fig. 127).
1962, Jewett, Pan-Pac. EntomoL, 38 (1) : 20.
1963, Ricker, Photo Offset, presented to third international symposium on
Plecoptera, pp. 14, 15, 19, and distribution map.
1964, Gaufin, Proc. Utah Acad. Sci., 41 (2) : 225.
1964, Gaufin, Gewasser Abwasser, 34/35: 39.
1964, Ricker, Gewasser Abwasser, 34/35: 55 and 68, fig. 11 (distribution map).
1965, Knight, Nebeker and Gaufin, EntomoL News, 76 (4) : 108, fig. 21 (egg,
lateral) .
1966, Nebeker and Gaufin, EntomoL News, 77 (2) : 42.
1966, lilies. Das Tierreich, 82: 428, fig. 19 (distribution map).
1966, Gaufin, Nebeker and Sessions, Univ. Utah Biol. Ser., 14 (1) : 12, 17, 19, 21,
72, 75, 77, figs. 2 (after Knight, Nebeker and Gaufin 1965, fig. 21), 203
(head, dorsal) , 235 (male terminalia, lateral) , 236 (female terminalia,
ventral), 238 (male terminalia, dorsal).
Ricker (1952) originally described and figured this species. Gaufin,
et al. (1966) recently refigured the head as well as the male and female
terminalia of JJ. sopladora and slightly paraphrased the text of Ricker’s
earlier work. Although both the description of Ricker and the figures
of all the above workers are adequate for the recognition of this species,
certain critical and fine structural details that might make this species
better known have heen overlooked. Therefore, it seems that further
comment on U. sopladora would be useful.
Male. — Length of the body varies from 5 to 7 mm. Supra-anal
process (fig. 5, 7) and its surrounding cowl are situated largely between
the tenth hemitergites. The process is composed of two major portions.
The very short “inner part” (Smith, 1917) is truncated at its anterior
margin and is held to the body by membrane from the tenth segment
(fig. 5 ip) . From its apex two very thin and sharply pointed lateral
braces arise one on each side (fig. 5). The “free part” (Smith, 1917)
viewed laterally is hook-shaped and is held to the body by membrane
from the cowl which attaches to its basal lateral sides. The free part
can be further divided into a single basal piece (bp) and two apical
pieces (ap). The hasal piece extends from the lateral braces and is a
long slender sclerite bearing a bluntly rounded apical margin. The two
apical pieces arise one on each side from the lateral apical margins of
the basal piece and each is a narrow sclerite which dorso-apically be-
comes slightly expanded. It is possible that the apical pieces may have
been derived from “lateral stylets” such as those associated with the
supra-anal process of P. frontalis. The cowl of the tenth segment is com-
JANUARY 1969] NELSON & HANSON UTAPERLA REVISION
31
Fig. 7. Utaperla sopladora Ricker, male supra-anal process, lateral view. Fig. 8.
Utaperla sopladora Ricker, female terminalia, ventral view. Fig. 9. Utaperla orien-
talis Nelson and Hanson, wings.
UI1IIS20
32
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
posed of membrane and two distinct paragenital plates. Ricker (1952)
mistakenly thought that the paragenital plates were the darkly sclerotized
inner mesal margins of the hemitergites. However, close examination
of this area reveals that these plates are separated from the true inner
mesal margin of each hemitergite by a thin strip of membrane.
Female. — Length of the body varies from 6 to 8 mm. Subgenital plate
(fig. 8) extending over the anteriormost portion of the ninth sternite,
with posterior margin convex. The specimens examined during this
study conform with Ricker’s figure of the allotype in having a slight
median emargination in the posterior margin of the subgenital plate.
No such emargination is shown in the figures of female terminalia of
U. sopladora by Gaufin, et al. (1966), thus this characteristic may be
variable.
Material Examined. — 10 males, 7 females, North fork of 12-mile creek, Steese
Highway, milepost 94, Alaska, 11 July 1952 (C. P. Alexander, M. E. Smith, D. L.
Carson) ; 1 male, Chatanika River, Steese Highway, milepost 39, Alaska, 10 July
1952 (D. L. Carson) ; 1 male, Alaska Highway, milepost 413, British Columbia,
Canada, 26 June 1954 (M. E. Smith) ; 1 male, Alaska Highway, milepost 430,
British Columbia, Canada, 27 July 1954 (C. P. Alexander) ; 1 male, Mendeltna
Creek, Glen Highway, Alaska, 8 August 1954 (C. P. Alexander). Of the preceding
material 5 males and 3 females of the “North fork” collection are in the C. H.
Nelson collection, the remaining material is in the J. F. Hanson collection.
Utaperla orientalis Nelson and Hanson, new species
(Figs. 9-12)
Only one specimen of this species, a male, is known. The important
differences by which U. orientalis can be distinguished from U. sopladora
concern the supra-anal process (see figs. 4-7, 10-12) : (1) the sharp
pointed lateral braces of U. orientalis are much the larger and more
distinct; (2) the basal piece of the hook-shaped free part is much the
broader and shorter in U. orientalis ; (3) the apical pieces of U. orientalis
are broad basally, become narrower apically and are more heavily
sclerotized as well as longer than those of U. sopladora. In all other
morphological details U. orientalis, with the possible exception of the
missing head and prothorax, is nearly identical to U. sopladora.
Holotype Male. — Yalu Station, altitude 2200 feet. Greater
Khingan Mountains, Manchuria, China, July 1940 (M. A. Weymarn) .
The type will be retained in the collection of the senior author.
Distribution of Utaperla
The present known distribution of U. sopladora is from Alaska to
Utah and within this range this species is closely associated with the
JANUARY 1969] NELSON & HANSON — UTAPERLA REVISION 33
Figs. 10-12. Utaperla orientalis Nelson and Hanson. Fig. 10. Male terminalia,
ventral view. FiG. 11. Male terminalia, dorsal view. Fig. 12. Male terminalia,
lateral view. (ap= apical piece; bp = basal piece; ip = inner part.)
34
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO, 1
Rocky Mountains. Ricker (1963, 1964) speculated that during the
Pleistocene glaciation JJ. sopladora was confined in the ice-free areas of
Yukon River Valley and as the ice melted this species moved southward
from its unglaciated refuge. Ricker considered the north to south move-
ment of U. sopladora more likely than a south to north one since this
species lacks close relatives in southwestern North America and becomes
rarer in the southern part of its range. The discovery of closely related
U. orientalis in the Greater Khingan Mountains of China lends addi-
tional support to Ricker’s conjecture concerning the dispersal of U.
sopladora.
Quite possibly, to expand slightly upon Ricker’s thinking, during the
Pleistocene glaciation the progenitor species of V. sopladora and V.
orientalis took refuge either in the unglaciated areas of northwestern
North America or those of northeastern Siberia. This ancestral species
subsequently divided, conceivably, into two populations, one of which,
U. orientalis, extended southward into Asia, and the other, U. sopladora,
extended southward in North America. However, any more secure
conclusion concerning the distribution and origin of Utaperla will have
to wait until more information on the range of both species, especially
that of TJ. orientalis, and on the probability of the existence of more
species belonging to this genus is available.
Acknowledgments
The authors would like to thank Mr. F. Christian Thompson and the
wife of the senior author, Elaine S. Nelson, for critically reading and
correcting this manuscript.
Literature Cited
Gaufin, a. R., a. V. Nebeker and J. Sessions. 1966. The Stoneflies (Plecoptera)
of Utah. Univ. Utah Biol. Ser., 14: 1-93.
Ricker, W. E. 1952. Systematic Studies in Plecoptera. Indiana Univ. Puhl. Sci.
Ser., 18: 1-200.
1963. Distribution of Canadian Stoneflies. Photo Offset (paper presented to
third international symposium on Plecoptera published in shorter form
as Ricker 1964) . 24 pp. 46 distribution maps.
1964. Distribution of Canadian Stoneflies. Gewasser Abwasser, 34/35: 50-71.
Smith, L. W. 1917. Studies of North American Plecoptera (Pteronarcidae and
Perlodini). Amer. Entomol. Soc. Trans., 43: 433-489.
JANUARY 1969] BRIGGS — A NEW PHALANGID FAMILY
35
A New Holarctic Family of Laniatorid Phalangids
(Opiliones)
Thomas S. Briggs
Galileo High School, Lux Laboratory, San Francisco
The taxonomy of phalangids of suborder Laniatores in America and
Europe has been impeded by the presence of a moderately large un-
described family on both continents. Many workers have been aware
of this group but its presentation into the literature is difficult without
considerable revision in adjacent families and reexamination of poorly
described Laniatorid phalangids from all parts of the world. This
problem has been particularly acute in the United States where several
abundant species which properly belong in this family have been in-
correctly placed in a genus that differs at the superfamily level. There-
fore, the designation of this new family is needed for most future studies
of American Laniatores lest past errors be perpetuated and perhaps
compounded.
This paper will assign the name Erebomastridae to the undescribed
family and will present the revisions and additions of genera and species
for known members of the family in the United States. European place-
ment in the family will be limited to two species in my possession, but
additional described species may belong here. The European members
of subfamily Tricommatinae in family Phalangodidae should be re-
examined. The new family differs from described families by having
a distally bifurcated single claw on the hind tarsi. It belongs in super-
family Travunoidea because musculature exists in the penes of males.
The only previous effort to place a member of this family in an ap-
propriate category was made by J. Hadzi (1935) in bis monumental
description of Cladonychium corii Hadzi (synonymized under Ereho-
master acanthina (Crosby and Bishop) in this paper). Among other
distinctive characters he noted that this species had bifurcated single
posterior claws and musculature in the penis. Family Triaenonychidae
Soerensen was the only member of Travunoidea described at this time,
so Hadzi established the subfamily Cladonychinae in Triaenonychidae
for his species. Unfortunately, this subfamily was not widely recognized.
Because Travunoidea has been subsequently subdivided on the basis of
the structure of the posterior claws, and typical Triaenonychids lack a
juvenile aroleum, I do not believe Erebomastridae can now be con-
sidered a subfamily of Triaenonychidae.
The known Ameriean Erebomastrids include established species from
eastern United States and a remarkable new species encountered in dense
The Pan-Pacific Entomologist 45: 35-50. January 1969
36
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
forests along the Oregon Coast. While some of these species have heen
described from caves, none are significantly troglohitic. The Oregon
species is larger than all United States Laniatorids except the Cosmetids.
It is found among dense fern and rhododendron litter in well established
spruce forests, often clinging to the undersurface of fallen bark.
The two European species studied differ at the generic level, but have
in common several structures including a shortened stem on the posterior
claws. It is difficult to distinguish these claws from those of certain
members of Phalangodidae in which the normally separate claws are
fused well before their point of attachment. However, by careful posi-
tioning under the compound microscope I have found a suture between
the fused claws of each such Phalangodid examined. In any case, exami-
nation of the claws of early juvenile instars reveals obvious differences
in structure and development. Phalangodid claws are embedded in a
central pad of soft tissue while Erebomastrid claws are firmly attached
to a central stem in each instar. The juvenile of at least one species of
Erebomastridae has a posterior claw with the characteristic six branches
of juvenile Triaenonychids, but they also possess an aroleum. Need-
less to say, many more juvenile examples are needed and should not
be overlooked by collectors.
Key to the Families of Suborder Laniatores of the World
1. Third and fourth tarsi of adults with two simple claws having separate points
of attachment or fused at proximal extreme, penis without muscles between
dorsal and ventral plates (Oncopodoidea) 2
Third and fourth tarsi of adults with a complex claw having a single point
of attachment, penis with muscles between dorsal and ventral plates
(Travunoidea) 7
2. Last two to four abdominal tergites moveable, the anterior tergites forming
an abbreviated scute 3
Only the last tergite (the anal plate) moveable, the remaining eight forming
a dorsal scute Oncopodidae Thorell, 1897 (Southeast Asia)
3. Last four abdominal tergites moveable 4
Last two abdominal tergites only are moveable
Paralolidae Kratochvil, Balat and Jaroslav, 1958 (Bulgaria)
4. Third and fourth tarsi of adults with a dorsal projection between the claws
(a pseudonychium) , fourth coxae broad 5
Third and fourth tarsi of adults without a pseudonychium (except for
Trionyxellinae of Assamiidae) , fourth coxae normal 6
5. Tibia of palpi compressed, armed with fine setiform spines
Cosmetidae Simon, 1879 (New World)
Tibia of palpus not compressed, armed with stout spines
Gonyleptidae Sundevall, 1833 (Central and South America)
6. Anterior margin of scute with three, five or seven prominent spines, palpi
weakly armed Assamiidae Soerensen, 1884 (Old World)
JANUARY 1969] BRIGGS — A NEW PHALANGID FAMILY
37
Anterior margin of scute without prominent spines, palpi armed with
elongate spines Phalangodidae Simon, 1879 (Cosmopolitan)
7. Third and fourth tarsi with a claw bearing a median prong, usually with
paired branches 8
Third and fourth tarsi with a bifurcate claw having no median prong
Erebomastridae new family (Europe and United States)
8. Claws of third and fourth tarsi with a broad median plate or thickened
prong having more than two branches 9
Claws of third and fourth tarsi with a continuous median prong, usually bear-
ing two branches Triaenonychidae Soerensen, 1886 (Old and New World)
9. Claws of third and fourth tarsi with two well developed branches that
approach length of median prong
Synthetonychidae Forster, 1954 (New Zealand)
Claws of third and fourth tarsi with pairs of small dorsolateral branches
which are shorter than median prong
Travuniidae Absolon and Kratochvil, 1932 (Europe)
Superfamily Travunoidea Absolon and Kratochvil
Erebomastridae Briggs, new family
Anterior margin of scute with recess above each chelicera bordered by a spur,
three spurs present on margin. Scute with five indistinct, undivided areas. Eye
tubercle well behind anterior margin of scute. Four moveable tergites on abdomen.
Fourth coxa only slightly broader than anterior coxae, spiracles exposed. Sternum
with slender anterior section joining depressed broadened posterior portion lying
between fourth coxae, not extending along posterior of fourth coxae.
Third and fourth coxae of adults with bifurcated claws having single stem
entering distal tarsal segment, stem and prongs of claws slender and tubular, stem
folds upward into distal tarsal segment. Calcaneus present on all metatarsi. Juve-
niles with aroleum and additional branches on claw during early instars.
Palpal tarsi with more than four prominent spines, palpal tibia spinose, palpi
robust.
Ovipositor with four distal divisions; one anterior, one posterior and two lateral.
Penis with articulated distal segment controlled by longitudinal muscles. Basal
and distal segments slender.
Type genus . — Erebomaster Cope.
Key to Genera of Erebomastridae
1. Claws of third and fourth tarsi with stem short, less than half length of
lateral prongs (Europe) 2
Claws of third and fourth tarsi with stem nearly equal in length to lateral
prongs (United States) 3
2. First tarsus with four segments Querilhacia (Lucas)
First tarsus with five or more segments Holoscotolemon Roewer
3. Prongs on hind claws meet in 180° opposition, male with distal swelling on
tibia of second leg Cryptomaster, new genus
Prongs on hind claws meet at angle near 90°, male without distal swelling
on tibia of second leg 4
4. Ovipositor with setose lateral lobes, penis cultriform Erebimaster Cope
38
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
/
^ 3
4
5 ' 6
Figs. 1-7. Holoscotolemon unicolor Roewer: Fig. 1. Dorsum. Fig. 2. Ventral
view of penis. Fig. 3. Lateral view of penis. Fig. 4. Ventral view of ovipositor.
Fig. 5. Lateral view of fourth claw. Fig. 6. Dorsal view of fourth claw. Fig. 7.
Lateral view of male.
Ovipositor with but one seta on lateral lobes, penis not cultriform
Theromaster, new genus
Genus Holoscotolemon Roewer
Holoscotolemon unicolor Roewer
(Figs. 1-7)
Holoscotolemon unicolor Roewer, 1915, Arch. Naturg., 81A: 5. Roewer, 1923, Die
Weberknechte der Erde, p. 102.
Male. — Total body length, 3.14 mm. Scute length, 2.60 mm. Length of eye
JANUARY 1969] BRIGGS — A NEW PHALANGID FAMILY
39
tubercle, 0.26 mm. Scute width, 2.00 mm. Length of second leg, 7.12 mm. Width
of eye tubercle, 0.38 mm.
Anterior margin of scute with deep recesses above chelicerae bordered laterally
by sharp projections; shoulders without tubercles. Scute yellow, mottled with
light brown pigment which decreases anteriorly allowing yellow background to
appear. Areas bordered by tubercles. Tergites yellow, with light brown pigmenta-
tion and lateral rows of small tubercles. Eye tubercle a low mound recessed from
anterior margin by space equal to its length. Eyes ringed by narrow circle of dark
pigment.
Sternum medially narrow, broadens slightly anteriorly and becomes depressed
posteriorly into a broad pentagonal zone. Maxillary processes from second coxa
large, separate and setose.
Chelicerae with smooth basal segments having socket-shaped juncture with
scute. Distal segments tuberculate.
Palpal tarsus with four prominent posterior lateral spines and two somewhat
reduced anterior lateral spines; tibia with seven lateral spines; patella with two
mesial spines; femur with one mesial spine, three dorsal spines and four ventral
spines; trochanter with one mesial spine; coxa with small spines.
Tarsal formula for male 7, more than 13, 4, 6. Distitarsi of first legs with three
segments, of second with three segments. Basal segment of third tarsus stout.
Tarsal claw of hind legs bifurcate, branches meet at angle near 90°. Stem pivots
upward into tarsal apex.
Penis a narrow stalk having articulated distal portion with arrowhead-shaped
apex bearing two setae. Musculature articulates distal portion of penis.
Female. — (Possibly not same species as male.) Similar to male but dorsal spines
on palpal tarsus much reduced, tarsal count 5 to 6, more than 13, 5, 6. Distitarsi
of first legs not well defined, tarsi of third legs normal.
Ovipositor with four distal projections as in fig. 4. Lateral projections bear
inwardly curved tips.
New records. — Austria. Karnten. One male, Sudhang der Koralpe bei St.
Vinzens (X1625), 14 August 1965, H. Franz; one female, Koralpen: Sudhang,
St. Lorenzen, Magdalensberg, August 1965?, H. Franz. W.-Steiermark. Two
females, St. Oswald ob Eibiswald, Mischwald nake Krummbach: graben, 850m.
(X1632), 21 August 1966, H. Franz. Deposited in the Naturhistorisches Museum,
Wien, Austria.
Genus Querilhacia Roewer
Querilhacia querilhaci (Lucas)
(Figs. 8-14)
Scotolemon querilhaci Lucas 1866 (non Simon 1872) , Ann. Soc. Entomol. Fr, ser.
4, 6: 216. Roewer, 1912, Arch. Naturg., 78A: 140. Roewer, 1923, Die Weberk-
nechte der Erde, p. 102.
Phalangodes querilhaci Simon, 1879, Arach. France, 7: 153. Simon, 1913, Arch.
Zool. Exp., 52: 382 .
Querilhacia querilhaci Roewer, 1935, Arch. Zool. Exp. Gen., 78: 12, 38. Dresco,
1959, Notes Biospeol., 7 : 81.
Male. — Total body length, 2.22 mm. Scute length, 1.69 mm. Length of eye
0.19 mm. Scute width, 1.80 mm. Length of second leg, 4.21 mm. Width of eye
tubercle, 0.34 mm.
40
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 1
Figs. 8-14. Querilhacia querilhaci (Lucas) : Fig. 8. Dorsum. Fig. 9. Lateral
view of penis. Fig. 10. Ventral view of penis. Fig. 11. Ventral view of ovipositor.
Fig. 12. Lateral view of fourth claw. Fig. 13. Dorsal view of fourth claw. Fig. 14.
Lateral view of male.
Anterior margin of scute with three prominent projections edging two socket
shaped recesses above chelicerae, thin dark line parallels edge of each recess. No
tubercles on shoulders. Scute smooth with areas poorly delineated with small spines.
Scute narrows behind eye tubercle. Tergites bordered by small tubercles. Eye
tubercle lightly tuberculate, low, with eyes widely separated. Tubercle recessed
from anterior margin of scute.
JANUARY 1969] BRIGGS — A NEW PHALANGID FAMILY
41
Sternum narrow with slight anterior broadening, depressed posteriorly. Maxillary
processes on second coxa prominent. Palpal coxa with two ventral projections.
Operculum small, rounded anteriorly and truncated posteriorly.
Chelicerae elongate, distal segment slightly roughened by small tubercles and
spines. Basal segment with prominent dorsal spine.
Palpal tarsus with four large posterior lateral spines and two smaller anterior
lateral spines; tibia with seven lateral spines; patella with two mesial spines;
femur with one mesial spine, four ventral spines and four dorsal spines; trochanter
with one mesial spine and coxa with small spines.
Tarsal formula 4, 8 to 9, 4, 5. Distitarsi of first legs with two segments, of second
with three segments. Tarsal claw of hind legs with short solid stem and two
elongate branches which meet at angle close to 90°. Stem retracts into apex of
tarsus by pivoting upward.
Penis slender, with articulated distal section having pointed apex bearing two
setae.
Female. — Similar to male. Ovipositor with four distal lobes, each lateral lobe
bearing inwardly curved tip.
Specimens examined. — France. Correze, three males and one female, C. Juberthie.
Deposited in the Naturhistorisches Museum, Wien, Austria.
Cryptomaster Briggs, new genus
Scute smooth, with segmental areas obliterated. Eye tubercle smooth, low mound
well separated from anterior margin of scute. Sternum with setae on center of
broad posterior plate. Operculum small.
Palpal tarsi with five prominent spines, two anterior pairs and one posterior
mesodorsal. First tarsus with five segments, second tarsus with more than eight
segments, third tarsus with less than six segments, fourth tarsus with more than
five segments. Distitarsus of first legs with two segments, of second with three
segments. Tibia of second leg with distal swelling. Tarsal claw of hind legs with
two branches, meeting at 180° on stout stem nearly equal in length to branches,
no medial spurs on branches.
Penis with cultriform distal appendage. Ovipositor with few setae on lateral lobes.
Juveniles with aroleum on posterior claws.
Type species. — Cryptomaster leviathan Briggs.
Cryptomaster leviathan Briggs, new species
(Figs. 15-25)
Male. — Total body length, 3.98 mm. Scute length, 3.18 mm. Scute width, 3.33
mm. Length of eye tubercle, 0.44 mm. Width of eye tubercle, 0.65 mm. Length
of second leg, 10.85 mm.
Anterior margin of scute with three prominent projections edging two socket
shaped recesses above chelicerae. One tubercle on each shoulder. Scute broad,
uniform, slightly granular, sparsely adorned with setae which poorly delineates
dorsal areas. Tergites sparsely bordered with small spines. Eye tubercle a low
mound separated by distance equal to its length from anterior margin of scute.
Eyes widely separated, bordered by black pigment and elevated above eye tubercle.
Sternum broadened anteriorly and broadly depressed posteriorly. Pentagonal
section between fourth coxae bears two setae. Posterior maxillary processes appear
42
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 1
Figs. 15-24. Cryptomaster leviathan Briggs: Fig. 15. Lateral view of male.
Fig. 16. Mesial view of palpal tarsus. Fig. 17. Dorsal view of fourth claw. Fig. 18.
Lateral view of fourth claw. Fig. 19. Distal portion of second tibia of male. Fig. 20.
Ventral view of penis. Fig. 21. Lateral view of penis. Fig. 22. Dorsum. Fig. 23.
Ventral view of fourth claw of juvenile. Fig. 24. Ventral view of ovipositor.
disconnected from second coxa and fused to sternum. Operculum small and
rounded.
Chelicerae with elongate basal segment and spinose distal segment.
Palpal tarsi with five prominent lateral spines, two ectoventral and three meso-
JANUARY 1969] BRIGGS — A NEW PHALANGID FAMILY
43
dorsal; tibia with five large and four small spines; patella with two mesial spines;
femur with three dorsal spines, one mesial spine and ventral row of spines with
prominent basal pair; trochanter with small mesial spine; coxa with small spines.
Tarsal formula 5, 11 to 15, 5, 6. Distitarsi of first legs with two segments, of
second with three segments. Tibia of second leg with distal swelling bearing en-
larged setae, with largest seta twisted. Tarsal claw on hind legs with two branches
meeting at 180° on solid stem equal in length to branches.
Penis with articulated distal section, broad in lateral view, narrow in ventral
view. Ventral plate slightly flared laterally at apex, with dorsal process extending
beyond apex of ventral plate.
Female. — Similar to male except tibia of first leg is normal. Ovipositor with four
distal lobes. Lateral lobes bear one to two setae.
Juvenile. — Instars at about half adult size lightly pigmented; posterior legs
possess prominent aroleum under bifurcated claw having ectomedial tooth on each
branch and ribbed central fiber supporting aroleum.
Holotype male, allotype female and eleven paratypes, 4.5 MILES SOUTH
Gold Beach, Curry County, Oregon, 29 January 1967, under fallen
bark and in litter in primary spruce forest, T. Briggs, V. Lee and K.
Horn. One juvenile, same data as holotype, 28 January 1967, T. Briggs.
Adults collected, same data as holotype, 19 June 1966, K. Horn, T.
Briggs and V. Lee. Deposited in the California Academy of Sciences.
Genus Erebomaster Cope
Erebomaster Cope, 1872, Amer. Natur., 6: 420. Roewer, 1923, Die Weberknechte
der Erde, p. 107.
Phalangodes (part) Simon, 1879, Arach. France, 7: 156. Goodnight, 1958, Proc.
Indiana Acad. Sci., 67 : 323.
Scotolemon (part) Banks, 1901, Amer. Natur., 35: 672.
Scute smooth, with segmental areas delineated by shallow grooves. Eye tubercle
well separated from anterior margin of scute and well elevated. Sternum with
narrow anterior and broad, depressed posterior. Maxillary processes from second
coxae large. Operculum small.
Palpal tarsus with three prominent pairs of spines. Sexual dimorphism in palpal
or cheliceral spination absent. First tarsus with four to six segments, second tarsus
with more than eight segments, third tarsus with less than five segments, fourth
tarsus with less than five segments. Tibia of second leg normal. Tarsal claw of
hind legs with two branches meeting at angle near 90° on tubular stem nearly
equal in length to branches. Juveniles with aroleum. Distitarsus of first leg with
two segments, of second leg with three segments.
Penis with cultriform distal appendage which lacks lateral setae. Ovipositor with
numerous setae on lateral lobes.
Note. — This genus has recently been synonymized with Phalangodes
Tellhampf, the type species of which is Phalangodes armata Tellkampf.
This species has been examined by me and clearly is a member of
Oncopodoidea.
Type species. — Erebomaster flavescens Cope.
44
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 1
Fig. 25. Sternum and opercular opening of Cryptomaster leviathan Briggs,
operculum removed.
Erebomaster flavescens Cope
(Figs. 26-32)
Erebomaster flavescens Cope, 1872, Amer. Natur., 6: 420. Roewer, 1923, Die
Weberknechte der Erde, p. 107.
Phalangodes flavescens Simon, 1879, Arach. France, 7 : 156. Packard, 1888, Mem.
Acad. Natur. Sci. Wash., 4: 52. Roewer, 1912, Arch. Naturg., 78A: 142. Good-
night, 1958, Proc. Indiana Acad. Sci., 67 : 323.
Scotolemon flavescens Banks, 1901, Amer. Natur., 35: 362.
Male. — Total body length, 2.00 mm. Scute length, 1.44 mm. Scute width,
JANUARY 1969] BRIGGS — A NEW PHALANGID FAMILY
45
Figs. 26-32. Erebomaster flavescens Cope: Fig. 26. Dorsum. Fig. 27. Lateral
view of penis. Fig. 28. Ventral view of penis. Fig. 29. Dorsal view of third claw.
Fig. 30. Lateral view of third claw. Fig. 31. Lateral view of male. Fig. 32. Lateral
view of ovipositor.
1.25 mm. Length of eye tubercle, 0.34 mm. Width of eye tubercle, 0.44 mm.
Length of second leg, 5.37 mm.
Anterior margin of scute with shallow recesses above chelicerce. No tubercles
on shoulders. Scute smooth with shallow grooves bounding areas. Little pigment
beneath light yellow chiton. Tergites smooth, with inconspicous setae. Eye tubercle
46
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
conical, separated by its length from anterior margin of scute. Eyes present at
base of cone. Sternum narrow anteriorly with a broad pentagonal posterior be-
tween fourth coxae. Maxillary processes from second coxae prominent, separate
and setose. Operculum setose, small, rounded anteriorly and truncated posteriorly.
Chelicerae elongate, basal segments smooth, anterior segments with distal tuber-
cles. Palpal tarsus with six major spines; tibia with seven major spines; patella
with two mesial and one ectal spine; femur with one mesial, four to five ventral
and four blunt dorsal spines; trochanter with one ventral spine. Ectoventral spines
of palpal tarsus in advance of mesodorsal spines.
Tarsal formula: 4, more than 8, 4, 4. Tarsal claw of hind legs with central
stem and two prongs about equal in length to stem which meet at angle near 90°,
no spurs on prongs.
Penis with short, articulated distal section on narrow stem which curves upward
to acute apex. Apical section with jagged, cultriform dorsum.
Female. — Similar to male. Ovipositor with four distal projections, ventral pro-
jection narrowed at tip, lateral projections rounded and bearing numerous elongate
setae.
Specimens examined. — Indiana. Crawford County. One male and one female.
Crater Room of Wyandotte Cave, 9 May 1948, C. and M. Goodnight. Deposited in
the American Museum of Natural History.
Erebomaster ACANTHINA (Crosby and Bishop) , (new combination)
(Figs. 33-41)
Phalangodes acanthina Crosby and Bishop, 1924, J. Elisha Mitchell Sci. Soc., 40:
11. Roewer, 1926, Abh. Naturwiss. Ver. Bremen, 26: 285. Goodnight and
Goodnight, 1942, Amer. Mus. Novitates, No. 1188.
Cladonychium corii Hadzi, 1935, Biologia Generalis, 11: 49-72.
Male. — Total body length, 1.69 mm. Scute length, 1.30 mm. Scute width,
1.27 mm. Length of eye tubercle, 0.22 mm. Width of eye tubercle, 0.28 mm.
Length of second leg, 4.80 mm.
Anterior margin of scute with recesses above chelicerae bordered laterally by
sharp projections. Shoulders smooth. Scute bears light brown mottling with circu-
lar spaces anteriorly and behind eye tubercle, smooth in texture with shallow
grooves bounding dorsal areas. Tergites with smooth margins, and brown mottling
becoming a uniform band posteriorly. Eye tubercle conical, eyes surrounded by
dark pigment, tubercle very lightly pigmented. Body reddish orange. Sternum
narrow, broadens posteriorly, with small pentagonal region on posterior area
which differs in surface texture from sternum and bears two setae. Maxillary
processes on second coxa directed prominently forward. Operculum small, sub-
triangular, and setose.
Chelicerae with smooth basal segment and tuberculate distal segment.
Palpi with three pairs of prominent spines on tarsus; two mesial spines on
patella; one mesial spine on trochanter; four to five dorsal spines on femur.
Tarsal formula: 4, 9 to 15, 4, 4. False articulations present on metatarsi of
posterior legs. Tarsal claw of hind legs with central stem and two branches about
equal in length to stem; each branch bears ventral spur near midpoint.
Penis with narrow stalk, articulated distal section narrower in ventral view than
in lateral view with sharp, jagged dorsal edge.
JANUARY 1969] BRIGGS — A NEW PHALANGID FAMILY
47
Figs. 33-41. Erebomaster acanthina (Crosby and Bishop) : Fig. 33. Dorsum.
Fig. 34. Ventral view of penis. Fig. 35. Lateral view of penis. Fig. 36. Ventral view
of ovipositor. Fig. 37. Dorsal view of fourth claw Fig. 38. Lateral view of fourth
claw. Fig. 39. Lateral view of fourth claw of juvenile. Fig. 40. Dorsal view of
fourth claw of juvenile. Fig. 41. Lateral view of male.
Female. — S imilar in structure to male. Ovipositor with four rounded distal
lobes, lateral lobes bear numerous setae.
Juvenile. — Instars less than half adult size lack pigment; claws of posterior
legs possess distal aroleum on central prong having three pairs of lateral branches.
Specimens examined. — North Carolina. Wake County. Three fe-
males and one male, Walnut Creek near Raleigh, 26 October 1923, S. C.
Bishop (Paratypes). Durham County. One male and one juvenile, 7 mi
S.W. Durham in beech-oak litter, 5 December 1965, N. Heshima and
P. Rubtzoff. Paratypes are in the American Museum of Natural History.
Notes. — Erebomaster acanthina differs from Erebomaster flavescens
in possessing a distinct medial spur on the prongs of the claws on the
hind tarsi as well differing in several minor features. Hadzi (1935)
describes specimens of Erebomaster acanthina having five or six seg-
48
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 1
Figs. 42-48. Theromaster brunnea (Banks) : Fig. 42. Dorsum. Fig. 43. Lateral
view of penis. Fig. 44. Ventral view of penis Fig. 45. Ventral view of ovipositor.
Fig. 46. Lateral view of male. Fig. 47. Dorsal view of fourth claw. Fig. 48. Lateral
view of fourth claw.
JANUARY 1969 ] BRIGGS A NEW PHALANGID FAMILY
49
ments on the first tarsi. Other workers have not noted these variations
and they were not present in the specimens examined hy me. Pending
further investigation these variations are acknowledged in the generic
description of Erebomaster.
Theromaster Briggs, new genus
Scute smooth, with segmental areas obliterated. Eye tubercle smooth, low
mound well separated from anterior margin of scute. Sternum with narrow anterior
and broad, depressed posterior. Operculum broad.
Palpal tarsus with three prominent pairs of spines. Palpi and chelicerae with
sexual dimorphism in spination. First tarsus with four segments, second with seven
to nine segments, third and fourth with less than five segments. Distitarsus of first
legs with two segments of second legs with three segments. Tibia of second leg
normal. Tarsal claw of hind legs with two branches meeting at angle near 90° on
tubular stem nearly equal in length to branches.
Penis with setose, blunt distal appendage. Ovipositor with one or no setae at
apex of lateral lobes.
Type species. — Theromaster brunnea (Banks).
Notes. — Theromaster differs significantly from Erebomaster in the
lack of demarked dorsal areas on the scute, sexual dimorphism and
genital structure. It differs from Cryptomaster in segmentation on the
first tarsi, structure of the posterior claws, sexual dimorphism and struc-
ture of the penis.
Theromaster brunnea (Banks), (new combination)
(Figs. 42-48)
Scotolemon brunnea Banks, 1902, J. New York Entomol. Soc., 10: 142.
Phalangodes brunnea Roewer, 1912, Arch. Naturg., 78A: 142. Roewer, 1923, Die
Weberknechte der Erde, p. 105.
Male. — Total body length, 2.34 mm. Scute length, 1.57 mm. Length of eye
tubercle, 0.23 mm. Scute width, 1.52 mm. Length of second leg, 4.95 mm. Width
of eye tubercle, 0.36 mm.
Anterior margin of scute with recesses above chelicerae bordered laterally by
sharp projections. Shoulders smooth. Scute with numerous small tubercles. Color
dark, with brown mottling beneath surface of chiton which diminishes anteriorly
Tergites with surface similar to that of scute, with posterior tubercles. Eye tubercle
rounded mound with eyes widely separated. Eyes ringed with black pigment.
Area between eyes with less brown mottling than area posterior to eyes. Dorsal
areas scarcely outlined by small tubercles. Sternum narrowed anteriorly, broadened
and depressed posteriorly. Maxillary processes from second coxa truncate and
meet along their entire length. Operculum overlaps slightly with fourth coxa;
rounded anteriorly, slightly rounded posteriorly and bears setae.
Chelicerae with smooth basal segment, two elongate anterior tubercles and
swollen movable finger on distal segment.
Palpal tarsus with six prominent lateral spines, two anterior ectoventral spines
fused at their bases; terminal spine of tarsus bears posteriorly directed spur which
50
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
prevents spine from being extended forward. Femur of palpus with dense row of
reduced ventral spines.
Tarsal formula: 4, 8 to 9, 4, 4. Distitarsi of first legs with two segments, of
second with three segments. Tibia of first leg normal. Tarsal claw of hind legs
with central stem and two branches about equal to length to stem; each branch
bears small ventral spur near midpoint.
Penis with narrow basal segment. Ventral plate of distal segment with two to
six prominent setae and two lateral projections near apex which arch beyond
blunt apex.
Female. — Similar to male except palpi and chelicerae normal. Anterior ecto-
ventral spines on palpal tarsus not fused at base, terminal spine of tarsus without
posterior spur, femur with four elongate ventral spines; chelicerae with normal
movable finger and without two elongate anterior tubercles. Ovipositor with pointed
lateral lobes with or without apical setae.
Specimens examined. — North Carolina. Watauga County. Two
males and one female, Blowing Rock, 10 October 1923, Crosby and
Bishop. Buncombe County. Four females. Montreat, 16 October 1923,
S. C. Bishop. Alabama. Jackson County. One male, McFarland Cave
near Garth, 22 August 1966, K. Horn. North Carolina specimens de-
posited in the American Museum of Natural History, Alabama specimen
deposited in the California Academy of Sciences.
Notes. — Phalangodes archeri Goodnight and Goodnight (1942) was
not examined by me, but is likely a member of Theromaster on the basis
of iheir description.
Acknowledgments
I wish to express my gratitude to Dr. W. J. Gertsch of the American
Museum of Natural History for his efforts in securing essential material
from Eastern United States and to Mr. Jurgen Gruber of the Naturhis-
torisches Museum, Wein, Austria for his systematic advice and loan of
European material. Art work was performed by Jack Tom and Kevin
Horn.
Literature Cited
Goodnight, C. J. and M. L. Goodnight. 1942. New Phalangodidae (Phalangida)
from the United States. Amer. Mus. Novitates, No. 1188.
Hadzi, j. 1935. Ein eigentumlicher neuer Hohlen-Opilionid aus Nord-America,
Cladonychium corii g.n. sp. n. Biologia Generalis, 11: 49-72.
JANUARY 1969] ALEXANDER — NEW CRANE FLY SUBFAMILY
51
A New Subfamily of Winter Crane Flies
(Diptera ; Trichoceridae)
Charles P. Alexander
Amherst, Massachusetts
The Trichoceridae, or so-called winter crane flies, include five genera
with approximately one hundred species, the great majority being in
the typical genus Trichocera. One of these genera, Kawasemyia Alex-
ander, is so different from the others that a new subfamily, the Kawase-
myinae, is here proposed for it.
The four genera retained in the older subfamily Trichocerinae are
T richocera Meigen, 1803, which is chiefly holarctic in distribution, with
slight extensions southward into the oriental and neotropical regions,
and with certain species that have been transported evidently through
commerce to certain remote islands in the southern hemisphere. One
of these, T. regelationis (Linnaeus) was placed in a supposed new genus,
T richocerodes, by Brethes, 1925 from the South Georgia and Falkland
islands in the south Atlantic, while another, T. maculipennis Meigen,
similarly was described in a supposed new genus, Palaeopetaurista, by
Seguy, 1940, from the Kerguelen islands in the Indian ocean. The sec-
ond genus is Diazosma Bergroth, 1913, with a single species that is
widespread throughout the Holarctic region. Paracladura Brunetti,
1911, with relatively few species, has a very discontinuous distribution
that includes the oriental and eastern palaearctic regions, western North
America, southern South America, eastern Australia and New Zealand.
The fourth genus, N othotrichocera Alexander, 1926, has a few species
in eastern Australia and the subantarctic islands of New Zealand. The
outstanding present student of the family is Dr. Christine Dahl, of the
Zoological Institute of the University of Lund, Sweden (Dahl, 1966,
1967).
Dr. Masaaki Tokunaga (1935) described as Alfredia imanishii, a
remarkable subapterous Trichocerid fly from the Japanese Alps. The
following year Tokunaga (1936) placed this species in the genus Tri-
chocera based on the female sex, the male being unknown to him at that
time. The discovery of this sex in 1935 by Mr. Eiji Kawase and its
later study by Alexander (1952) showed that in wing venation and
structure the fly was entirely different from all other members of the
family and a new genus Kawasemyia was proposed for it. More recently
Dr. Kintaro Baba secured further materials of both sexes and the
fly is known satisfactorily (Alexander, 1956: 79-80). The range of
The Pan-Pacific Entomologist 45: 51-53. January 1969
52
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 1
SCi
Fig. 1. Venation of Trichocera hiemalis (De Geer). Figs. 2, 3. Venation of
Kawasemyia imanishii (Tokunga). Venational symbols: A, Anal; Cu, Cubitus;
M, Media; R, Radius; Sc, Subcosta.
Kawasemyia imanishii (Tokunaga) in Japan as known to me is re-
stricted to the province of Echigo in Honshu but undoubtedly it will be
found to have a wider range. From analogous distribution patterns of
certain other Diptera, the genus might conceivably be found to occur in
western North America.
As has been emphasized in earlier papers, the general features are
much as in Trichocera, including the antennae, legs, and genitalia of
both sexes. The subapterous condition of the female is unique in the
family while the wing structure and venation precludes its inclusion in
the Trichocerinae. The venation of Trichocera and Kawasemyia are
shown herewith for comparison.
Trichocera, and all other genera in the Trichocerinae (fig. 1) show
the venation with twelve veins that attain the margin, with R and M
each with four branches, and with two complete Anal veins. Vein Sc is
elongate, with Sc 2 far retracted, vein R 2 always preserved, and with the
prearcular field short but with all elements of venation preserved and
distinct. Kawasemyia (figs. 2, 3), the only genus in the Kawasemyinae,
has only seven veins attaining the margin, R with three branches,
M with two branches, and without Anal veins. Vein Sc is short, with
JANUARY 1969] ALEXANDER — NEW CRANE FLY SUBFAMILY
53
Sc 2 close to the tip of Sci, vein R 2 lacking, and the prearcular field
greatly reduced. There evidently is much variation in the venation, as
shown, for example, by the figure provided earlier (Alexander, 1952:
15) as contrasted with the two provided at this time. The short series
of males available to me appears to indicate that the normal venation is
about as in fig. 2. In fig. 3 cell Rs shows an adventitious vein. Also of
note in the wing are the elongate marginal trichia that completely en-
circle the wing, these being very long and sparse, widely spaced and in a
single row, being of approximately equal length on both the costal and
posterior margins. In the Trichocerinae the costal trichia are short and
very numerous and in several rows, those of the posterior margin longer
and more delicate, approaching the condition found in Kawasemyia but
more numerous and crowded.
Additional important references include: Alexander (1927), Alexan-
der and Alexander (1967), Esaki (1950), and Tokunaga (1938).
Literature Cited
Alexander, C. P. 1927. The Trichoceridae of Australia (Diptera). Proc.
Linnean Soc. N. S. W., 51: 299-304, 11 figs.
1952. A new genus of Trichoceridae from Japan (Diptera). Trans. Shikoku
Entomol. Soc., 3: 14^16, 1 fig. {Kawasemyia).
1956. Undescribed species of Nematocerous Diptera. Part IV. Bull. Brook-
lyn Entomol. Soc., 51: 75-81.
Alexander, C. P. and M. M. Alexander. 1967. A catalogue of the Diptera of
the Americas south of the United States (Sao Paulo). Ease. 3. Family
Trichoceridae, pp. 1-4.
Dahl, C. 1966. Notes on the taxonomy and distribution of Swedish Trichoceridae
(Dipt., Nemat.). Opuscula Entomol., 31: 93-118, 72 figs.
1967. Notes on the taxonomy and distribution of Arctic and Subarctic Tricho-
ceridae (Dipt. Nem.) from Canada, Alaska and Greenland. Opuscula
Entomol., 32: 49-78, 83 figs.
Esaki, T., et al. 1950. Iconographia Insectorum Japonicorum, Ed. 2: 1-1738,
4937 figs., 15 col. plates (reference p. 1512, fig. 4337; adult 2, as
Trichocera ) .
Tokunaga, M. 1935. Three new crane-flies from Japan. Annot. Zool. Jap., 15:
194-198, 6 figs., 2 plates (pi. 14, fig. 2, adult 2, as Alfredia imanishii) .
1936. Three marine crane-flies from Japan. Annot. Zool. Jap., 15: 460-468,
18 figs.; Appendix, p. 468 (as Trichocera imanishii).
1938. New or little-known Trichoceridae from Japan (Diptera). Tenthredo,
2: 137-148, 2 pis., with 31 figs.
54
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Horn’s Bruchidae Type-Material in the Ulke Collection
(Coleoptera)
Clarence D. Johnson
Northern Arizona University, Flagstaff
As pointed out by Bottimer (1968) and Johnson (1968), problems
exist concerning the location of type-specimens of G. H. Horn’s bruchid
species. Both Bottimer and Johnson designated lectotypes and indicated
that specimens forming the basis for Horn’s species descriptions were
deposited in the LeConte Collection, Harvard University, Cambridge,
Mass. (MCZ) ; The Academy of Natural Sciences, Philadelphia, Pa.
(ANS) ; The California Academy of Sciences, San Francisco, Calif.;
and the Ulke Collection, Carnegie Museum, Pittsburgh, Pa. This paper
is an attempt to clarify further where bruchid types are located by de-
scribing Horn bruchid type-material in the Ulke Collection.
A study of the Bruchidae in the Ulke Collection revealed that it con-
tains specimens of 27 Horn species. Of these, one is the holotype of
Sennius discolor (Horn), one is perhaps a type of Cercidiestes ulkei
(Horn), others are possible syntypes of other Horn species and some
cannot be considered as types. According to G. E. Wallace, the small,
red, square or rectangular pieces of paper attached to some of the speci-
mens are possibly the specimens examined by Horn. These bits of paper,
the locality labels, the determination labels, and a comparison of the
specimens with the original descriptions were used in determining
whether the specimens were indeed types.
Horn Bruchid Type-Material in the Ulke Collection
(1) Below are the Horn specimens that merit special discussion to
clarify their status as type-material.
Acanthoscelides perforatus (Horn), 1873. Two specimens. One has
the labels “Va.” and ^^Bruchus perforatus Horn” attached to it. The
other has the labels “Md.” and ‘‘^perforatus H. type” attached to it.
Neither of these conform to Horn’s published type-locality of “Arizona”
and an “unknown locality” and neither can be considered as a type.
Cercidiestes ulkei (Horn), 1873. One specimen. Attached labels:
“Ariz.,” “red square,” “ulkei Horn.” From Horn’s original description
of this species it is not possible to determine whether more than one
specimen was described. As both this specimen and the specimen labeled
type and deposited in the MCZ (Johnson 1968) are members of the
same species, problems should not develop concerning species concepts.
The Pan-Pacific Entomologist 45: 54^56. January 1969
JANUARY 1969] JOHNSON HORN’s BRUCHIDAE TYPE-MATERIAL
55
Megacerus impiger (Horn), 1873. Two specimens. One has the
labels “Cal.,” “red square,” ‘‘‘‘Bruchus ramicornis Boh. = impiger Horn”
attached to it. This specimen and the specimens deposited in the MCZ
and ANS (Johnson 1968) should he examined by the first revisor of
the genus before a lectotype is selected. The other specimen is labeled
“Ariz.” and cannot be considered as a type as it is not from Horn’s
published type-locality.
Merohruchus julianus (Horn), 1894. One specimen. Attached labels :
“San Julio,” “Typical spec,” ‘‘^Bruchus julianus^ This specimen is
evidently a syntype.
Sennius discolor (Horn), 1873. One specimen. Attached labels:
“Tex,” “red square,” “Bruchus discolor Horn type” I consider this
specimen to be the holotype of this species as it conforms to Horn’s
original description and statement “One specimen from Texas in the
cabinet of Mr. Ulke.”
(2) Listed below in the genus to which they are presently assigned
are specimens of species that could be considered as syntypes because
they have both the correct type-locality label and a red, square or rectan-
gular label attached to them. (The publication date of the original
species description, if other than 1873, is in parentheses in the follow-
ing lists. )
Acanthoscelides alboscutellatus, A. aureolus, A. floridae, A. fratercu-
lus, A. longistilus, A. seminulum, Merohruchus placidus, Mimosestes
amicus, Sennius cruentatus. Stator pruininus and Stylantheus macro-
cerus.
(3) Specimens of the following species have labels attached to them
which indicate they are from the type-locality, and, for this reason may
be syntypes, but they do not have a red, square or rectangular label
attached to them.
Acanthoscelides calvus, A. exiguus and Sennius bivulneratus.
(4) The following species are represented by specimens but the
locality label is not that published by Horn and they cannot be con-
sidered as types.
Acanthoscelides bisignatus, A. distinguendus, Sennius nigrinus. Stator
limbatus, S. sordidus, Zabrotes ohliteratus (1885), Z. spectabilis (1885)
and Z. suhnitens (1885) .
Specimens of Species of Other Authors
Specimens of species of authors other than Horn (some with red,
square or rectangular labels attached to them) are pinned after the
following labels in the Ulke Collection.
56
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Bruchus pisi Linnaeus, B. mimus Say, B. scutellaris Fabricius, B.
quadrimaculatus Fabricius, B. discoideus Say, B. coryphae Olivier, B.
desertorum LeConte, B. pauperculus LeConte, B. prosopis LeConte, B.
uniformis LeConte, B. ohsoletus Say, B. hihisci Olivier and B. musculus
Say.
Acknowledgments
I wish to thank G. E. Wallace, Carnegie Museum, for lending me the
specimens for this study.
Financial assistance for this paper was provided by the Entomology
Research Division, U. S. Department of Agriculture, under Grant 12-14-
100-9187 (33).
Literature Cited
Bottimer, L. J. 1968. On the location of types of five species of Bruchidae with
notes on early American literature of Acanthoscelides ohtectus. Can.
EntomoL, 100 (3) : 284-9.
Horn, G. H. 1873. Revision of the Bruchidae of the United States. Trans.
Amer. EntomoL Soc., 4: 311-42.
1885. Contributions to the coleopterology of the United States. Ibid., 12:
128-62.
1894. The Coleoptera of Baja California. Proc. Calif. Acad. Sci., 4: 302^49.
Johnson, C. D. 1968. Bruchidae type-specimens deposited in United States
museums, with lectotype designations. Ann. EntomoL Soc. Amer., 61
(5) : 1266-72.
ZOOLOGICAL NOMENCLATURE: Announcement A. (n.s.) 82
Required six-month’s notice is given on the possible use of plenary powers by the
International Commission on Zoological Nomenclature in connection with the follow-
ing names listed by case number:
(see Bull. zool. Nomencl. 25, pt. 2/3, 27 September 1968) :
1838. Type-species for Oligolophus C. Koch, 1872 (Arachnida).
1843. Validation of Aphis gossypii Glover, 1877 (Insecta, Hemiptera).
1845. Type-species for Anoplius Dufour, 1834; Neotypes for Sphex niger Fabricius,
1775, and Sphex nigerrimus Scopoli, 1763 (Insecta, Hymenoptera) .
(see Bull. zool. Nomencl. 25, pt. 4/5, 17 January 1969) :
1852. Suppression of Siphocoryne angelicae del Guercio, 1911 (Insecta, Hemiptera).
1853. Validation of emendation to Hyposmocoma of Hyposmochoma Butler, 1881
Insecta: Lepidoptera) .
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, Acting Secretary to the International Commission on Zoological Nomen-
clature.
JANUARY 1969] BOHART NEW DUFOUREA FROM CALIFORNIA
57
A New Species of Dufourea from California
(Hymenoptera ; Halictidae)^
George E. Bohart
Entomology Research Division, Agr. Res. Serv., USD A, Logan, Utah
The following description of a new species of Dufourea is published
to make the name available to J. W. MacSwain of the University of
California for his study of bees pollinating the onagraceous genus
Clarkia.
Dufourea macswaini G. E. Bohart, new species
Holotype male. — Length 9 mm, length of anterior wing 6 mm; integument in-
tense black, shining, without distinct microscopic crenulation or tesselation;
pubescence of dorsum mostly dark, that of pleura, venter and legs, mostly pale;
body elongate. Head'. Face and antenna seen from in front as in fig 1, in profile
with eye four-fifths as broad as cheek; the segments proportionately broader
laterally; mouthparts in profile as in fig. 2, in dorsal view with galea slightly
over 3 times as long as broad, lacinia equal in length to hindtibia; pubescence
dark brown but mixed with long, pale hairs on vertex, dorsal margin of scape,
ventral portion of cheek; punctation of clypeus close and fine, elsewhere close
and moderately fine except a little sparser between scape and eye, quite sparse
below median ocellus, irregular in size and distribution between lateral ocellus
and margin of vertex. Thorax: Pronotal angles unusually produced (right-angled
in dorsal view, narrowly rounded in lateral view) ; scutum with punctures irregular
in size and distribution, averaging slightly more than one large puncture width
apart, sparser postmedially, those of scutellum sparse sublaterally, otherwise
moderately close, those of metanotum fine, close, uniform, those of pleura fine,
mostly very sparse, except absent ventrally on hypoepimeral area; forefemur a
little more than, and mid- and hindfemora a little less than twice as long as broad;
foretibia more than twice and hindtibia about three times as long as broad, the
latter longer than hindfemur (7:5.8) ; midtibial and outer hindtibial spurs as in
figs. 3, 4; tarsomere I of all legs slightly more than three times as long as broad,
that of hind leg gently bowed and with posterior hair fringe over twice as long
as width of segment; tarsomeres II-IV of foreleg, and II-III of mid- and hindlegs
broader than long; tarsomeres II-IV of hindleg produced posteroapically ; pubes-
cence of scutum dark, sparse, mostly short, that of pleura pale, sparse, longer;
wings uniformly brownish infuscated, the veins dark brown; venation as in fig. 5.
Propodeum: Sides, posterior face with pale, rather long, sparse pubescence; dorsal
enclosure carinate behind, coarsely striate throughout, with about 12 striae on
each side. Abdomen: Terga shining, with moderate-sized, deep punctures, close
basally, becoming sparse apically, absent on posterior impressed borders which
are dark piceous; pubescence of terga I-IV dark, short, semierect except each
with single row of short, stiff, white hairs basally; sterna without sublateral hook-
1 In cooperation with the Utah Agricultural Experiment Station.
The Pan-Pacific Entomologist 45: 57-59. January 1969
58
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
lets; sterna VI-VIII and genitalea as in figs. 6-13, the ventral tuft on sternum VI
with mixed dark and light hairs.
Female. — Differs from holotype in usual characters influenced by sex but other-
wise similar except as follows: Postocellar area more rounded, only slightly de-
pressed sublaterally, and only slightly over twice as long as an ocellar diameter;
facial punctures sparser between scape and eye (about two puncture diameters
apart) ; clypeus more convex, very sparsely punctured distally but coarsely, more
closely punctured basally; width of cheek subequal to that of eye in profile;
flagellomere VII unusually broad, at least twice as broad as long; pronotal angles
more rounded but unusually prominent for female; short spines on apical margins
of fore- and midtibiae numerous and well developed; tibial scopa light brown to
pale, suberect, unusually long, sparse, not at all concealing integument in any view;
dorsal fringe of hind tarsomere I light brown, slightly appressed, about half as
long as tarsomere; tarsomeres II-IV of all legs at least slightly longer than broad;
dorsal enclosure of propodeum more closely striate on median half; tergal pubes-
cence very short and sparse, anal fimbria dark brown; tergum III definitely broader
than others.
Holotype male, (deposited in Calif. Acad. Sci.) 4-8 miles north of
North Fork, Madera County, California, 20 June 1963, on Clarkia
dudleyana (Abrams) Macbr. (J. W. MacSwain). Paratypes: (Housed
at U. C. Berkeley, U. C. Riverside, Utah State U., Logan) four females
and one male, same locality and date as holotype, on Clarkia dudleyana
and C. purpurea (Curt.) A. Nels Macbr. (J. W. MacSwain, J. A. Chemsak,
and R. W. Thorp) ; one female, 1.5 miles northwest of Tollhouse, Fresno
Co., Calif., 20 June 1963, on Clarkia cylindrica (Jepson) Lewis & Lewis
( J. A. Chemsak) .
The type series is uniform. The single paratype male, like the females,
has closer median striations on the propodeum than has the holotype.
D. macswaini is probably related most closely to D. echinocacti Tim-
berlake but it differs in having darker integument, shorter, darker
pubescence, shorter antennae, less modified legs, less flattened face, and
no modified flagellar hairs on the male. Other species in the same group
include D. vernalis Timberlake and D. vandykei Bohart.
Fig. 1. Face and antenna of male in frontal view. Fig. 2. Mouthparts distal to
car do in lateral view. Fig. 3. Midtibial spur of male. Fig. 4. Outer hindtibial spur
of male. Fig. 5. Forewing of female. Fig. 6. Sixth abdominal sternum of male in
ventral view. Fig. 7. Sixth abdominal sternum of male in lateral view. Fig. 8.
Seventh abdominal sternum of male in dorsal view (left) and ventral view (right) .
Fig. 9. Seventh abdominal sternum of male in lateral view. Fig. 10. Eighth ab-
dominal sternum of male in ventral view (left) and dorsal view (right). Fig. 11.
Eighth abdominal sternum of male in lateral view. Fig. 12. Genital capsule of
male in ventral view (left) and dorsal view (right) . Fig. 13. Genital capsule of
male in lateral view.
JANUARY 1969] BOHART NEW DUFOUREA FROM CALIFORNIA
59
6
7
12
13
60
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
New Species of the Genus Eurytoma from California
(Hymenoptera : Eurytomidae)
Robert E. Bugbee
Department of Biology, Allegheny College, Meadville, Pennsylvania
The following two new species and one subspecies were included in a
large sample of specimens sent to me for determination by Mr. D.
Charles Dailey of Davis, California. The new material was collected and
reared by him from six species of cynipid galls occurring on oaks
[Quercus] in the central valley of California.
Eurytoma f oligalla Bugbee, new species
(Fig. 3)
Female. — Length 2.7 mm (average 2.9, range 2.3-4.0) . Black except for yellow
scape, light brown to brown wing veins, and femora and tibiae which may vary
from all orange yellow, to yellow on distal and proximal extremities only. Pile
on face silvery white. Abdomen 1.5 mm in length (average 1.5, range 1.1-1. 9) ;
shape, deeply oval to circular from lateral view; lateral compression moderate;
wide sixth tergum 0.62 mm (average 0.57, range 0.47-0.70) at widest point; lateral
surface of sixth with fine scaling that thins out dorsally, but extends over dorsal
surface covering up to anterior one-half of surface; ninth tergum, short, pointed,
and 0.13 mm in length (average 0.17, range 0.12-0.22) with small, oval yellow
cercus on either side; ninth tergum and ventral valves extend upward at about 45
degrees from horizontal axis of abdomen. Internal genitalia average 1.4 mm (1.1-
1.7) in length and 1.1 mm (0.90-1.3) in height; dorsal valves narrow and black
for horizontal length but turn dorsally, anteriorly, with ventral valves at approxi-
mately a right angle; stylet arch in vertical plane. Propodeum with wide median
furrow narrowing gradually ventrally; central carina complete to base of furrow
or limited to upper one-third to one-half; lateral surfaces irregularly ridged and
punctate giving rough surface. Lower face not striate. Antennal flagellum filiform
with first five segments truncate distally and longer than wide, and terminal unit
of three segments, with most proximal separated from two closely fused distal seg-
ments by shallow but distinct annulation. Marginal and postmarginal veins of
wings linear, marginal 0.27 mm (average 0.30, range 0.22-0.35) and postmarginal
0.25 mm (average 0.23, range 0.17-0.30) in length, stigmal club small and usually
rectangular.
Male. — Length 2.4 mm (average 1.8, range 1.5-2. 5). Pile on face silvery white.
Flagellum of antenna with five pedicellate, dorsally produced segments and termi-
nal unit of two closely fused segments. Legs with black infuscation on all femora
and tibiae, but occasionally front tibiae all yellow.
Types. — Holotype female and allotype are from Folsom Lake, Placer
County, California, emerged 20 and 24 May 1962 from Callirhytis
pomiformis (Ashm.), (bisex, generation), on Q. wislizeni. Paratypes
include 15 females and 65 males, from the same locality, host gall and
The Pan-Pacific Entomologist 45: 60-64. January 1969
JANUARY 1969] BUGBEE NEW EURYTOMA FROM CALIFORNIA
61
3
Fig. 1. Abdomen, lateral view, female, Eurytoma flavifacies Bugbee. Fig. 2.
Flagellum of antenna, male, Eurytoma flavifacies Bugbee. Fig. 3. Abdomen, lateral
view, female, Eurytoma foligalla Bugbee.
oak, but emergence dates ranged from 14 and 29 April, 1, 20, 22, 24,
and 27 to 29 May, 2 June 1962 and 21 February 1967. Types in the
United States National Museum, Washington, D. C. with paratypes in
the Bugbee collection, Meadville, Pennsylvania and in the University of
California at Davis. Additional paratypes include: Rocklin, Placer
County, California, 13 females, 7 males, emerged 5 December 1966,
16 and 22 April 1962 and 3 May 1962, from galls of Antron douglasii
(Ashm.) on Q. lobata, and 1 female, 3 males, emerged 12 and 16 April
1962, and November and December 1961 from galls of Disholcaspis
eldoradensis (Beuten.) on Q. lobata; Folsom Lake, Placer County, Cali-
fornia, 10 females, 4 males, emerged December 1961, 19 and 21 April,
and 27 May 1962 from galls of Antron echinus (Ashm.) on Q. douglasii,
and 2 females reared March 1962 from galls of Besbicus multipunctatus
(Beuten.) on Q. douglasii.
Hosts. — Callirhytis pomiformis, bisexual generation on Q. wislizeni;
Antron douglasii on Q. lobata-, A. echinus and Besbicus multipunctatus
on Q. douglasii and Disholcaspis eldoradensis on Q. lobata.
Range. — Known only from the Sierran foothills east of Sacramento,
California.
Remarks. — In the key to the females of the genus Eurytoma (Bugbee
62
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
1967) this new species would run to E. querciglobuli (Fitch). It differs
from E. querciglobuli, however, in averaging noticeably smaller in
practically all of its dimensions, especially the very short and stubby
ninth abdominal tergum. The sixth abdominal tergum is wider both
dorsally and laterally than the fifth, in contrast to the condition in E.
querciglobuli where the fifth and sixth are about equal in width
dorsally.
Eurytoma flavifacies Bugbee, new species
(Figs. 1 and 2)
Female. — Length 2.8 mm (average 3.0, range 2.4-3.4). Black except for yellow
tegula, scape, frons from base of scape, ventrolaterally to below compound eye
and surrounding a black spot just above clypeus, narrow band along anterior
and posterior margin of compound eyes, legs except for black spot on outer face
towards distal end of hind femur or some black may occur on base of fore femur
and in middle of mid and hind femur and sometimes on middle of mid tibia,
medial edge and base of fore coxa and mid coxa, exposed tips of ventral valves
and dark brown wing veins. Abdomen oval from lateral view and with slight lateral
compression so it appears quite plump; 1.6 mm in length (average 1.5, range 1.3-
1.7) ; wide sixth tergum 0.60 mm (average 0.62, range 0.57-0.67) at widest point;
sculpturing on lateral surface of sixth recedes towards anterior margin not reaching
dorsal surface; ninth tergum short and blunt, 0.10 nun (average 0.12, range 0.07-
0.15) in length. Abdominal petiole about half the length of the hind coxa. Internal
genitalia widely spread; wide dorsal valves are black along lower ventral margin
for about three-fourths horizontal length; stylet arch oblique; total length averages
1.5 mm (1. 4-1.6) and height averages 0.85 mm. Propodeum with shallow concavity
containing wide median furrow, furrow complete to base or indicated by lateral
Carina in upper one-third only; central carina distinct in upper one-third, and
below, surface crossed by irregular horizontal ridges; lateral areas narrow and
surface irregularly ridged. Head with wide, smooth, shiny area from below angle
of compound eye to base of mandible. Antennal flagellum six segmented; first
longer than wide; two to five about equal in length and width and moniliform;
sixth separated by shallow annulation from two closely fused terminal segments.
Wing veins with marginal wider than postmarginal; marginal 0.37 mm (average
0.35, range 0.32-0.40) and postmarginal 0.22 mm (average 0.17, range 0.15-0.22)
in length.
Male. — Length 2.0 mm (average 1.8, range 1. 3-2.0). Black except for yellow
scape, tegula, frons, and median surface of fore coxa; front legs may be all yellow,
or black infuscation on all femora and mid and hind tibia; mid coxa may be all
yellow, black, or combination of both. Flagellum with first four segments longer
than wide, dorsally raised, and pedicellate; fifth longer than wide and separated
from two closely fused terminal segments by deep annulation instead of short
pedicle Wing veins dark brown and relationships of lengths of veins about as
in female.
Types. — Holotype female and allotype from 5 miles s.w. of Madison,
Yolo County, California. Emerged 9 and 20 April 1967 from gall of
Andricus chrysolepidicola (Ashm.) on Q. douglasii. Paratypes 16 fe-
JANUARY 1969] BUGBEE — NEW EURYTOMA FROM CALIFORNIA
63
males and 7 males from same locality. Emerged 3 to 22 April 1967,
from same host gall maker and oak. Holotype and allotype in United
States National Museum, Washington, D. C. Paratypes in Bugbee col-
lection, Meadville, Pennsylvania and University of California at Davis.
Host. — From gall of Andricus chrysolepidicola (Ashm.) on Q.
douglasii.
Remarks. — This new species runs to Eurytoma flavovultus Bugbee
in the key to Eurytoma (Bugbee 1967). It may be easily distinguished
from E. flavovultus by the very short stubby ninth tergum, the dark
brown wing veins with the much wider marginal vein in relation to the
linear postmarginal and the dark brown antennae.
Eurytoma californica nana Bugbee, new subspecies
Female. — Length 2.7 mm (average 2.5, range 1.9-3.0). Black except for yellow
tegula and lower half of scape, yellow extremities of femora and tibia and brown
wing veins. Abdomen deeply oval in lateral view; 1.2 mm (average 1.3, range
1.2-1.4) in length; sculpturing covers lateral surface of sixth tergum but recedes
towards anterior margin on dorsal surface; ninth tergum very short and stubby,
0.07 mm (average 0.07, range 0.05-0.10) in length; ninth tergum and tips of
ventral valves extend dorsally forming angle of 45-50 degrees with the horizontal
axis of the abdomen. Internal genitalia average 1.2 mm in length and 1.0 mm in
height; narrow dorsal valves black for entire length and with ventral valves turning
dorsally, anteriorly, at approximately a right angle; stylet arch in a vertical plane.
Propodeum with shallow, wide, complete median furrow that narrows gradually
ventrally; central carina complete to base; lateral areas irregularly ridged, with
punctations between. Head showing a few weak striae converging on clypeus from
below and between compound eyes. Flagellum of antenna filliform and consisting
of five, longer than wide, distally truncate segments, and sixth segment separated
by distinct annulation from terminal unit of two closely fused segments. Black
on femora and tibia of all legs, sometimes present as a splotch on larger specimens
or more diffuse in smaller specimens. Wing veins with marginal wider and usually
twice as long as postmarginal; marginal 0.25 mm (average 0.27, range 0.25-0.30)
and postmarginal 0.12 mm (average 0.12, range 0.11-0.15) ; stigmal club approxi-
mately square.
Male. — Length 3.0 mm (average 2.0, range 1.5-3.0). Black except for yellow
tegula and lower one-half or one-third of scape; leg color quite variable but often
all femora and tibia with brown to black infuscation and only extremities yellow;
front and mid coxa may be yellow or only front coxa; less often front legs all
yellow and black present as spots on mid and hind femora and tibia. Antenna
with flagellum of five pedicellate, dorsally raised, longer than wide segments;
sixth segment separated from the terminal segment by deep annulation. Wing
veins brown and length relationships same as in the female.
Types.— Holotype female and allotype from Folsom Lake, Placer
County, California. Emerged 3 June and 24 May 1962 from galls of
Callirhytis pomiformis (bisexual generation), on Q. wislizeni. Para-
64
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO, 1
types, 14 females and 6 males from same locality, emerged in March,
April and May 1962 from the same host gall on the same oak species.
Types in United States National Museum, Washington, D. C, Paratypes
in Bugbee collection, Meadville, Pennsylvania and University of Cali-
fornia at Davis.
Host. — Callirhytis pomiformis (Ashm.) (bisexual generation).
Remarks. — This new subspecies differs from Eurytoma californica
Ashmead chiefly in its average smaller dimensions in all of the char-
acteristics measured (total length, length of abdomen, sixth and ninth
terga, wing veins, and internal genitalia) . Wing veins are lighter brown
and the sculpturing on the surface of the broad sixth abdominal tergum
spreads onto the anterior dorsal surface.
Literature Cited
Bugbee, R. E. 1967. Revision of the chalcid wasps of the genus Eurytoma in
America north of Mexico. Proc. U. S. Nat. Mus., 118 (3533) : 433-552.
SCIENTIFIC NOTE
Occurrence of Labidura riparia (Pallas) in Baja California, Mexico
(Dermaptera: Labiduridae). — On 9 April 1968, the Riparian earwig, Labidura
riparia (Pallas) was collected near kilometer marker 100, highway number 5,
south of Mexicali, Baja California, Mexico, by William Clark and Gene Ralston.
Twenty specimens were collected about 10 a.m. The site was below the highway
on the east side at an elevation near sea level. There was a small puddle of water
here at the end of a highway culvert. The earwigs were collected around the
puddle where the soil was damp. All of the specimens were collected under flat
rocks. There was no vegetation in the immediate area.
No mention could be found in the literature of Labidura riparia being collected
in Baja California, Mexico.
Heifer (1963, How to Know the Grasshoppers, Cockroaches and Their Allies,
Brown, Dubuque, Iowa) states that Labidura riparia has been found from Texas
to North Carolina. Schlinger et al. (1959, Jour. Econ. Ent., 52(2) : 247) mentions
that this species is known from Alabama, Florida, Georgia, Texas, and Louisiana.
Schlinger et al. also reports that the first collection of Labidura riparia in Califor-
nia was at Calexico on 21 November 1952. Nutting (1960, Pan-Pac. Entomol.,
36: 203) documented several collection sites in Arizona for Labidura riparia. The
first collection was at Yuma on 24 July 1952.
The authors would like to thank Irving J. Cantrall for the identification of
Labidura riparia. Two specimens were sent to Dr. Cantrall, University of Michigan,
and the remaining specimens are in the collections of Clark and Ralston. —
William H. Clark and Gene L. Ralston University of Nevada, Reno.
JANUARY 1969]
PROCEEDINGS
65
PACIFIC COAST ENTOMOLOGICAL SOCIETY
K. S. Hagen
President
H. L. McKenzie
President-elect
M. S. Wasbauer
Secretary
P. H. Arnaud, Jr.
T reasurer
Fig. 1. Edwin R. Leach — Ninetieth Birthday.
66
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Edwin R. Leach — Ninetieth Birthday. — The Pacific Coast Entomological
Society extends Congratulations and Best Wishes to its Honored Member, Mr.
Edwin R. Leach of Piedmont, California, on his 90th Birthday. Both, at the Execu-
tive Committee meeting, and in the General Meeting held on 15 November 1968,
motions were made and unanimously passed to extend these greetings to Mr. Leach.
Mr. Leach was born in Vallejo, California, on 3 May 1878. His family moved to
Oakland in February 1886. It was in this city that he attended grammar and high
school. He is a graduate of the University of California, Berkeley. He majored in
the field of mining and received the Bachelor of Science degree in 1901.
Mr. Leach, a 52 year member of the Pacific Coast Entomological Society, joined
the Society in 1916. His friendship with the late F. W. Nunenmacher, “won him
over to insects” and led to his attending the annual field trip of the Society held on
7 May 1916, at Pilarcitas Lake, San Mateo County, California. On this occasion he
recalls meeting Dr. Edwin C. Van Dyke, Dr. F. E. Blaisdell and Mr. Lawrence
Reynolds. At the sixty-first meeting of 26 August 1916 he was proposed (by Mr.
Nunenmacher) and elected to membership in the growing Society. Even though Mr.
Leach was interested in Entomology and Natural History at an early age, it was not
until he joined the Society that he decided to specialize in Coleoptera, particularly
the Lucanidae and cetonid Scarabaeidae. At this time he acquired the Lawrence
Reynold collection which also contained the Carl Fuchs generic series of Coleoptera.
Testimony of this zeal in these groups of Coleoptera is his collection which is one
of the finest in the country.
Mr. Leach served as treasurer of the Pacific Coast Entomological Society from
1931 through 1942. He has been active on its committees and also interested in the
finances of the Society. He became a Life Member in 1943. At the one-hundred
and ninety-seventh meeting, held 17 January 1948, Mr. Leach was elected to Hon-
ored Membership, the Society’s highest award.
The Pacific Coast Entomological Society extends to Mr. Leach best wishes for
many years of continued good health and activity. — Paul H. Arnaud, Jr., California
Academy of Sciences, San Francisco.
Proceedings
Three Hundred and Sixteenth Meeting
The 316th meeting was held Friday, 16 February 1968 in the Morrison Auditorium
of the California academy of Sciences, Golden Gate Park, San Francisco, with
President Hagen presiding.
Members present (35) : R. P. Allen, P. H. Arnaud, Jr., J. D. Birchim, T. Briggs,
R. M. Brown, R. Brownlee, G. Buckingham, G. M. Buxton, L. E. Galtagirone, W. E.
Ferguson, W. Gagne, M. R. Gardner, A. R. and Anita Gillogly, G. A. Gorelick,
J. Guggolz, J. F. Gustafson, K. S. Hagen, K. Horn, D. S. Horning, Jr., C. R. Kovacic,
Jr., W. H. Lange, R. L. Langston, V. Lee, H. B. Leech, K. Lorenzen, P. Lum, A. R.
Molenke, J. A. Powell, D. C. Rentz, E. S. Ross, R. Schoeppner, R. 0. Schuster, T.
A. Sears, W. E. Simonds, R. W. Thorp, J. W. Tilden, M. S. Wasbauer, R. F. Wilkey.
Visitors present (21) : Karen Allen, R. C. Gardner, W. Goodman, Carol Horning,
G. Jeung, Pauline Lange, Shari Lewis, R. T. Miller, R. P. Papp, R. R. Pinger, Jr.,
Kathy Rentz, R. C. Riley, R. B. and M. G. Roberts, Suriya Sampunth, R. Laud,
Cathy Tassan, Joyce Thorp, B. Tilden, J. Tom, M. J. Wong.
JANUARY 1969]
PROCEEDINGS
67
The minutes of the meeting held 15 December 1967 were summarized.
Mr. Hugh B. Leech noted with profound regret the passing of the well-known
Hymenopterist, Francis X. Williams, on 16 December 1967 at Chula Vista, Cali-
fornia.
The following note was presented:
Prey hunting and courtship of Cyrtopogon vanduzeei Wil. & Mart. — Ob-
servations were made on the perch behavior of this small asilid at Blodgett Forest,
13 miles east of Georgetown, El Dorado County, where the flies were active along a
logging road in late June, 1967. Prey consisting of leafhoppers and small nema-
tocerous Diptera, was captured in flight, and was held on the beak during feeding.
Courtship followed a regular pattern in which a male followed a female in flight,
then perched behind her, oriented in the same direction, usually about 5 mm back.
The positioning was usually repeated several times by a male after flights by the
female. The sequence often progressed to one in which the male slowly approached
so as to rest his fore tarsi on the folded wings of the female. At this point the
female invariably flew off, terminating the association. Although mating pairs, in
typical tail-to-tail position, were observed, consumation of the courtship sequence
in mating was not seen. It is interesting that the same courtship pattern obtained
in a larger robberfly, C. montanus Loew, which was active at the same site, yet
several other species of Cyrtopogon have been reported to carry out courtship in
a face-to-face position (Wilcox and Martin, 1936, Ent. Amer., 16:4). Some members
of the genus have the legs ornamented in the male, and at least one of these species
is included in the latter behavioral group. The two Blodgett species have no obvious
secondary features in the male. (Determinations by J. Wilcox). — J. A. Powell,
University of California, Berkeley.
The principal speaker of the evening was Mr. G. M. Buxton, California Depart-
ment of Agriculture, Sacramento. His illustrated talk was entitled “Grasshopper
collecting in montane California.”
Three Hundred and Seventeenth Meeting
The 317th meeting was held Friday, 15 March 1968 in the Morrison Auditorium
of the California Academy of Sciences, Golden Gate Park, San Francisco. President
Hagen presided.
Members present (44) : R. P. Allen, R. Anderson, P. H. Arnaud, Jr., L. N.
Bell, F. L. Blanc, R. Briggs, J. S. Buckett, W. L. Chase, Jr., H. V. Daly, D. G.
Denning, Carol L. Duba, J. G. Edwards, D. P. Furman, M. R. Gardner, Anita and
A. R. Gillogly, K. S. Hagen, K. Horn, A. Jung, W. H. Lange, R. L. Langston, V. Lee,
H. B. Leech, P. Lum, H. Marensten, A. E. Michelbacher, W. W. Middlekauff, W. D.
Murray, P. A. Opler, J. E. Prine, Sister Eugenie Rohner, S. B. Ruth, F. Santana,
T. A. Schulz, Joanne Slansky, C. Slobodchikoff, R. E. Stecker, A. Stephen, J. W.
Tilden, W. H. Tyson, M. S. Wasbauer, R. E. Wheeler, Barbara Wilson, E. C. Zim-
merman.
Visitors present (38) : S. Anderson, H. A. Appleton, H. R. Appleton, Mrs. H.
Appleton, Connie Bell, R. Bruce, D. Byrne, P. Gammer, R. Chavez, Loretta Den-
ning, J. Doyen, L. L. Duba, Alice and Janie Edwards, Mr. and Mrs. F. Ennik, Kay
Furman, Priscilla Gallego, R. Gardner, W. Goodman, E. E. Grissell, Kent Hagen,
Harriette Jung, P. Jung, Paulene Lange, D. W. Lee, Mr. and Mrs. Manuel Marquis,
68
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Mr. and Mrs. D. A. Ramsey, Roberta Sears, Mr. and Mrs. D. A. Shrout, Jan Snyder,
Anne Sugintas, R. Tenaga, Hazel, Jan and Bruce Tilden, J. Tom, Eleanor Wheeler,
Mrs. E. C. Zimmerman.
The minutes of the meeting held 16 February 1968 were summarized.
Seven new members were elected; Steven B. Ruth, R. E. Rice, Richard W. Fin-
ger, Robert R. Finger, Brian S. Cheary, Don Calvert, James Robertson.
Dr. E. S. Ross introduced Dr. E. C. Zimmerman, a long time member of the So-
ciety, visiting from New Hampshire. Fresident Hagen pointed out that Dr. Zimmer-
man is the founder of the series “Insects of Hawaii.”
Mr. Hugh Leech announced some accessions to the Society’s historical file: A
plate of A. Boving’s original drawings, a small file of letters from M. R. Smith to
Thomas Cook, donated by Mrs. Cook, and some photographs donated by John
Thomas.
The principal speaker of the evening was Dr. E. S. Ross, California Academy of
Sciences. His illustrated lecture was entitled “An Entomologist’s View of South-
west Africa.”
A social hour was held in the entomology rooms following the meeting. — M. S.
Wash AUER, Secretary.
Three Hundred and Eighteenth Meeting
The 318th meeting was held at 7 ;45 p.m. on Friday, 19 April 1968 in the Morrison
Auditorium of the California Academy of Sciences, Golden Gate Fark, San Fran-
cisco. Fresident Hagen presided.
Members present (36) : R. L. Adlakha, F. H. Arnaud, Jr., T. S. Briggs, R. G.
Brownlee, J. S. Buckett, G. R. Buckingham, D. Calvert, H. V. Daly, T. W. Davies,
B. J. Donovan, J. G. Edwards, W. E. Ferguson, W. Gagne, M. R. Gardner, E. E.
Grissell, J. Guggolz, J. F. Gustafson, K. S. Hagen, K. Horn, A. Jung, V. Lee, H. B.
Leech, F. Lum, R. E. Main, R. W. Finger, R. R. Finger, J. A. Fowell, S. B. Ruth,
F. J. Santana, R. 0. Schuster, T. A. Sears, Joanne E. Slansky, R. L. Tassan, R. W.
Thorp, J. S. Tilden, M. S. Wasbauer.
Visitors present (17) : Michele Benson, J. A. Caylor, Mary Davies, R. Gardner,
G. Jeung, H. Meyer, L. M. Middleman, Mrs. L. M. Middleman, R. B. Roberts, Bessie
D. Ruth, Suriya Sampunth, L. Stotelmyre, Joyce Thorp, Tina Vargas, S. C. Williams,
Charlene Williams.
The minutes of the meeting held 15 March 1968 was summarized.
Four new members were elected: Mr. Lee Middleman, Mr. Jule A. Caylor, Mr.
Robert Lame and Mr. David N. Ferro.
Mr. Hugh B. Leech showed Kissinger’s recently published monographic revision
of the curculionid subfamily Apioninae. This remarkable work contains over 2800
illustrations. It was published privately by the author and is available from him.
The following notes were presented:
New distribution records of two Nearctic Acalypta Westwood (Heterop-
tera: Tingidae). — A brachypterous $ of this primarily muscicolous genus of
tingids was collected from a berlese sample of a stream-side moss by Mr. G. L.
Rotramel. He had taken the sample from The Geysers, Sonoma Co., Calif., 13 April
1968. The species runs readily A. saundersi (Downes) in Drake and Lattin (Froc.
U. S. Nat. Mus. 115:334) where it is cited as feeding and breeding on moss. In the
1965 world catalogue of the family (Drake and Ruhoff, U. S. Nat. Mus. Bull. 243)
its distribution is listed as B. C., Wash., and Ore. This then constitutes a new
JANUARY 1969]
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69
state record and its most southern point. A search through the exhibitors material
revealed another, macropterous d running to A. barberi Drake from Thessalon,
Ontario, 10 June 1965, W. Gagne, taken in an insect pitfall trap from an aban-
doned pasture invaded by moss and Carex. Its host preferences are listed as moss
and hops. The species is previously recorded from B. C., Oreg., N. Y., and N. B.
This then constitutes a new provincial record. — Wayne C. Gagne.
In Memoriam — ^Elwood Wendell Molseed (5 August 1938 — 4 April
1967) . — Dr. Elwood Wendell Molseed, 29, Assistant Professor of Botany at the
University of San Francisco, died on 4 April 1967 of cancer, after an illness of
several months. He was a keen field botanist and one of those rare modern taxono-
mists with a green thumb. During his all-too-short career he made significant con-
tributions to our knowledge of Mexican and Central American Iridaceae. He did
undergraduate work at the University of San Francisco and received his degrees
at the University of California, Berkeley.
Dr. Molseed’s research interests centered in the family Iridaceae, particularly
the genera Tigridia and allies, such as Rigidella and Nemastylis, and their insect
pollinators. He made several extensive trips to Mexico, from Chihuahua to Chiapas,
to study these plants. His last field trip, assisted by three students, Luis F. Baptista,
Steven Kirchanski, and Robert Cruden, was in the summer of 1966. They collected
in areas along the main road from Durango to Mazatlan, the states of Nayarit,
Jaliseo, Colima, Michoacan, Puebla, Oaxaca, and Chiapas. The Kodachrome slides
here projected (loaned by Mr. Luis Baptista) include pictures of Tigridia areas in
Oaxaca, Nayarit, and Chiapas; the flower of Tigridia pavonia with a honey bee at
nectary; fields of Rigidella orthantha in the Sierra de Juarez, Oaxaca, at 7,000
feet elevation; flower of the red Rigidella orthantha with a new species of wasp
and a variegated morph of this species; and the flower of Tigridia dugesii var.
passif lor aides taken at Campastella, Nayarit. The southern portion of the trip
was to collect flowers and insect pollinators of Iridaceae, while the return trip
north was to collect seeds and corms.
Deposited with the Department of Entomology, California Academy of Sciences,
are approximately 830 specimens collected as insect pollinators or flower visitors
of Iridaceae. Each specimen is labeled with flower collection data. The majority
of specimens belong to the families of Apoidea, with various wasps, Coleoptera,
Diptera, and Hemiptera represented by smaller numbers. Regrettably, Dr. Molseed
did not have the time to publish on this planned phase of his research. — Paul H.
Arnaud, Jr., California Academy of Sciences, San Francisco.
A 1968 flight of Vanessa cardui Linnaeus in San Francisco and Pacifica,
California (Lepidoptera : Nymphalidae). — April 9, 1968 was Vanessa cardui
day in San Francisco and vicinity. This was an especially warm day with some
breeze. The official high temperature recorded by the weather bureau at Civic Cen-
ter in San Francisco was 78°F., with a light wind averaging 4 miles per hour from
the northeast at 1100 to 1200 hours and averaging 5 to 6 miles per hour from the
north at 1200 to 1600 hours. Telephone calls received by the Department of
Entomology, California Academy of Sciences, included those from local residents
as well as from newspapers in San Francisco and Pacifica. One at 1200 hours re-
ported a flight in the vicinity of Taraval Street and 30th Avenue, another at 1230
hours reported a flight at the stables between Skyline Boulevard and the Pacific
Ocean just south of the San Francisco city and county line flying from “southwest
70
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
to east of northeast into a warm breeze,” and another at 1320 hours reported a
flight in the vicinity of Ortega Street and 9th Avenue moving in a north easterly
direction. Earlier a lady had telephoned Mr. George Draper (reporter of the San
Francisco Chronicle) that butterflies were flying in the vicinity of Twin Peaks.
In the afternoon a reporter from the Pacifica Tribune reported butterflies in flight
in the coastal community of Pacifica.
The article in the Pacifica Tribune on April 10th, “Pacifica Goes Buggy with
Painted Ladies” contained much misquoted information. An article written by
Mr. Draper entitled “A Butterfly Invasion” and “ ‘Painted Lady’ Migration” ap-
peared in the April 10th San Francisco Chronicle. Because of general interest
among the citizenry, newspaper articles and television coverage was made of the
details of this flight.
In San Francisco, personal observations of the cardai flight were made with
the assistance of Mr. Dennis Murphy in Golden Gate Park, from the Morrison
Planetarium roof of the California Academy of Sciences. This roof overlooks to the
southwest the Academy court, and to the southeast Middle Drive. At 1315 hours,
89 cardui were counted to pass between the Academy buildings and Middle Drive,
in a one-minute period. The flight was oriented into the wind in a north-north-
west direction. In another one-minute period, at 1325 hours, 91 cardui flew through
the Academy court and over the Planetarium building. A total of ten specimens
(seven females and three males) were successfully netted as they flew over the top
edge of the building wall and over the roof. Cardui does not appear to be deterred
from flying up and over objects in its path of flight. The roof of the Planetarium
is about 40 feet above ground and numbers of specimens flew over the dome of
the Planetarium which is over fifty feet above the surrounding land surfaces. A
single male Monarch, Danaus plexippus Linnaeus, flew over the roof and was also
collected while flying with the cardui. On April 10th no further flight of cardui was
observed at the Academy. It was a cooler day and there was a coastal morning fog.
All times given are Pacific Standard Time. — Paul H. Arnaud, Jr., California
Academy of Sciences, San Francisco.
Occurrence of Cacoeciamorpha pronubana (Hbn.) in Oregon. — This
Palearctic tortricine moth has distinctive bright orange hindwings. Its presence in
Oregon (and in North America) was first noted three years ago when Kenneth
Goeden sent two female specimens which had been reared from pine in a Portland
nursery, in May, 1964. Establishment and spread of the species was confirmed this
past year, when specimens were received from Stanley G. Jewett, Jr. which had
been collected in August, 1967, near Elsie, Clatsop County, about 50 miles north-
west of Portland. The species is a polyphagous foliage feeder which is widespread
in the Palearctic and is known to be introduced in other areas, having first made its
appearance in England in 1905. — J. A. Powell, University of California, Berkeley.
President Hagen called for reports of the standing committees. Dr. H. V. Daly,
Chairman of the Membership Committee, made a motion that Drs. E. G. Linsley, A.
E. Michelbacher and R. L. Usinger be given honored member status in the Society.
It was further moved that an account of the contributions each of these men has
made to the Society be submitted to the Secretary for publication in the Proceed-
ings. Both motions passed unanimously.
Mr. J. S. Buckett, Chairman of the Program Committee, reminded the member-
ship that the Society’s annual picnic will be held at Russelmann Park on May 11.
JANUARY 1969]
PROCEEDINGS
71
Dr. J. F. Gustafson, Chairman of the Salt Marsh Committee, gave a report on his
studies on salt marsh habitats and made a plea for help in collecting and identifi-
cation of insects that occur in salt marsh habitats. He pointed out that perhaps a
third of the insects of this habitat are undescribed.
The principal speaker of the evening was Dr. Radcliffe B. Rorerts. His in-
formative talk was entitled, “Opportunites in Tropical Biology.”
A social hour was held in the entomology rooms following the meeting. — M. S.
Wasbauer, Secretary.
Three Hundred and Nineteenth Meeting
The three hundred and nineteenth meeting was the annual field day and picnic
and was held Saturday, 11 May 1968 at Russelmann Park, near Mount Diablo.
Members present (12) : P. H. Arnaud, Jr., H. V. Daly, J. Haddock, K. Hagen,
W. W. Middlekauff, A. Moldenke, P. A. Opler, R. E. Stinner, R. W. Thorp, M. S.
Wasbauer, P. Welles, R. F. Wilkey.
Visitors present (36) : J. Ball, D. Beaver, Barbara and Diane Daly, L. Edson,
Patricia Felch, Alan Garren, Steve and Donna Gary, Mr. and Mrs. Andy Gutierrez,
Helen Haddock, Maxine and Kent Hagen, G. M. and M. Hassell, J. Hendrickson,
Shari E. Lewis, Mrs. A. Moldenke, Chris, Sandra and Tim Opler, Terri Orr, R. P.
Papp, D. Sanders, Marilyn Sanders, D. Sehwartz, K. P. Shea, H. Stainbrook, Judy
Stinner, Y. Tanada, Joyce Kelly, Jeff and Katy Thorp, Patricia Welles.
The weather on picnic day was unsettled. The early morning was clear but later
there were clouds and some intermittent rain. The inclement weather seemed
not to dampen the spirits of those attending, and baseballs, bats and other sports
equipment were much in evidence. Between showers, some interesting insects were
collected, especially from the chaparral slopes. Very little swimming was noted,
however. — M. S. Wasbauer, Secretary.
Three Hundred and Twentieth Meeting
The 320th meeting was held Friday 13 September 1968 in tbe Morrison Audito-
rium of the California Academy of Seiences, Golden Gate Park, San Franeisco.
President Hagen presided.
Members present (31) : P. H. Arnaud, Jr., Mary F. Benson, F. L. Blanc, D.
Briggs, R. M. Brown, J. S. Buckett, G. M. Buxton, T. W. Davies, D. G. Denning,
J. G. Edwards, M. R. Gardner, J. Guggolz, J. Gustafson, K. S. Hagen, K. Horn, R.
L. Langston, V. Lee, H. B Leech, K. Lorenzen, P. Launibos, P. Lum, A. E.
Michelbacher, J. A. Powell, D. C. Rentz, E. S. Ross, R. 0. Schuster, Joanne
Slansky, C. Slobodchikoff, R. W. Thorp, J. W. Tilden, M. S. Wasbauer
Visitors present (18) : T. Bartolomei, R. Chavez, P. R. Grimstad, A. B.
Gurney, B. Hill, G. Julian, R. Lem, Martha Michelbacher, P. L. and Mary E. Nel-
son, R. P. Papp, Kathy Rentz, Joyce Thorp, B. Tilden, J. Tom, D. Ubick, L. Wehin,
M. J. Wong.
The minutes of the meeting held 19 April 1968 were summarized.
Thirteen new members were elected: Ian A. Boussy, R. L. Blickle, Annette
F. Braun, Robert W. Cruder, Eric M. Fisher, Melville H. Hatch, Stanley G.
McGregor, Theodore B. Mitchell, Charles H. Nelson, David R. Smith.
President Hagen announced the appearance of the Society’s third volume of
the Memoirs series entitled “Revisionary Studies in the Nearctic Decticinae,” by
72
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
David C. Rentz and James D. Birchim (173 pp. illus.). The publication sells for
$4.00 to the public and is available to Society members at a 20% discount. He
also pointed out that back issues of the Pan-Pacific Entomologist (vols. 1-16)
are available to the membership at $1.50 per volume. This is a large saving over
the regular price of back issues.
Dr. J. W. Tilden announced that the recent annual meeting of the Pacific Slope
Section of the Lepidopterists’ Society was held at the Berkeley campus of the
University of California September 6 to 8 and was well attended.
Mr. Hugh B. Leech noted with profound regret the passing of the well-known
Coleopterist, John Wagner Green on 20 June 1968 at Orinda, California.
Mr. J. S. Buckett, Program Committee Chairman, announced that the next meet-
ing of the Society is scheduled for 11 October and will consist of several short
papers. He suggested that those interested in participating contact him with their
suggested topic as soon as possible.
Dr. Joel Gustafson, Chairman of the Salt Marsh Study Committee, reported on
faunal surveys currently being conducted at the Audubon Sanctuary, Kent Island in
Bolinas Lagoon and some of the difficulties encountered in collecting insects in
this habitat.
The principal speaker of the evening was Dr. Ashley B. Gurney, U. S. National
Museum, Washington, D. C. His illustrated talk was entitled “Faunal Relation-
ships of Some Orthoptera Encountered During Field Experiences in the U. S. and
Abroad.”
A social hour was held in the entomology rooms following the meeting. —
M. S. Wasbauer, Secretary.
Three Hundred and Twenty-First Meeting
The 321st meeting was held Friday, 11 October 1968 in the Morrison Audito-
rium of the California Academy of Sciences, Golden Gate Park, San Francisco.
President Hagen presided.
Members present (40) : P. H. Arnaud, Jr., 1. A. Boussy, T. Briggs, R. M.
Brown, D. J. Burdick, G. M. Buxton, C. Dailey, T. W. Davies, R. L. Doutt, J. G.
Edwards, W. E. Ferguson, G. T. Ferrell, M. R. Gardner, G. A. Gorelick, J. Guggolz,
J. L. Gustafson, K. S. Hagen, K. Horn, D. H. Kistner, R. L. Langston, V. Lee, H.
B. Leech, K. Lorenzen, P. Lum, A. E. Michelbacher, A. R. Moldenke, P. A. Opler,
R. R. Pinger, Jr., J. A. Powell, D. C. Rentz, D. W. Ribble, S. B. Ruth, L. A. Ruud,
Jr., L. G. Santana, T. A. Sears, J. Shepard, C. N. Slobodchikoff, A. Stephen, J.
W. Tilden, M. S. Wasbauer.
Visitors present (48) : A M. Abrams, C. Adamson, Mellissa Boussy, J. G.
Brandon, P. Gammer, Carolyn Dailey, J. De Lana, E. Jimaeus, T. Dimock, J. F. Em-
mel, D. A. Evans, C. Ferrell, Helen Ferrell, S. Ferrell, R. C. Gardner, J. Geddes, C.
S. Glaser, Betty Guggolz, T. E. Hewton, Jr., J. Jones, P. Jung, N. M. Kindig, R.
Lem, D. Machado, Martha Michelbacher, Alison Moldenke, Sandra Opler, R. P.
Papp, J. M. Pasteels, R. Peterson, W. Rauscher, Kathy Rentz, Anne Rotramel,
Suriya Sampunth, Paula Schaller, Sigrid Shepard, Melissa Sheehan, W. E. Stewart,
Jr., Sharon Tanner, B. Tilden, J. Tom, D. Ubick, W. Velasquez, R. Vierhus, M. J.
Wong, W. Wright, W. L. Wright.
The minutes of the meeting held 13 September 1968 were summarized.
Eight new members were elected: Kenneth W. Cooper, Johannes L. Joos, Ray
JANUARY 1969]
PROCEEDINGS
73
Peterson, Billy H. Pettit, Michael H. Robinson, R. G. Rosenstiel, Alan R. Stephen,
Stanley C. Williams.
President Hagen announced with profound regret the death on 1 October 1968
of Dr. R. L. Usinger, an honored member of the Society. Memorial services will
be held at the First Congregational Church of Berkeley at 11 a.m. on Monday, 14
October 1968.
President Hagen announced that the New York Entomological Society is cele-
brating its 75th anniversary on 29 October 1968 with a membership dinner. He
noted that the Pacific Coast Entomological Society will be celebrating its 75th
anniversary in 1976.
Dr. W. E. Ferguson introduced Dr. William G. Evans, University of Alberta,
Edmonton, and Mr Louis Ruud, Arvin, California.
Dr. D. H. Kistner introduced Dr. Jacques M. Pasteels, visiting from Brussels
University.
Mr. Leech mentioned the newly formed Coleopterists’ Society, sparked by Dr.
Lee H. Herman, Jr., and open to all interested persons. Upon payment of dues,
members will receive the journal The Coleopterists’ Balletin, currently in its
twenty-second volume. Locally, people may contact Dr. J. Gordon Edwards, San
Jose State College. Full information may be had from Dr. Robert T. Allen, Ento-
mology Department, University of Arkansas, Fayetteville, Arkansas 72701.
Dr. P. H. Arnaud announced that volumes 1-16 of the Pan-Pacific Entomologist
are now available to the membership at $1.50 per volume.
President Hagen called for notes and exhibits. The following note was presented:
Cave adaptations in phalangids of genus Taracus (Opiliones: Ischyrop-
salidae). — Specimens of the genus Taracus taken from two widely separated caves
(Snake Cave in eastern Nevada and the White Chief Caves in the southern Sierra
Nevada Mountains) have similar external morphology. They both have entirely
black cephalothoraces and chelicerae with white or off-white abdomens, legs and
palpi. Their size is two to three times greater than known species of Taracus, all of
which are epigaeic. This increased size agrees with the usual directions of cavernic-
olous adaptation.
The White Chief specimens were discovered during a National Speleological
Society (San Francisco Bay Chapter) trip to the Mineral King area limestone caves
(Tulare Co., Calif.). They were encountered by Warren Rauscher, Bruce Rogers,
William Lum and I on the walls of passages in areas of total darkness or near
darkness. In spite of the high elevation of 11,000 ft, and the presence of snow
in the caves throughout the year, much life is evident in the caves. — Thomas S.
Briggs.
The principal speakers of the evening and their topics are given in the order
of their appearance:
J. A. Powell and W. J. Turner — Observations on oviposition behavior in
Orussus accidentalis . — Observations on Orussus occidentalis Cresson, in El Do-
rado County, California, during late June and early July, 1967, corroborate previous
findings on the behavior of 0. sayii Westwood, of the eastern United States. The
methods of patrolling (a means of reconnaissance), selection of oviposition sites,
and oviposition are exactly like those described for 0. sayii. The latter two
processes were observed about 20 times for 0. occidentalis. This species selected
bark-free areas on conifer logs of widely variable ages. The target of oviposition
74
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
was, in all four cases where it was traced, a frass-packed gallery of a wood-boring
insect, at distances from 59 to over 150 cm from the living occupant. The galleries
were those of Siricidae, Cerambycidae, and Buprestidae. One larva of 0. occiden-
talis was recovered from a target gallery where oviposition had been observed
15 days previously. The immature larva was found feeding on the distal abdominal
segments of a buprestid larva. The data suggest that suppositions of both
Burke (1918, Proc. Ent. Soc. Wash., 19:87) and Cooper (1953, Proc. Roches. Acad.
Sci., 10:38) were correct and not necessarily incompatible, i.e., oviposition and
possibly early instar feeding of Orussus occurs in frass-packed galleries, and
feeding by at least later instars takes place in the form of an external para-
sitoid on larvae and pupae of wood-boring insects.
G. Rotramel — Orientation and coupling in Dilophus orhatus (Diptera:
Bibionidae). — Dilophus orhatus Osten-Sacken (Det. confirmed by D. E. Hardy)
appeared in abundance from the turf in front of Wellman Hall on the Berkeley
campus this autumn. Daily observations revealed that copulation takes place in
flight with the male beneath the female. Male D. orhatus fly in a characteristic
pattern, dancing up and down. When there is a breeze, they head into it and use
swarm markers to maintain their positions. In still air they may be oriented in any
direction. Virgin female D. orhatus fly more slowly and tend to drift with pre-
vailing air currents. In the absence of a breeze, they fly shorter distances, fol-
lowing a straight path. Females have blackish wings and are distinguishable in
flight from males, which have transparent wings. This dimorphism may aid males
in visual discrimination of the sexes. The simple flight pattern of females
probably facilitates orientation and coupling by the males.
A male, which has recognized a female flying above him, makes an abrupt
approach from below and slightly to the rear. A male which recognizes a female
flying beneath him must drop below her before approaching. Striking modifica-
tions of the compound eyes and forecoxae are associated with orientation and copula-
tory behavior in the males. The eyes are composed of an upper turbaniform portion
which is made up of larger facets, and a lower portion which is set off by a band
of undifferentiated cuticle and consists of smaller facets. The coxal bases have an
enlarged membranous area which permits the males to fly with the forelegs
above their heads. The males appear to grasp the thorax of females before geni-
talic contact is achieved. I believe that each portion of the divided eye of the
male plays a special role in the sexual behavior of D. orhatus^ The lower portion
appears to assist in orienting to swarm markers on the ground and in recognizing
females at a distance. The upper portion of the eye apparently operates as a
wide angle scanning device allowing the male to home on the female flying above
him. I attempted to test this idea by presenting crude models of females to males in
flight, using elliptical black spots painted on white bristol board. When the models
were presented just beneath or slightly to on side of flying males, I frequently
obtained a response identical to the one elicited by a flying female.
P. A. Opler — Status of knowledge of the leaf-mining Lepidoptera of Cali-
fornia. — Tonight I would like to briefly summarize the current state of knowledge
about the leaf-mining Lepidoptera that occur in California. These creatures, which
include representatives of over twenty families of Lepidoptera in the state, are
known to feed on plants of at least forty-five families in California. This sizeable
fauna has been almost universally ignored by entomologists in general and lepidop-
JANUARY 1969]
PROCEEDINGS
75
terists in particular. These insects are very rarely collected as adults due to their
small size and the fact that they fly primarily at dusk in the near proximity of
their hosts. They are seldom attracted to light and even then they can be easily
overlooked on a light sheet.
Fortunately several features make leaf-mining Lepidoptera attractive to the
entomologist. These are as follows: 1) Many are beautifully colored and patterned
— ^the most beautiful lepidoptera in the state are leaf -miners; 2) their small
size makes the rearing of a large number of individuals a relatively easy task;
3) most species are quite specific in their host choice, thus making them ideal
subjects for host specificity, ecological or phyletic studies; 4) their biologies
are varied, often bizarre, and always intriguing; 5) most of the primitive groups
of Lepidoptera are leaf -miners; and finally 6) because these insects have been
little studied, one is not confronted with a ceaseless parade of literature muddles,
thereby keeping library hours to a minimum and field and laboratory hours
at a maximum.
My own research dealing with microlepidoptera associated with oaks has raised
the leaf-mining fauna on these hosts from twelve to more than fifty, a fourfold
increase ; approximately seventy per cent of these species are undescribed. A
similar ratio for the rest of the leaf-mining fauna in the state should prevail
when attention is eventually given to it.
J. F. Gustafson — Rearing observations on the madrona shield bearer Cop-
todisca arbutiella (Heliozelidae). — One hundred and thirty pupae of the
madrona shield bearer were collected during April of 1967. Observations were made
daily until 2 June 1967, and occasional observations were made thereafter. The
larva of this moth is a leaf-miner. Prior to reaching its full larval growth the cater-
pillar completely mines a small (2x3 mm) area of the leaf. The remaining upper
and lower epidermis are cut from the leaf and sewn together by the caterpillar.
This “shield” containing the caterpillar is lowered by a strand of silk toward
the ground, frequently interrupted in its descent by branches or other structures.
Pupation presumably occurs after the shield case is attached.
The moths emerge 22-34 days following “pupation.” The shield cases contain
strands of silk which protrude from the case and therefore the handling of these
tiny objects is made awkward because they stick to whatever object touches them.
Consequently 2 of the cases were lost during the examination period — probably
dropping off of a camel’s hair brush used to transfer the cases during a number
of observations. Cases were reared in units of 10 in small stoppered vials. Sixty-
four adult moths emerged — an emergence rate of 49%. Four parasitic wasps
also emerged during this period, for an emergence rate of 3%. One of the 4 wasps
was a chalcid, the other 3 were not identified. Wasps emerged 23-27 days fol-
lowing host pupation. At some time between October, 1967, and August of 1968,
31 additional chalcid wasps emerged. This raised the parasitism rate to 26.6%.
Of 21 of the host cases, examination of the contents disclosed: 7 were empty —
no evidence of the caterpillar; 10 semi-dehydrated moth larvae; 4 moth pupae —
dead. No evidence of parasites in these 21 were noted.
Several alternative interpretations are suggested to account for the delayed
emergence of the parasites. All of the suggestions were suppositions.
One point is made clear from this report. That is, if the combined emergence
of hosts and parasites is low in a rearing experiment, it is strongly suggested that
the specimens be kept for an additional year to permit possible later emergences.
76
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
A social hour was held in the entomology rooms following the meeting.- — M.S.
Wasbauer, Secretary.
Three Hundred and Twenty-Second Meeting
The 322nd meeting was held Friday 15 November 1968 in the Morrison Audito-
rium of the California Academy of Seiences, Golden Gate Park, San Franeisco, with
President Hagen presiding.
Members present (39) : R. P. Allen, P. H. Arnaud, Jr., N. Bell, F. L. Blanc,
G. Brady, D. L. Briggs, T. Briggs, R. M. Brown, R. G. Brownlee, J. S. Buckett,
T. W. Davies, D. G. Denning, M. R. Gardner, G. Gorelick, J. Guggolz, J. Haddock,
K. S. Hagen, K. Horn, W. H. Lange, R. L. Langston, V. Lee, H. B. Leech, C. D.
MacNeill, A. R. Moldenke, P. A. Opler, R. W. Pinger, R. R. Pinger, Jr., J. A.
Powell, D. C. Rentz, D. W. Ribble, E. S. Ross, T. A. Schulz, R. 0. Schuster,
Carolee Ann Start, R. E. Stecker, R. W. Thorp, M. S. Wasbauer, S. P. Welles, Jr.,
S. C. Williams, Barbara Wilson.
Visitors present (35) : Dr. and Mrs. A. J. Beattie, Connie Bell, J. Benet,
Michele Benson, P. Cammer, Renae Cottam, W. Davies, Loretta Denning, Mr. &
Mrs. John Doyen, J. Geddes, H. Gibbs, Mrs. D. A. Haddock, Mrs. J. D. Haddock, J.
Henrickson, A. Jung, R. Lem, Shari E. Lewis, T. Lucas, M. Marquis, Mrs. A. R.
Moldenke, S. R. Montague, M. Montgomery, Sandra Opler, R. P. Papp, W. Raus-
den, Kathy Rentz, Pam Robie, Suriya Sampunth, Barbara Subak, Joyee Thorp, J.
Tom, Darrell Ubick, Mrs. S. C. Williams, M. J. Wong.
The minutes of the meeting held 11 Oetober were summarized.
President Hagen announced with deep regret the death of Mr. H. L. McKenzie
on 17 October 1968 at Sacramento.
Letters of acceptance for honored member status by Drs. Usinger, Linsley and
Michelbacher were then read to the membership.
It was announced that the executive board has elected Dr. R. Miller as an hon-
ored member.
Dr. P. H. Arnaud, Jr. exliibited a photograph of Mr. E. R. Leach and made a
motion that it be published in a forthcoming issue of the Pan-Pacific Entomologist.
The motion passed by unanimous vote.
President Hagen announced the appointment of Dr. J. A. Chemsak as Chairman
of the Membership Committee to succeed Dr. Daly who is retiring from this post
at the end of the year, and his appointments of the Auditing and Nominating
Committees. These committees will be: Nominating: Dr. R. L. Doutt, Chairman,
Mr. H. B. Leech and Dr. W. E. Ferguson; Auditing: Mr. H. V. Davis, Chairman,
Dr. J. F. Gustafson and Mr. D. G. Denning.
Mr. H. B. Leech drew attention to part 5 of Lindroth’s series on “The ground
beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska” (Opuscula Entomo-
logica, supplements). Part 5 includes Amara, Harpalus, Selenophorus, Anisodacty-
lus, Dicheirus, Bradycellus, Stenolophus, Acupalpus, etc. As usual, there are keys
to species, descriptions with ecological and other data, and excellent illustra-
tions. There are almost 300 pages ; price 60 Swedish crowns.
Mr. Leech showed kodachrome slides of the larva, pupa and adult of the omniv-
orous looper, Sabulodes caberata Guenee, from a specimen feeding on a garden rose-
bush in San Francisco.
The following notes were presented:
Callophrys dumetorum (Bdv.) and Callophrys viridis (Edw.) in Califor-
JANUARY 1969]
PROCEEDINGS
77
nia. — A series of twelve color slides were shown of the adults and young stages of
Callophrys dumetomm and Callophrys viridis, two commonly-occurring Bay Area
bycaenid butterflies, characterized by the green scaling of the wing undersides.
The green color seen on the undersides of C. dumetorum appears grass-green in
most cases whereas the color of C. viridis is almost always dark green or blue-green.
The latter, found along the immediate coast from San Mateo County to Mendocino
County in California, oviposits on both Lotus scoparius and Eriogonum latifolium
latifolium (predominantly the latter). Callophrys dumetorum is more widely dis-
tributed, occurring primarily in the Coast Ranges and in the Sierra Nevada foot-
hills of California. Unlike C. viridis, however, it never occurs along the immediate
coast and oviposits mainly on Lotus scoparius. Although the two entities were
believed to be synonymous by many lepidopterists in past years, the segmentation
and color patterns seen in the larvae of both species differ from one another,
providing additional evidence that the two allopatric populations are sibling species.
— Glenn A. Gorelick, University of California, Berkeley.
Compsilura concinnata (Meigen) (Diptera: Tachinidae) reared from
Agraulis vanillae incarnata (Riley) (Lepidoptera : Nymphalidae) at San
Leandro, California. — Compsilura concinnata (Meigen) is one of the most widely
distributed species of Tachinidae. It was described from Europe in 1824, and is
known to occur naturally in Europe, Asia, Japan and South Africa. Compsilura
concinnata was first intentionally introduced into eastern North America in 1906 as
a parasite of gypsy moth larvae. Subsequently, it was liberated in the western
United States — in New Mexico, Arizona, California, Idaho, and Oregon. It is rarely
encountered in California collecting; there are no determined California specimens
of this species in the California Academy of Sciences or California Insect Survey
Collections.
In Eurpoe, according to Herting (1960, Biologic der westpalaarktischen
Raupenfliegen, Dipt., Tachinidae, Monog. z. Angeiv. Ent. 16:55-57) C. concinnata
is recorded as parasitizing about 100 different species of lepidopterous hosts
and a few Tenthredinoidea. In the United States and Canada it is recorded from
over 200 lepidopterous host species and also some Tenthredinoidea.
Fifty mature caterpillars of the Passion Flower butterfly, Agraulis vanillae
incarnata (Riley) were collected on Passiflora caerulea Linnaeus at San Leandro
Alameda County, California, in October, 1968. A mature larva of the tachinid
issued from the side of the chrysalis and was found as a puparium in the rearing
container on the 28th of October. The adult female emerged at some time during
the period of November 4th through 8th. Only this one female C. concinnata was
obtained, so parasitism of the Agraulis larvae was very low. — P. H. Arnaud, Jr.,
California Academy of Sciences, San Francisco.
The principal speakers of the evening were Ron Gardner and Terry Sears,
University of California, Davis. Their illustrated talk was entitled “Collecting
in the Grand Canyons of Northern Mexico.”
A social hour was held in the entomology rooms following the meeting. — M.S.
Wash AUER, Secretary.
Three Hundred and Twenty-Third Meeting
The 323rd meeting was held Friday, 20 December 1968 in the Morrison Audito-
rium of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Hagen presiding.
78
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
Members present (28) : R. J. Adams, P. H. Arnaud, Jr., J. Benet, F. L. Blanc,
1. Boussy, T. Briggs, P. Gammer, J. A. Caylor, D. G. Denning, J. G. Edwards, W.
E. Ferguson, M. R. Gardner, J. F. Gustafson, K. S. Hagen, C. R. Kovacic, Jr. V.
Lee, H. B. Leech, K. Lorenzen, P. Lum, R. E. Main, A. E. Michelbacher, D. W.
Ribble, E. S. Ross, S. D. Smith, Carolee Ann Start, R. W. Thorp, D. Ubick, M. S.
Wasbauer.
Visitors present (11) : C. Adamson, J. G. Brandon, Midori Caylor, R. C.
Gardner, W. C. Heath, M. Lim, W. Rauscher, G. Resnick, Joyce Thorp, B. Tilden,
T. D. White.
The minutes of the meeting held 15 November 1968 were summarized.
Twelve new members were elected: Richard J. Adams, Jason Benet, Ralph J.
Bushnell, Paul A. Gammer, Robert S. Erdmann, Paul R. Grimstad, Kenneth Groeppe,
Theodore W. Jones, David G. Marqua, Tom Newton, Gaylan Queirola, Russell A.
Rahn.
Two student members were elected: Richard Anderson and Darrell Ubick.
Mr. Leech exhibited two recent publications: Invertebrate Zoology, volume
2, by Alfred Kaestner (Translated by Herbert W. and Lorna R. Levi. Interscience
Publishers, John Wiley & Sons, Inc., New York; released November 11, 1968.
x-)-472 pp., 252 figs. $22.95. Treats of the Phyla Onychophora, Tarigrada, Penta-
stomida, Arthropoda, the last from the Triobites through Araneae to Symphyla,
with a final 3 pp. on Insecta.) ; and The carabid beetles of New Guinea Part III.
Harpalinae (continued) : Perigonini to Pseudomorphini, by P. J. Darlington, Jr.
Bulletin of the Museum of Comparative Zoology, vol. 137, no. 1, pp. 1-253, inch
186 figs, and 3 pis. in color. 30 July 1968.
The following notes were presented:
Salt marsh insects. — As part of the activity of the salt marsh habitat committee
of the Society, an insect survey of a local salt marsh has been initiated.
Standard methods of sampling insect populations have been employed. Of the
three major plants of the salt marsh community, cord grass, Spartina foliosa Trin.,
by far supports the highest insect populations.
It was our custom to make insect collections during periods of low tide.
Late in November, 1968, aerial net sweeping was conducted on an in-coming tide.
The rising waters drove the insects up high on the plants. Consequently, the
November collections show an apparent great increase in the populations of the
insects. Actually, we witnessed what must be a commonplace event in this habi-
tat. The respective populations of insects on each of the three major plants of
this salt marsh community, all demonstrated the same response — that of insects
being concentrated toward the tops of the plants by the rising tide.
We take this opportunity to express our great appreciation for assistance of
the classification of the salt marsh insects as indicated:
Dr. Stanley F. Bailey (Thysanoptera) ; U. C. Santa Cruz
Dr. A. C. Cole, Jr. (Formicidae) ; University of Tennessee
Dr. Frank Cole (Diptera ) ; U. C., Berkeley
Dr. Ralph H. Davidson (Cicadellidae) ; Ohio State University
Mr. Wayne C. Gagne (Miridae) ; Bishop Museum, Honolulu
Mr. Gail Grodhaus (Chironomidae) ; Calif. Dept, of Public Health, Bureau of
Vector Control, Berkeley
Mr. Hugh Leech (Coleoptera) ; Calif. Academy of Sciences, San Francisco
Dr. Edward L. Mockford (Psocoptera) ; Illinois State University
JANUARY 1969]
PROCEEDINGS
79
Mr. Lawrence J. Pinter (Salticidae) ; Museum of Comparative Zoology, Harvard
Univ.
Mr. John T. Polhemus (Saldidae) ; Englewood, Colorado
Dr. Jerry Powell (Lepidoptera) ; U.C., Berkeley
Dr. Edward S. Ross (Embioptera) ; Calif. Academy of Sciences, San Francisco
Mr. Curtis W. Sabrosky (Chloropidae) ; U.S. National Museum
Mr. David B. Scott (Collembola) ; Soilserv, Inc., Salinas, Calif.
Dr. Willis W. Wirth (Ephydridae) ; U.S. National Museum.
— Robert S. Lane and Joel F. Gustafson, San Francisco State College.
Occurrence of the genus Enlinia Aldrich in the western Nearctic (Dip-
tera: Dolichopodidae). — The genus Enlinia, which contains some of the smallest
Dolichopodidae, is known only from the Americas. There are three species pres-
ently known from the United States. The genus was described in 1932 by Aldrich.
The type species is magistri Aldrich, described from specimens collected in the
state of New York and the District of Columbia. Two additional species are
known from the southern United States. The new Enlinia displayed this evening,
collected at Redwood City, San Mateo County, California, on 21 March 1952, by
the author, is the first to be discovered in the western Nearctic. A series of
specimens was collected on the rock wall of a pond near the top of a serpentine
hillside. This species, at the present time, is known only from specimens collected
during California’s rainy season. — Paul H. Arnaud, Jr., California Academy of
Sciences, San Francisco.
President Hagen called for the 1968 reports from the chairmen of the stand-
ing committees. Dr. J. G. Edwards, Membership Committee, reported that 60 new
members were elected in 1968 and four members — E. G. Linsley, A. E. Michel-
bacher, R. C. Miller, and R. L. Usinger — were elected to honored status. Mr. Leech,
Chairman of the Historical Committee, reported on important acquisitions to the
Society’s historical files during 1968, which included in addition to those things
mentioned in previous meetings, original photographs by A. L. Melander; from E.
S. Ross, a large photograph of Dr. E. C. Van Dyke, Mr. J. W. Green and of Dr.
Ross himself; from E. C. Zimmerman, miscellaneous colored plates and figures
of insects and from the Department of Entomology of the Academy, the original
figures from a posthumous paper on the thrips of Australia by Dudley Moulton,
published this year by the Academy; and from the Secretary of our Society, various
records and papers, some having to do with the founding of the Pan-Pacific
Entomologist. The files should obtain a new lease on life in 1969; additional
rooms in the Entomology Department should allow them to be brought up from the
basement, spread out and sorted.
Dr. W. E. Ferguson gave a brief report of the Publication Committee and
mentioned that the first 16 volumes of the Pan-Pacific Entomologist are on sale
to members at very reduced prices. He also mentioned that the Society produced,
during 1968, one of the Memoirs series, volume 3 entitled “Revisionary Studies
in the Nearctic Decticinae,” by David C. Rentz and James D. Birchim.
Dr. P. H. Arnaud, Jr. read the financial report of the Treasurer, and Mr.
D, G. Denning gave the report of the Auditing committee stating that the finan-
cial records for 1968 are in good order.
Mr. J. S. Buckett, Chairman of the Program Committee, announced that the
next meeting will be held 10 January 1969 which may seem rather soon, but fits
80
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 1
the schedule of our guest, Dr. Schlinger, who will be coming up from Riverside to
speak to us.
Dr. J. F. Gustafson reported that the Salt Marsh Study Committee has com-
pleted its annotated bibliography of the insects of salt marsh habitats which
will appear in volume 44, no. 4 of the Pan-Pacific Entomologist. He reported
that approximately 90 notices of the activities of this project of the Society
have been sent out to journals throughout the coimtry, with the hope that it will
engender similar activities in other parts of the United States.
Mr. Leech, representing the Nominating Committee, presented the slate of
nominees for offices in the Society during 1969. Because of the untimely death
of Howard L. McKenzie, President Hagen agreed to run for the office of President
for another year. The nominees were: Dr. Hagen, President; Dr. R. W. Thorp,
President-Elect; Dr. M. S. Wasbauer, Secretary; and Dr. P. H. Arnaud, Treasurer.
There were no nominations from the floor. The nominees were unanimously elected
to office for 1969.
President Hagen thanked all the committees for their work on behalf of the
Society during the year.
The principal speaker of the evening was Dr. K. S. Hagen, Department of
Entomology and Parasitology, University of California, Berkeley, and President
of the Society. His illustrated talk was entitled “Some beetle relics from the shores
of ancient inland seas.”
A social hour was held in the entomology rooms following the meeting. — M. S.
Wasbauer, Secretary.
CORRECTION AND ADDITION
Fisher, T. W. and R. E. Orth. 1966. A new species of Pteromicra from western
North America and resurrection of Pteromicra pleuralis (Cresson). Pan-Pac.
Entomol., 42(4) : 307-318.
Page 309, fourth line above legend: change 45° to 41° 30'. The holotype and
allotype of Pteromicra siskiyonensis F & 0 have been consigned to California
Academy of Sciences as CAS # 10168. — T. W. Fisher, University of California,
Riverside.
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PUBLICATIONS
OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY
PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL
SOCIETY.
Vol. 1 (16 numbers, 179 pages) and Vol. 2 (numbers 1-9, 131
pages) . 1921-1930. Price $2.50 per volume.
PAN-PACIFIC ENTOMOLOGIST.
Vol. 1 (1924) to present. Price $6.00 per volume of 4 numbers,
or $1.50 per single issue.
MEMOIR SERIES.
Volume 1. The Sucking Lice by G. F. Ferris. A 320-page book
which summarizes the knowledge of the Anoplura of the world.
Published October 1951. Price $6.00. (Plus 350 postage.)*
Volume 2. The Spider Mite Family T etranychidae by A. Earl
Pritchard and Edward W. Baker. This worldwide treatment deals
with the systematics, identification, and economics of the “Red
Spiders” and includes descriptions of 33 new species. 472 pages.
Published July 1955. Price $10.00. (Plus 500 postage.)*
Volume 3. Revisionary Studies in the Nearctic Decticinae hy David
C. Rentz and James D. Birchim. This 173-page revision of Nearctic
shield-back katydids includes descriptions of one new genus and
13 new species and subspecies. Published July 1968. Price $4.00.
(Plus 250 postage.)*
* (Add 5% sales tax on all California orders. Members of the Society will
receive a 20% discount.)
Send orders to:
Pacific Coast Entomological Society
c/o California Academy of Sciences
Golden Gate Park
San Francisco, California 94118
IX
ENTOMOLOGICAL EXCHANGES AND SALES
Small Carpenter Bees (Ceratina) of the World. — ^Wanted for purchase or
exchange for California insects. Write H. V. Daly, Division of Entomology, Univ.
California, Berkeley 94720.
SOCIETY NOTICES
Advertisement of Exchanges. — A special advertising page is,: now available to
members who wish to advertise entomological exchanges, sales, announcements,
etc. We can offer Society members 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 California, Davis, California
95616.
Sale of Past Issues. — We are overstocked in early issues of the Pan-Pacific
Entomologist, and need to reduce our inventory to provide more storage space.
Therefore, we can offer Society members complete sets of volumes 1 through 16
(1924 through 1940) at the extremely reduced rate of $1.50 per volume or $24.00
for the entire first 16 volumes. These are available to Society members only,
on a first-come, first-serve basis as long as the surplus lasts. Contact the Treasurer,
P. H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San
Francisco, California 94118.
New Memoir. — The third volume of the Memoir Series published by the Pacific
Coast Entomological Society was published on 12 July 1968. It is entitled “Re-
visionary Studies in the Nearctic Decticinae” and is authored by David C. Rentz
and James D. Birchim- The authors revise the Nearctic shield-back katydids
(Orthoptera) in this 173-page memoir which is illustrated by 37 figures. Twenty-
two genera and 112 species and subspecies are discussed. One genus and 13 species
and subspecies are described as new. The price is $4.00 plus 250 postage (and
5% sales tax on all California orders). Society members will receive a 20%
discount. Contact the Treasurer, P. H. Arnaud, Jr., California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118.
The Salt Marsh Habitat Committee. — At a meeting in 1966 of the Society,
a lecture on the ecology of the mud flat-salt marsh habitat was presented. Follow-
ing this meeting a number of the members expressed concern regarding the rapid
disappearance of this habitat due to commercial development. Subsequently, the
salt marsh habitat committee was appointed.
The first recommendation of the committee was that the Society encourage
studies concerning the salt marsh habitat in central California. The Society
approved this in 1968 and recommended that notes and observations of salt marsh
insects (and other terrestrial arthropods) be submitted to the editor of the Pan-
Pacific Entomologist for publication.
It is our hope that this action, an indication of scientific concern and interest,
will stimulate similar concerns by entomological societies in other geographic
locations. — Joel F. Gustafson, chairman, San Francisco State College, California.
THE PAN-PACIFIC ENTOMOLOGIST
Information for Contributors
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1958. Insects and mites of western North America. Rev.
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Vol. 45 APRIL 1969 No. 2
THE
Pan-Pacific Entomologist
CHEMSAK — Notes on the habits of Astromula nitidum Chemsak and Linsley
with descriptions of the immature stages (Coleoptera: Cerambycidae) 81
THORP — The identity of Bombus vandykei (Hymenoptera: Apidae) 86
WIRTH AND BLANTON — A new Nearctic species of the genus Paradasy-
helea Macfie (Diptera: Ceratopogonidae) 97
GILLOGLY — Taxonomic notes on Nitidulidae of California (Coleoptera) 100
MAYO — Nymph of Thraulodes speciosus Traver with notes on a symbiotic
Chironomid ( Ephemeroptera: Leptophlebiidae) 103
STRUBLE — Gall wasp infestations in forest trees, chiefly pines, of Californa
(Hymenoptera: Eurytomidae) 112
HYNES — The immature stages of Gonomyodes tacoma Alex. (Diptera: Tipu-
lidae) 116
FENDER — New California Malthodes and distribution notes on other species
in California (Coleoptera: Cantharidae) 119
CRUDEN — A new species of Cordulegaster from the Great Basin region of
the United States (Odonata: Cordulegasteridae) 126
DAILEY — Synonymy of Dryocosmus attractans (Kinsey) and Callirhytis
uvellae Weld (Hymenoptera: Cynipidae) 132
ZIMMERMAN — Francis Xavier Williams, 1882-1967 — 135
PHILIP — New or little-known Neotropical Tabanidae (Diptera) 147
FISHER AND ORTH — Two new species of Sepedon separated from S. armipes
Loew in western North America (Diptera: Sciomyzidae) 152
SAN FRANCISCO, CALIFORNIA • 1969
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
THE PAN-PACIFIC ENTOMOLOGIST
EDITORIAL BOARD
R. W. Thorp, Editor
C. G. Linsley R. O. Schuster, Asst. Editor E. S. Ross
\ D. Hurd, Jr. D. L. Briggs, Editorial Assistant H. B. Leech
P. H. Arnaud, Jr. Treasurer W. H. Lange, Jr. Advertising
Published quarterly in January, April, July, and October with Society Proceed-
ngs appearing in the January number. All communications regarding nonreceipt
»f numbers, requests for sample copies, and financial communications should be
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Sciences, Golden Gate Park, San Francisco, California 94118.
Application for membership in the Society and changes of address should be
iddressed to the Secretary, Dr. Marius S. Wasbauer, Bureau of Entomology, Cali-
ornia Department of Agriculture, 1220 N. St., Sacramento, California 95814.
The annual dues, paid in advance, are $5.00 for regular members of the Society
)r $6.00 for subscriptions only. Single copies are $1.50 each or $6.00 a volume.
Hake checks payable to Pan-Pacific Entomologist.
The Pacific Coast Entomological Society
Officers for 1969
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Statement of Ownership
Title of Publication : The Pan-Pacific Entomologist.
Frequency of Issue: Quarterly (January, April, July, October).
Location of Office of Publication, Business Office of Publisher, Publisher and Owner: Pacific Coast
Entomological Society, California Academy of Sciences, Golden Gate Park, San Francisco, California
94118.
Editor: Dr. Robbin W. Thorp, Department of Entomology, University of California, Davis, California
95616.
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This issue mailed 18 July 1969.
Second Class Postage Paid at Lawrence, Kansas, U.S.A. 66044.
ALLEN PRESS, INC. LAWRENCE, KANSAS
The Pan-Pacific Entomologist
VoL. 45 April 1969 No. 2
Notes on the Habits of Astromula nitidum Chemsak and
Linsley with Descriptions of the Immature Stages
(Coleoptera : Cerambycidae)
John A. Chemsak
University of California, Berkeley
The elaphidionine ceramhycid, Astromula nitidum Chemsak and
Linsley (1965), is known thus far only from several localities in south-
ern California, Joshua Tree National Monument, Riverside County, and
Pearblossom and Valyermo, Los Angeles County. Adults are rarely en-
countered in the field but are occasionally attracted to ultra-violet lights.
Until Tyson (1966) reported the rearing of this species from trunks of
Yucca brevifolia Engelman, the habits were completely unknown.
During the first part of May, 1968, numerous infested trunks of
Y. brevifolia were observed at Valyermo and near vicinity. Successful
rearings of adults were accomplished during August and September,
1968 and observations were made on adult and immature behavior.
In the field, infested trees were relatively easy to spot after the initial
encounter. In all observed cases, the trees were standing and at least
partially alive. In most cases, most of the leaves of the trees were either
chlorotic or partially dead and the initial attack by the beetles had oc-
curred at least one season before. Evidence of infestation was readily
apparent by the emergence holes of the adults. These were visible near
the bases of the trees and very rarely above 2-3 feet from the ground.
These emergence holes were the only visible indications of beetle injury
on the trunk.
The larvae bore longitudinally through the heartwood, filling the
galleries with hard, compacted frass (fig. 2, upper) . Although the con-
dition of initially attacked trees is not known, the areas of larval boring
become necrotic and quite hard. Apparently after several successive
years of attack, the trees die and eventually fall over. The dead centers
of the trunks become reddish brown and filled with larval frass (fig. 1,
upper) . A number of trees had numerous longitudinal galleries in them.
These were very close to each other but never transecting. Infestation
apparently begins near ground level and proceeds up the trunk to about
halfway during successive years.
The Pan-Pacific Entomologist 45: 81-86. April 1969
£2
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Pupation occurs in the heartwood at the end of the larval gallery.
I he pupal chambers are constructed longitudinally and are about 40
r im long and 15 mm in diameter. The upper end of the chamber is
stuffed with very fine loose frass and the pupa is oriented with the head
toward the top of the tree (fig. 2, lower). Adult emergence is through
the thick bark out of round holes about 10 mm in diameter.
Mating behavior of freshly emerged adults was observed in the labora-
tory. A. nitidum is characterized by the very slow, sluggish movements
t)f the adults. During the day or in a brightly lighted room, the beetles
made no attempts at activity and remained motionless in niches and
crevices in the Joshua tree trunks (fig. 3, upper). Mating was induced
]iy darkening the room and placing the males and females next to each
((ther.
Upon contact the male exhibited signs of excitement and mounted
the female. She remained motionless and appeared to be unaware of
the male’s presence. The position of the male’s legs was as follows:
ihe hind legs around the female and on the substrate; the middle pair
vere positioned around the elytra and under in front of the middle legs
of the female; the front legs were on the female pronotum (fig. 3,
lower) . After assuming this position, the male backed down, extended
he penis and inserted it between the apical abdominal sclerites of the
emale. After joining, the male moved the apex of his abdomen slowly
and gently up and down. The body was gently rocked back and forth
also. The initial motions of the male were rather fast. These gradually
slowed down, almost ceasing for a brief time and became rapid again.
These motions consist of a series of short and gentle pullings by the
nale. In the first pair observed, the male pulsations were very slow
after about five minutes. After eight minutes, the motions quickened
For a period and ceased in about two minutes. The male withdrew his
genitalia at this time but remained on the female. During this entire
process the female remained motionless. If she was disturbed and
moved, the male immediately attempted to couple again. If he was suc-
cessful, the entire mating procedure was repeated. No palpating by
the male was observed but the mouthparts were placed on the female’s
pronotum when she moved.
Mating behavior was very similar both in action and duration in
Fig. 1. Upper: Cut away section of Yucca brevifolia showing dark, necrotic
center resulting from larval borings of Astromula nitidum. Lower: Full grown
larva of A. nitidum.
APRIL 1969] CHEMSAK — ASTROMULA NITIDUM HABITS
83
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 2
r
c
4
Fig. 2. Upper: Larva of 4. nitidum in gallery in trunk of Y. brevifolia. Lower:
Pupa and pupal chamber of A. nitidum in trunk of Y. brevifolia.
3ther pairs. All coupled again after a period of no activity and all re-
mained in the mating position for at least 20 minutes after disjoining.
Attempts to observe oviposition were unsuccessful. This was probably
due to the fact that A. nitidum is nocturnal or that the host material
APRIL 1969] CHEMSAK — ASTROMULA NITIDUM HABITS
oc
oD
Fig. 3. Upper: Adult of A. nitidum resting in bark crevice of Y. hrevifolia.
Lower: Mating position of A. nitidum before coupling.
was unacceptable. Judging from the locus of infestation in the field,
eggs are probably laid in crevices in the bark at the base of the host
trees. No immature larvae were found in any of the trees which sug-
gests a one year life cycle for the species. However, as is the case in
156
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
]iiany xerophytic species, some individuals do not emerge until the
r econd year.
Description of larva (fig. 1, lower). — Form robust, rather elongate; integu-
ment tough, shining, milky white. Head rectangularly rounded, slightly narrowed
it apical half; mouth frame heavily corneous, dark brown; labrum oblong rounded,
arownish hairs dense, erect, arranged in an arc over apical one-third; mandibles
: hort, stout, apex roxmded, inner face shallowly excavated, basal piece one-half
length of distal, outside face of distal piece smooth; antennae slender, first seg-
nent arising out of rather large fleshy lobes, third segment very small; ocelli large,
oaired, not enveloped by genae; ventral mouthparts heavily sclerotized, rather
lensely clothed with stiff setae; hypostoma shallowly transversely rugose; gula
anteriorly emarginate, about as long as broad; epistoma projecting over mandibles;
jides and edge of front with numerous long erect setae. Prothorax depressed, over
wice as hroad as long; hairs along lateral and apical margins dense, rather long,
lark golden; apical margin broadly sclerotized, interrupted at middle and sides;
pronotum much broader than long with a sparse band of long hairs along anterior
iiargin, surface shallowly rugose, median suture deep; prosternum rectangular,
broad, median line elevated, rather densely pubescent except for central basal
portion; meso and metasternum transversely, arcuately impressed. Legs rather
short, four segmented, two basal segments with a small tuft of erect setae on
inside. Abdomen with dorsal ampullae prominent, oval, marked by one lateral
and two transverse impressions; epipleurum non-protuberant. Spiracles oval, peri-
treme corneous.
Pupa. — Pronotum transversely, shallowly rugulose, anterior margin with a short
median impression, each side of impression with about 20 very small, seta-bearing
asperites, a small band of short setae also present on each side just before middle.
Abdomen dorsally with acute, recurved, sclerotized spines which increase in size
posteriorly, these arranged somewhat circularly, several spines present on eighth
segment.
The field assistance of J. A. Powell is gratefully acknowledged. These
studies were made during the course of National Science Foundation
Grant GB-4944X on North American Cerambycidae.
Literature Cited
Chemsak, j. a. and E. G. Linsley. 1965. New genera and species of North
American Cerambycidae. Pan-Pac. EntomoL, 41 : 141-153, 4 figs.
Tyson, W. H. 1966. Notes on reared Cerambycidae. Pan-Pac. EntomoL, 42;
201-207.
APRIL 1969]
THORP BOMBUS VANDYKEI IDENTITY
87
The Identity of Bomhus vandykei
(Hymenoptera: Apidae)
Robbin W. Thorp
Department of Entomology, University of California, Davis
The descriptions, sex association and discussion of color variation in
Bomhus (Pyrohomhus) vandykei (Prison) are presented at this time in
order to clarify the use of the name for subsequent publications in prep-
aration on the pollination of Clarkia (Onagraceae) and on the system-
atics and ecology of the Bombini of California.
Prison (1927) described Bomhus vandykei as a “new variety” of B.
flavifrons Cresson on the basis of two workers from Mt. Adams, Wash-
ington. He correctly recognized the affinities of his new variety to the
Plavifrons group, as well as the superficial similarities in color pattern
to both R. hifarius nearcticus Handlirsch and B. edwardsii Cresson.
Stephen (1957) elevated B. vandykei to specific rank after examining
an additional five workers and one queen all taken in northcentral
Oregon. His decision was based on several slight, but consistent, struc-
tural and color pattern differences and the lack of intergrades between
B. vandykei and variants of B. flavifrons. He suggested that B. vandykei
was more likely an “aberrant form” of B. caliginosus, and most closely
resembled certain “color variants” of this species from Riverside and
San Bernardino Counties, California.
Examination of Bombini in numerous collections in western North
America has produced a number of female specimens from California
and southern Oregon which bear a superficial color resemblance to the
color patterns found in B. vosnesenskii Radoszkowski and B. caliginosus
(Prison) and have been variously assigned by previous workers to
these two species. These specimens are the “color variants” of B.
caliginosus discussed by Stephen (1957). Prom females of both B.
vosnesenskii and B. caliginosus they may be readily separated by the
position of the band of yellow hairs on the metasoma. This band is on
tergum 4 in the former species and on tergum 3 in the “color variant”
females. It is my contention that these females are actually a southern
color form of B. vandykei. Recent field collections have not only con-
firmed this idea, but have allowed the association of these females with
the previously unknown male.
Bombus (Pyrobombus) vandykei (Prison)
Bremus flavifrons vandykei Prison, 1927: 375,
Bomhus vandykei, Stephen, 1957 : 106-109.
The Pan-Pacific Entomologist 45: 87-96. April 1969
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
c o
c a
I'ombus caliginosus, Stephen (non Prison, 1927), 1957: 10(1-104. (females, in part,
misidentification. )
I'remus centralis var, monardae, Prison (non Cockerell and Porter, 1899), 1927:
372. (males, misidentification; distribution.)
I'omhus flavifrons dimidiatus, Stephen (non Ashmead, 1902), 1957: 92-95. (males,
in part, misidentification.)
Queens and Workers. — Ocelli located at supraorbital line; malar space 1.3 to
1.5 times as long as apical width; first flagellomere 1.5 times as long as third;
tegula black; wing dark brownish stained; nervures black. Vertex and face with
ling, dense, yellow hairs, each with few dark hairs peripherally. Gena with black
1 air. Thoracic dorsum with broad black interalar band; long, dense, yellow hair
cn scutum anterior to tegular, scutellum and pleura. Metasomal tergum 1 with
predominantly black hair, extreme lateral margins with some yellow hair; meta-
somal tergum 2 with predominantly black hair, extreme lateral and apical fringes
with yellow hair; metasomal tergum 3 with predominantly yellow hair, black
liair on basal third extending apically as narrow V-shaped interruption of yellow
liand; metasomal terga 4-6 with hair black. Apical fringes of metasomal sterna
2-5 with yellow hair. Legs with hair all black; corbicular fringe black. Sting
capsule with inner margin of second valvular ramus slightly notched, sting of
queen with pair of narrow elongate sclerotized plates in coxosternal fascia.
Males. — Ocelli located at supraorbital line; malar space approximately 1.6
imes as long as apical width ; first and third flagellomeres subequal in length ;
egula black; wing lightly stained with brown. Vertex and face with long, dense,
ellow hairs, each with few dark hairs peripherally. Gena with predominantly
ellow hair, few black hairs along outer margin of eye. Thoracic dorsum with
narrow, moderately distinct black interalar band behind tegula and across posterior
end of scutum; rest of scutum, scutellum and pleura with long, dense, yellow hair,
iorne black mixed peripherally on scutellum. Metasomal terga 1-4 with long, dense,
ellow hair; terga 5-7 with predominantly black hair, except extreme apicolateral
nargins fringed with yellow. Legs with hair on coxae, trochanters and femora
"ellow; tibiae and tarsi with hair predominantly rust-brown, except posterior fringe
of mid and hind tibiae with yellow hairs.
Male genitalia. — Apex of penis valve sickle-shaped and broadly dilated; gono-
stylus broader than long; seventh metasomal sternum shallowly medially emargi-
late apically with subapical fringes of long hairs on lateral lobes; eighth meta-
somal sternum hairy and trilobed apically with apex almost membranous and often
lent over with subapical margin shallowly medially emarginate.
Type material. — Holotype worker from Mt. Adams, Yakima Indian
Forest Reservation, Washington, 3,000 feet elevation, taken on 30 June
1925 by E. C. VanDyke is in the collection of the California Academy
of Sciences, San Francisco. A paratype worker with the same collection
Map 1. Geographic distribution of Bombus vandykei. Dots represent the paler
Qorthern female color form of the type; circles represent the darker southern
female color form; circles containing an X are localities represented by males only.
APRIL 1969 ]
THORP — BOMBUS VANDYKEI IDENTITY
89
45
35
a
^0
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
c ata is in the collection of the Illinois Natural History Survey, Urbana.
'^^'he description of the male is based on a specimen from Surin
( = Swim ? ) , near Government Camp, Mt. Hood, Oregon, taken on 25
July 1937 by E. C. VanDyke and in the collection of the California
/tcademy of Sciences.
Distribution. — This species occurs in the Cascade Ranges from
southcentral Washington to northeastern California, the Klamath
Ranges of southern Oregon and northern California, the eastern slope
c f the Sierra Nevada Ranges of California, the Transverse Ranges of
southern California and sparingly in the Coast Ranges of California
I Map 1). Material examined includes 247 males, 85 queens, and 216
1 , orkers from the following localities :
CALIFORNIA: Alameda Co.: Berkeley. Butte Co.: Chico and 12 miles N.E. ;
Jarboe Pass and 6.2 miles N.E. Calaveras Co.: Mokelumne Hill. Contra Costa
Co.: Lafayette. El Dorado Co.: Fallen Leaf Lake; Pilot Hill, 6 miles S. ; Placer-
\ille; Pollock Pines; Riverton; Snow Line Camp. Fresno Co.: Dunlap; Hunting-
tm Lake; Orange Cove, 4 miles N. ; Pinehurst, 2 miles E. ; Tollhouse and 1.5
1 riles N.W. and 2.3 miles E. ; Trimmer, 7 miles S.W. and 10 miles W. Glenn Co.:
iJder Springs, 7 miles E. Lassen Co.: Bridge Creek Camp. Los Angeles Co.: Bell
Canyon, San Dimas Experimental Forest; Claremont; Crystal Lake; Glendale;
Prairie Fork, San Gabriel Mountains; Tanbark Flat. Madera Co.: Lake Millerton;
San Joaquin Experimental Range. Mariposa Co.: Bagby, 17.3 miles N. ; Coulter-
ville, 5 miles W. ; Exchequer Dam and 4 miles W. ; Mariposa; Mariposa Grove,
^ miles E.S.E. ; Midpines Summit; Peregoy Meadows; Wawona; Yosemite Na-
tional Park, Camp Curry, and Valley, and Wawona Tunnel. Merced Co.: Hilmar.
Modoc Co.: Canby, 17 miles W. Mono Co.: Sonora Pass. Monterey Co.: Arroyo
^eco; Arroyo Seco Camp; Carmel; Limekiln Creek. Nevada Co.: Grass Valley;
Nevada City; Sagehen, near Hobart Mills. Placer Co.: Auburn and 10 miles S.E. ;
Dutch Flat; Foresthill and 5 miles W. ; Green Valley; Lake Forest, 0.5 miles
N. ; and Tahoe City. Plumas Co.: Blairsden; Greenville, 2 miles W. ; Johnson-
^ille; Lake Almanor; Quincy and 1.3 miles N. and 4 miles W. ; Taylorville.
Pdverside Co.: Riverside; Tahquitz Peak. Sacramento Co.: Andrus; Folsom and
10 miles N.E. ; Sacramento; Sherman Island. San Bernardino Co.: Barton Flats;
liear Flat, Mt. San Antonio, 5,500 feet; Big Bear City; Big Bear Valley, Pine
I 'rest Golf Course; City Creek Road; Dollar Lake Trail; Falls Public Camp-
ground; Fallsvale; Lake Baldwin; Lytle Creek; Mill Creek, 6,000 feet; Mojave
Iiiver above Deep Creek, Mojave Desert; San Bernardino Mountains, Little Mojave
River; Sugar Loaf Mountain; Vivian Creek Trail, 6,500-6,600 feet; Wildwood
(!anyon, 5 miles E. Calimesa. San Luis Obispo Co.: Atascadero, 5 miles W. ;
Santa Margarita, 5 miles N.E. Santa Barbara Co.: Cachuma County Park; Canada
c el Venadito ; Canon del Refugio ; San Marcos Ranch Headquarters, Santa Inez
Mountains; Santa Ynez Mountains. Santa Clara Co.: Alum Rock Park; Los
iLltos; Los Gatos; Mount Hamilton; Palo Alto; San Antonio Canyon; San Jose;
Stanford University. Shasta Co.: Cassel; Hat Creek Post Office; Hell Creek, 16
iiiles E. Trinity Center. Sierra Co.: Downieville; St. Charles Hill. Siskiyou Co.:
Bartle, 3 miles E. ; Deep Lake ; Etna, 2 miles N. ; Hilt, 17 miles W. ; Klamath
I liver P. 0., 6 and 15 miles E. ; Mount Shasta City ; Sky High Lake, Marble Moun-
APRIL 1969]
THORP BOMBUS VANDYKEI IDENTITY
91
tains; Yreka, 8 miles N. Sonoma Co.: Glenn Ellen; Petaluma; Sobre Vista.
Stanislaus Co.: La Grange. Trinity Co.: Carrville; Coffee Creek R. S., and 10
miles N. ; Nash Mine. Tulare Co.: Aukland; Fairview, 9 miles S. ; Kaweah;
Mineral King; Porterville; Sequoia National Park, 2,000-3,000 feet, and Ash
Mountain, and Potwisha, 2,000-5,000 feet; Springville; Three Rivers. Tuolumne
Co.: Camp Bob MacBride, Pinecrest, 5,400 feet; Columbia, and at Old Catholic
Church; Dodge Ridge; Eleanor Lake; Groveland, 6 miles E. ; Jacksonville, 2.5
miles W.; Mather; Pinecrest; Stanislaus River; 5 miles N.W. of Columbia; Straw-
berry; Tuolumne Camp; Twain Harte; Yosemite National Park, near Crane Flat.
Ventura Co.: Wheeler Springs, Ojai. OREGON: Benton Co.: Corvallis; Marys
Peak. Clackamas Co.: Surin (=Swim?), near Government Camp, Mt. Hood.
Douglas Co.: Azalea; Crystal Spring, 1 mile E. Lemolo Reservoir; Roseburg, 10
miles N.W. and 15 miles E., Whistler’s Bend; Siskiyou Summit near Oakland;
Spencer Creek near Smith River. Hood River Co.: Cloud Cap Inn, Coopers Spur,
Mount Hood, 6,000 feet; Parkdale. Jackson Co.: Butte Falls, and 11 miles W.;
Gold Hill; Gold Ray Dam; Griffin Creek; Laurelhurst State Park; Medford;
Mt. Ashland, 2 miles W. ; Siskiyou Pass; Talent, 2 miles E. ; Trail, 4 miles E.
Jefferson Co.: Warm Springs. Josephine Co.: Grants Pass. Klamath Co.: Beatty,
7 miles S.W. ; Bonanza, 5 miles N. ; Crater Lake National Park; Gearhart Moun-
tain; Keno Springs Ranch; Klamath Falls, Berkeley Springs, and Geary Canal,
and above Geary Ranch; Klamath River, 12 miles S.W. Keno. Lane Co.: Blue
River, Lucky Boy Camp; McKenzie Bridge. Wasco Co.: Mark A. Meyer Park
near The Dalles, 600 feet; Rowena. WASHINGTON: Yakima Co.: Mount Adams,
Yakima Indian Forest Reservation, 3,000 feet; Nile.
Flight season. — Queens may be found in almost any month from
late March to early October with peak records for the spring queens
being during April and for the overwintering queens being late June
and early July. Workers have been recorded from early March to early
September with most records in late June and early July. Males fly from
early May to early September reaching their peak in early July (fig. 1) .
Flower records. — Floral associations are available for 16 queens,
62 workers, and 69 males. The records include flowers from 15 families
and 34 genera. Flower records for queens are mostly with Labiatae
(3), Leguminosae (3), Hydrophyllaceae (2), and Saxifragaceae (2);
those for the workers are predominantly with Leguminosae (16) , Hydro-
phyllaceae (14), Labiatae (12), and Scrophulariaceae (8); those for
the males are primarily with Onagraceae (20), Compositae (18), Legu-
minosae (15), and Labiatae (11). The following records are listed
alphabetically with the number and sex of the specimens associated:
Berberidaceae: Berberis aquifolium Pursch., 1 $. Boraginaceae: Amsinckia
sp. 1 Compositae: Anaphalis margritacea (L.) Beth. & Hook., Id ; Aster sp.,
1 ^ , 5 d ; Chrysopsis villosa (Pursh) Nutt., 1 $ ; Cichorium intybus L., 3 ^ ,
1 d ; Cirsium sp., 6 d ; Grindelia sp., 1 d ; Senecio douglasii DC., 3 d ; Taraxacum
sp., 1 2. Ericaceae: Arctostaphylos sp., 1 2- Hydrophyllaceae: Eriodictyon sp.,
2 ^ ; E. calif ornicum (H. & A.) Torr., 1 ^ ; E. trichocalyx Heller, 1 ^ ; Phacelia
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
^2
70-
Fig 1. Seasonal distribution of the castes of Bombus vandykei from March to
C'ctober with collection records combined into semimonthly segments.
sp., 9$ ; P. alba, 1 $ ; P. grandiflora (Benth.) Gray, 1$, 1^. Hypericaceae:
Pypericum formosum HBK var. scouleri (Hook.) Coult., 1 ^ ; H. perforatum L.,
1 Labiatae: Agastache urticifolia (Benth.) Kuntze, 2 1 $; Monardella
sp., 25 , 1 ^ ; Salvia carduacea Benth., 1 9 ; S. columbariae Benth., 1 $ ; S.
smomensis Greene, 1 $,69; Stachys albens Gray, 19,9^; S. rigida Nutt, ex
I enth., 19- Leguminosae: Cercis occidentalis Torr. ex Gray, 2 9 ; Lotus sp.,
1 $ ; L. scoparius (Nutt, in T. & G.) Ottley, 1 $ ; Lupinus albifrons Benth., 1 $,
1 9 ; Trifolium pratense L., 1 $ , 8 9 , 9 ; T. variegatum Nutt., 3 9,5^; T.
v'ormskioldii Lehm., 1 9 ; Vida sp., 1 9- Liliaceae; Calochortus sp., 1 $. Mal-
vaceae: Althaea fidfolia Cav., 1 9- Onagraceae: Clarkia biloba (Durand) Nels.
& Macbr., SS; C. dudleyana (Abrams) Macbr., ; C. unguiculata Lindl., 39,
13 ; C. rhomboidea Dough, 1 $ ; Epilobiuni sp., 2 Polygonaceae: Eriogonum
S D., 1 $. Rosaceae: Pyrus sp., 1 $ ; Rubus sp., 1 ^ ; R. parviflorus Nutt., 1 9-
S AxiFRAGACEAE : Ribes sp., 2 $. Scrophulariaceae: Collinsia concolor Greene,
1 9 ; Linaria sp., 1 $ ; Penstemon sp., 1 $ , 1 9 , 1 ^ ; P- brevifloris Lindl., 2 9 >
1 ^ ; P. palmeri Gray, 19;^- richardsonii Dough 3 9 •
APRIL 1969]
THORP — BOMBUS VANDYKEI IDENTITY
93
Systematics
Bomhus vandykei belongs to the Flavifrons group of Pyrobomhus
which includes B. flavifrons, B. centralis Cresson, B. caliginosus, B.
vagans F. Smith and their varieties in California, Oregon and southern
Washington. Within this group it appears most closely related to B.
caliginosus. It may be separated from the above species by the charac-
ters in the following key:
Females - 1
Males 6
1. Metasomal terga 3 and 4 with reddish hair 2
Metasomal terga 3 and 4 with yellow or black hair 3
2. Scutum anterior to tegula with black and yellow hair intermixed giving
clouded appearance flavifrons flavifrons Cresson
Scutum anterior to tegula with yellow hair only centralis Cresson
3. Metasomal tergum 1 with yellow hair 4
Metasomal tergum 1 with black hair 5
4. Face with black hair; thoracic dorsum with yellow hair, except for median
patch of black; metasomal terga 1 and 2 with yellow hair
vagans F. Smith
Face with grayish -yellow hair; thoracic dorsum with broad black interalar
band, some black hair intermixed over anterior face of scutum and
scutellum; metasomal terga 1 and 2 with black hair centrally
flavifrons dimidiatus Ashmead
5. Metasomal tergum 3 with yellow hairs apically; tergum 4 with black
hair vandykei (Frison)
Metasomal tergum 3 with hairs black; tergum 4 with predominantly yel-
low hair caliginosus (Frison)
6. Metasomal terga 3 and 4 with reddish hair 7
Metasomal terga 3 and 4 with yellow or black hair 8
7. Interalar band usually distinct laterally to wing base; scutum anterior to
base of tegula with yellow hair, little or no black intermixed
centralis Cresson
Interalar band obscure, abundant light hair intermixed laterally; scutum
anterior to wing base with some black hair intermixed, often giving sur-
face clouded appearance flavifrons flavifrons Cresson
8. Metasomal terga 1-3 with two or more segments bearing yellow hair 9
Metasomal terga 1-3 with black hair only caliginosus (Frison)
9. Metasomal terga 3 and 4 with black hair, at least basally with at most only
small fringes of yellow hair apicolaterally 10
Metasomal terga 3 and 4 with all yellow hair, rarely with few black hairs
medioapically on tergum 4 vandykei (Frison)
10. Metasomal terga 3-7 with black hair vagans F. Smith
Metasomal terga 3-7 with at least some segments bearing yellow hair
peripherally flavifrons dimidiatus Ashmead
I have not found any distinguishing characters for members of this
group in male genitalia or female sting. As in other members of the
9i THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
group, B. vandykei exhibits considerable color variation in and between
populations throughout its geographic range. The descriptions above
are based on the type worker and male from southern Washington and
northern Oregon.
The principal color variation in females of this species is due to the
r ^placement of yellow hair by black hair on the thoracic pleura, scutel-
lum and the lateral margins of metasomal terga 1 and 2 and the apex
cf tergum 2. This is the principal color form of B. vandykei in southern
C>regon and throughout California. A few workers from these popula-
t ons retain the apicolateral yellow fringes on the second metasomal
tergum. The color pattern of southern females places them in the
mimetic complex which includes: B. caliginosus, B. vosnesenskii and
F. californicus F. Smith as well as two species of Psithyrus. Bomhus
landykei is not abundant in the coastal areas where B. caliginosus
c ccurs, it is most abundant in collections from the Transverse Mountain
Ranges of southern California and along the western foothills of the
Sierra Nevada below 6,000 feet in elevation.
Most of the males in Southern Oregon and California possess the
same color pattern as the male from near Mt. Hood, Oregon. These
^'ere, in part, assigned by Stephen (1957) to B. flavifrons dimidiatus
/Lshmead. Earlier determinations by Frison (1927) assigned males of
B. vandykei from California to B. centralis monardae Cockerell and
I’orter. The “5. centralis^^ recorded by Stephen (1957) from western
e nd southern California are also males of B. vandykei. This color vari-
ant of B. centralis was described from northern New Mexico and I have
not seen any valid specimens from California. Fresh males of B.
vandykei possess brilliant lemon-yellow pile as opposed to the more
straw-yellow hairs in other males of the Flavifrons group.
A few males from California exhibit additional darkening through
the replacement of yellow hairs with black hairs on the scutellum. A
laale from Bear Flat and another from Lake Baldwin have mostly
])lack hairs on the scutellum, except apically, and medially on metasomal
terga 1 to 3. A male from near Bagby, Mariposa County, California
lias additional areas of dark hairs completely replacing the yellow on
1 he metapleura, metasomal tergum 1 and basally on terga 2 and 4. Two
males from near Tollhouse, Fresno County, and Grass Valley, Nevada
County are darker with the pleura almost completely covered with black
hair except for an anterior fringe of yellow, the dorsum of the thorax
posterior to the tegulae and metasomal tergum 1 are completely covered
vith black hair, metasomal tergum 2 has only an apicolateral fringe of
, ellow hair, metasomal tergum 3 has considerable black hair intermixed
APRIL 1969]
THORP BOMBUS VANDYKEI IDENTITY
95
with the yellow basally and metasomal terga 4 and 5 have yellow only
on the extreme lateral margins the rest of these segments being covered
with black hair. Typical males predominate in all these localities.
One male labeled “Imperial Co., Calif.” and assigned by Stephen
(1957) to B. centralis belongs to B. vandykei, but the locality is doubtful.
Discussion
Although the present paper attempts to clarify the variations in color
and to associate the sexes of B. vandykei, its taxonomic rank in the Flavi-
frons group of Bomhus cannot at this point be determined. The entire
Flavifrons group contains a number of taxa variously recognized as
species, subspecies and color variants. The taxa recognized as species
and subspecies are for the most part allopatric. In addition there appear
to be no consistent morphological features other than patterns of hair
color which will allow separation of the various taxa. Structural features
in the females such as: the malar spaces being longer than their apical
widths, the first flagellomere being longer than the third, the paired
elongate sclerotized plates in the coxosternal fascia of the sting of the
queens; and in the males: the sickle-shaped, broadly dilated apex of
the penis valve with a distinctly serrate basolateral margin in the genital
capsule, and the shape of the apex of the eighth metasomal sternum; are
characteristics of the entire group which allow easy separation from
other species groups in the subgenus Pyrobombus, but are not diagnostic
within the Flavifrons group. The range of structural variation within
and between populations of any taxon completely overlaps the reported
differences between the taxa presently designated as species.
The figures of male genitalia and seventh and eighth ventral plates
of B. flavifrons, B. centralis, B. caliginosus and B. vagans in Stephen
(1957) exhibit the range of differences within the Flavifrons group.
However, the subtle differences between species expressed in these
diagrams do not hold. The serrate basolateral margins of the head of
the penis valve is reputed to be diagnostic for B. vagans according to
Stephen (1957). But I find this character in the penis valves of all the
species of the Flavifrons group, as well as in other North American
Pyrobombus, Cullumanobombus, Separatobombus and F raternobombus .
The figures of the eighth ventral plates in Stephen (1957) indicate
greater variation than really exists unless the situation is clearly under-
stood and closely examined. His figure for B. caliginosus is typical for
the group. But the trilobed or at least medially acutely pointed apex is
thin and almost membranous and is frequently folded over. The result-
ing subapical margin is truncate or shallowly emarginate medially. This
^6
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
condition is figured by Stephen (1957) for B. flavifrons and B. centralis.
Thus, the separation of these taxa is dependent on differences in
color pattern. In areas where their distribution overlap integradation
in color patterns occurs between some of the taxa. Stephen (1957)
mentions the occurrence of intergrades between the subspecies of B.
jlavifrons, B. /. flavifrons, and B. f. dimidiatus, pointing out that the
lypical forms of these taxa predominate. He also states, “A great num-
]»er of specimens from eastern slopes of the Cascades in southern Oregon
end northeastern California could be either /. flavifrons or centralis,
]»ut the blending of diagnostic characters applicable to northern popu-
lations of the 2 species makes separation impossible.” Bombus f.
dimidiatus and B. vagans are phenotypically similar, but can be sepa-
rated by the characters used in the key above. Females of B. vandykei
Irom southern Oregon and California are phenotypically similar to
ihose of B. caliginosus except for the position of the yellow band on the
inetasomal dorsum as indicated in the key. Males of B. vandykei exhibit
rome resemblances in color to the lighter males of B. f. dimidiatus, but
they always have the third metasomal tergum completely covered with
^^'■ellow hair, while the latter possess some black hairs at least basally
on this segment. In addition the fourth tergum in B. vandykei never
attains the extensive coverage of black hairs which are found in the
lightest males of B. f. dimidiatus.
At our present state of knowledge it is not possible to satisfactorily
solve the systematic position of the various taxa of the Flavifrons group.
Additional species, B. pleuralis Nylander and B. frigidus F. Smith, and a
number of varietal and subspecific designates of the species in this group
have not been discussed here since they are not in the geographic range
of B. vandykei. In order to clarify the limits and rank of the various
axa, we will need more extensive field collections and ecological studies
rom areas of sympatry among populations of the taxa, detailed biologi-
cal studies, laboratory colonizations and controlled cross matings.
Literature Cited
PRISON, T. H. 1927. Records and descriptions of western bumblebees (Bremidae).
Proc. Calif. Acad. Sci., 4th Ser., 16 (12) : 365-380.
Stephen, W. P. 1957. Bumble bees of western America (Hymenoptera: Apoidea).
Oreg. Agr. Exp. Sta. Tech. Bull., 40: 1-163.
APRIL 1969] WIRTH & BLANTON NEW NEARCTIC PARADASYHELEA 97
A New Nearctic Species of the Genus Paradasyhelea Macfie
(Diptera: Ceratopogonidae)^
Willis W. Wirth and Franklin S. Blanton
Entomology Research Division, Agr. Res. Serv., USD A" and
Department of Entomology, University of Florida, Gainesville
The genus Paradasyhelea has been known until now from only four
species restricted to the southern part of the Southern Hemisphere:
The type species D. brevipalpis (Ingram and Macfie) from Patagonia,
P. egregria (Macfie) from New Zealand, and P. albipunctata Wirth
and Lee and P. minuta Wirth and Lee from Australia. On a recent col-
lecting trip to the Pacific Northwest one of us had the good fortune to
pick up a fine series of an undescribed species belonging to this genus.
This new North American record extends the range of the genus in a
most unexpected and remarkable manner.
Instances of taxa with distributions mainly in the “Southern End of
the World” (Darlington, 1965) but with one or a few species on the
Pacific Coast of North America are extremely rare and of consequent
great zoogeographic interest. One example is the genus Bequaertomyia
with one Pacific Coast species (Philip, 1965), the only member of the
family Pelecorhynchidae found outside Australia (27 species) and
southern Chile (5 species) (Hennig, 1960). The genus Parochlus in
the chironomid subfamily Podonominae has 44 species in the temperate
parts of the Southern Hemisphere but only one in the northern part of
the Holarctic Region (Brundin, 1966). We have recently found a new
species of the biting midge genus Leptoconops in southern California
greatly resembling the species in the subgenus Styloconops found in
Australia, New Zealand, and the Oriental Region. The Pelecorhynchidae
and Podonominae are cold-adapted groups resembling Paradasyhelea
in distribution patterns, while Styloconops is warm-adapted with dis-
tribution accordingly. Whether the groups mentioned are “relicts” of
previously widespread ancestors of northern origin, or originated in the
south and are present in the north as “immigrants” (terminology from
Darlington, 1965, p. 153 and elsewhere) is still a moot question with
little data from animals as ancient as insects to form a basis for con-
clusions.
Paradasyhelea seems to be a relict ceratopogonid of an extremely
annectent type with characters cutting across three subfamily lines. It
1 This investigation was supported in part by U. S. Army Contract No. DA-49-193-MD-2177.
-Mail address: c/o U. S. National Museum, Washington, D.C. 20560.
The Pan-Pacific Entomologist 45: 97-100. April 1969
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
lesembles Forcipomyia (subfamily Forcipomyiinae) in the vestiture of
the wings, oblique r-m crossvein, broadly separated eyes, presence of
scattered, slender, byaline, sensory pegs on the antenna, and in structure
of the male basistyles and parameres. It resembles Dasyhelea (sub-
family Dasyheleinae ) in the slender 3-4 segmented palpus, hairy eyes,
short proboscis with rudimentary mouthparts, presence of basal verticils
on all of antennal segments three to fifteen, male antenna with last four
segments elongated, and the shape of the apicolateral processes of the
male genitalia. It resembles Culicoides (subfamily Ceratopogoninae)
m the presence of well-developed humeral pits, presence of minute distal
liensory pits bordered by fine setae on certain antennal segments, distal
antennal segment without terminal papilla, and the V-shaped or Y-shaped
aedeagus. It differs from all these genera by having both radial cells
obliterated and tbe costa extending less than halfway to wing tip.
Genus Paradasyhelea Macfie
^aradasyhelea Macfie, 1940, Ann. Trop. Med. Parasit. 34: 17. Type-species,
Dasyhelea egregria Macfie (by original designation) .
Wirth and Lee (1959) reviewed the genus with a diagnosis of the
male, female, and pupa, and a table comparing the four known species,
;o which the reader is referred for a full discussion of the genus.
Paradasyhelea olympiae Wirth and Blanton, new species
(Figs. 1-5)
Female. — Wing length 0.82 mm. Head-. Brown. Eyes broadly separated, with
dense, fine, interf acetal hairs. Antenna (fig. 3) with lengths of flagellar segments
in proportion of 25-15-15-15-15-15-15-16-18-20-23-25-36; proximal segments slightly
longer than broad, gradually becoming longer and slenderer through eleventh;
three distal segments stouter, each a little more than twice as long as broad; only
third segment bearing a pair of small sensory pits bordered by minute setae. Only
three distinct palpal segments (fig. 2), the first (primitive third) about as long
as distal two combined, bearing a small round distal pit containing a few long-
stalked hyaline sensilla. Thorax: Shining brownish black; scutum with sparse
vestiture of long, suberect, brownish hairs; scutellum with four long bristly hairs.
Legs yellowish brown, with sparse vestiture of short hairs; hind tibial comb with
five spines, the second from the spur longest; hind basitarsus 2.2 as long as
second tarsomere; claws short and sharp, nearly straight, a minute, branched
empodium present. Wing (fig. 4) : Uniformly grayish brown, with rather long,
fine, decumbent, brown hairs, dense distally but sparse on proximal portion of
wing; costa extending to 0.47 of wing length; radial cells obliterated. Halter
grayish. Abdomen: Pale brown. One retort-shaped spermatheca (fig. 1) present,
the large distal end directed caudad, the spermatheca thus bent back along the
slender neck at base of the duct; spermatheca relatively large, measuring 0.058
by 0.038 mm.
APRIL 1969] WIRTH & BLANTON NEW NEARCTIC PARADASYHELEA 99
Figs. 1-5. Paradasyhelea olympiae Wirth & Blanton. Fig. 1. Female spermatheca.
Fig. 2. Female palpus. Fig. 3. Female antenna. Fig. 4. Female wing. Fig. 5.
genitalia, ventral view
Male.^ — Similar to the female with the usual sexual differences; antennal plumes
well developed, distal three antennal segments elongate, in proportion of 62-44-44.
Genitalia (fig. 5) : Ninth sternum with moderately deep caudomedian excavation,
the ventral membrane spiculate for half the height of aedeagal arch; ninth tergum
long and tapering, with well separated, large, angular, apicolateral processes, the
eaudal margin between them nearly straight. Basistyle moderately stout, inner
margin with sharp dorsomesal process directed toward and nearly meeting the
triangular, well-sclerotized, median sclerite of parameres; dististyle with stout
base, distal portion slightly curved, tapering to quite slender, pointed tip. Aedeagus
V-shaped, basal arms nearly straight, the basal arch extending to about two-thirds
of total length of aedeagus; distal process pointed, with a few fine subapical
spicules and a pair of short, blackish, sharp-margined lips on ventral side proximad
of the spicules.
Types. — Holotype female, allotype male, on slides, Elwha River,
Olympic National Park, Washington, 6 July 1968, W. W. Wirth,
swept from boggy seepage area in open forest near Lake Mills (Type
no. 70340, USNM). Paratypes, 21 males, 2 females pinned; 37 males,
11 females on slides; same data as type. Paratypes will be deposited in
the U. S. National Museum, Washington; British Museum (Natural
History), London; California Academy of Sciences in San Francisco,
and the Canadian National Collection, Ottawa.
00
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Discussion. — Paradasyhelea olympiae, although agreeing well in the
generic characters of the male genitalia with the four known species,
differs from all of them in the shape of the apicolateral processes of the
ninth tergum, the aedeagus, and the median sclerite of the parameres.
J^emales are known of only the two Australian species; P. olympiae
iiffers from both of them in the shape of the spermatheca, in the com-
rlete reduction of the palpal segments proximad of the one bearing the
sensory pit, in the less hairy wings, in the shorter female antennal seg-
iients, and by the only slight lengthening of segment twelve in the male
antenna.
Literature Cited
Brundin, L. 1966. Transantarctic relationships and their signifieance, as evi-
denced by chironomid midges vfith a monograph of the subfamilies
Podonominae and Aphroteniinae and the Austral Heptagyiae. Kgl.
Svenska Vetenskapsakad. Handl., Band 11, Nr. 1, 472 pp., 30 plates.
Darlington, P. J., Jr. 1965. Biogeography of the Southern End of the World.
236 pp. Harvard Univ. Press, Cambridge, Mass.
Hennig, W. 1960. Die Dipteren-Fauna von Neuseeland als systematisches und
tiergeographisches Problem. Beitr. EntomoL, 10: 221-329. (translation
by P. Wygodzinsky in Pacific Insects Monograph 9: 1-81, 1966).
Philip, C. B. 1965. Family Pelecorhynchidae. p. 319. In Stone, et al. A catalog
of the Diptera of America North of Mexico. U. S. Dep. Agr. Handb.
276, 1696 pp.
WiRTH, W. W., AND D. J. Lee. 1959. The genus Paradasyhelea Macfie, with
descriptions of two new species from eastern Australia (Diptera:
Ceratopgonidae) . Bull. Brooklyn EntomoL Soc., 54: 114-121.
Taxonomic Notes on Nitidulidae of California
(Coleoptera)
Alan R. Gillogly
University of California, Riverside
In my studies of the nitidulids of California, two items of taxonomic
importance have come to my attention. The first of these is an error in
synonymy, and the second is the presence of an undescribed species in
the southwestern United States.
The genus Cateretes Herbst is represented in California by two sub-
species placed under the name Cateretes pennatus (Murray) by Dr. Hatch
in 1961. Although I agree with the subspecific status of these, I believe
that the resulting specific name should be the older Cateretes sericans
(LeConte), and that the subspecies should be called Cateretes sericans
The Pan-Pacific Entomologist 45: 100-102. April 1969
APRIL 1969]
GILLOGLY- — NOTES ON NITIDULIDAE
101
sericans (LeConte, 1859) and Cater etes sericans pennatus (Murray,
1864) .
The new species belongs to the genus Colopterus Erichson, bringing
to seven the number of known species in the United States. Two of these,
including the new species, occur in California.
Colopterus testaceus A. R. Gillogly, new species
Male (holotype ) . — Oblong oval, moderately depressed, testaceous, pubescence
thick, golden. Head coarsely punctate, punctures oblong, separated by one-third
diameter or less, intervals between punctures reticulate; terminal segment of anten-
nae acutely angled at apex.
Prothorax width at widest point to length at midline as 1.9 to 1, apex shallowly
emarginate, base feebly bisinuate ; sides of prothorax arcuate, most strongly in basal
third, widest at basal third; ciliate laterally, forming band as wide as widest part
of third antennal segment; hind angles broadly rounded, not projecting backward.
Disk of pronotum shallowly punctate, punctures oblong, distinctly separated by
one-half to one puncture diameter, intervals between punctures mixed reticulate
and transversely alutaceous. Scutellum moderately punctate, impunctate in apical
fifth, punctures separated by at least one-half diameter, intervals between punc-
tures transversely alutaceous, apex evenly, narrowly rounded.
Elytra closely, serially punctate and setose, intervals between punctures finely
reticulate; lateral margins eiliate, apices broadly rounded, sutural angles obtusely
rounded. Abdominal tergites sparsely punctate and setose, each puncture with a
small, seta-bearing granule at anterior end, intervals between punctures retieulate.
Pygidium truncate at apex, very shallowly emarginate at middle of hind margin.
Prosternum sparsely punctate, anterior one-half almost impunctate in middle,
intervals between punctures alutaceous. Prosternal process between coxae bearing
a number of strong setae. Abdominal sternites obsoletely, shallowly ptmctate,
densely setose, each seta on a small, raised granule, intervals between granules
mostly reticulate, almost transversely alutaceous near midline of each sternite.
Hypopygidium shallowly emarginate, distinctly sinuate; lateral margins with
seven small spines on left side and nine on right side. Length 3.7 mm, width 2.0
mm.
Female {allotype ). — Differs from holotype in the following respects: disk of
pronotum slightly more closely punctate, apex of scutellum broadly rounded, apex
of pygidium broadly rounded, hypopygidium not emarginate; length 4.0 mm, width
2.0 mm.
Holotype male. Sycamore Canyon, Atasco Mts., Arizona, 22 May
1954, bait trap, G. D. Butler. Allotype female same data as holotype.
Paratypes, 14 males, 7 females: Arizona: 1 male, 1 female, same data as holotype;
1 female, Arivaipa, 25 August 1933, Bryant; 1 male, Canelo, 10 July 1957, G. D.
Butler; 1 female, Chiricahua Mts., Cochise Co., 3.5 mi. S.W. Portal, 5000 ft.,
13 August 1952, H. B. Leech; 1 male, 2 females, Cochise Stronghold, Dragoon Mts.,
U.V. light trap, 21 July 1961; 4 males. Globe, rotten watermelon, 7 October 1949,
F. H. Parker; 1 male, Graham Mts., Moon Creek, bait, 8 July 1955, G. Butler and
F. Werner; 1 male Hk. Hwy. mi. 5, Santa Catalina Mts., U.V. lite, 11 August 1961,
]02
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
T7erner and Nutting; 1 male, 1 female, Madera Canyon, Santa Rita Mts., 5000 ft.,
U.V. lite and trap, 18-23 June 1962, F. Werner; 1 male, Patagonia, Santa Cruz Co.,
iO May 1955, G. Butler; 1 male, Santa Rita Mts., 5-6, Hubbard and Schwarz; 1
lemale, 9 mi. east Superior, 23 July 1956, Butler and Gerhardt. California: 1 male,
( .4 mi. S. E. Houser Creek Public Camp near Campo, San Diego Co., el. 1800 ft.,
August 1965, R. Somerby and B. Ruge. Utah: 1 male, Leeds Cyn., Washington
Co., light trap, 14 August 1961, G. F. Knowlton.
Additional specimens referred to this species: Mexico: 1 male, 1 female, 44 mi.
IJ. E. Villa Union, Sinaloa, el. 4400 ft., 9 July 1962, E. Sleeper, R. Anderson, A.
Hardy, R. Somerby.
The holotype and allotype have been deposited in the California
Academy of Sciences, and paratypes have been placed in the following
collections: California State College at Long Beach, A. R. Gillogly,
R. Gillogly, C. T. Parsons, United States National Museum, University
of Arizona, University of California at Berkeley, and University of
California at Davis.
The variation in the paratype series is limited, as follows: form
oblong oval to elongate oval; pronotal punctures from one-third to one
liameter apart; elytral apices occasionally slightly obliquely rounded;
nale pygidium sometimes shallowly emarginate; prosternum sparsely
o moderately punctate in middle; spines on margin of hypopygidium
sometimes very small, worn, variable in number; apex of female scutel-
lum narrowly to broadly rounded; color from light testaceous to casta-
tieous; male length 2.9-4.4 mm, width 1.7— 2.2 mm; female length 2.9-
4.0 mm, width 1. 7-2.2 mm.
C. testaceus is most similar to C. unicolor (Say) from the eastern
United States, but can be distinguished by its serially punctate and setose
elytra, and the broadly rounded hind angles of the pronotum. From
C. truncatus (Randall), the most common western species, C. testaceus
can be separated by its serially punctate and setose elytra, broadly
rounded elytral apices, and lack of maculation.
Literature Cited
Hatch, M. H. 1961. The Beetles of the Pacific Northwest, Part HI: Pselaphidae
and Diversicornia I. University of Washington Press, Seattle, 503 pp.
(specifically, pp. 127-128).
LeConte, J. L. 1859. Proc. Acad. Natur. Sci. Philadelphia, p. 69.
Murray, A. 1864. Monograph of the family Nitidulariae. Trans. Linn. Soc.
London, 24, p. 235.
APRIL 1969]
MAYO THRAULODES SPECIOSUS NYMPH
103
Nymph of Thraulodes speciosus Traver with Notes
on a Symbiotic Chironomid
( Ephemeroptera : Leptophlebiidae)
Velma Knox Mayo
Tucson, Arizona
Rustler Park, altitude 8,000 to 9,000 feet on the east side of the
Chiricahua mountains of Arizona is the type locality of the adults of
Thraulodes speciosus Traver. Near there, at an altitude of 5,400 feet at
the Southwestern Research Station of the American Museum of Natural
History in Cochise County, five miles west of Portal, Arizona, the
nymphs of this species were found in abundance and reared as follows :
one female imago, 25 June 1960; one male and one female imago, 10
July 1960; two female subimagos, 26 July 1960. They were the most
common mayfly nymph in the region.
Thraulodes speciosus Traver
(Figs. 1-17, 19-25)
Nymph (fig. 1). — Length: body 8-9 mm; caudal filaments 15-16 mm. General
color brown on dorsal surface. Head: color brown mottled with dark brown;
midline pale; epicranial suture white; ocelli white with wide black crescent at
base; area between medial ocellus and lateral ocelli dark brown; dark brown
around bases of antennae; first two segments brown, other segments pale yellow
basally, brown distally. Antennae twice as long as width of head. Mouthparts
(figs. 2-5, 7, 11). Labrum and mandibles prominent from dorsal view. Maxillae
with dense, thick brush of bright rust hairs (fig. 11) ; ventromedially with row
of about 14 large, toothed spines, inner margin with long, single row of pale
hairs; labrum (fig. 7) with apical margin slightly emarginate, with 5 small
crenulations. Thorax: pronotum brown with variable pattern of dark brown
streaks; posterior border pale yellow with pale area extending anteriorly at
midline; pale areas along anterior and lateral borders; anterior border slopes
rearward laterally; short hairs along anterior lateral borders. Intersegmental areas
of thorax pale yellow. Mesonotum dark brown, darker than pronotum, mottled
with lighter brown laterally; long, pale yellow, prominent U-shaped mark at
midline (fig. 1) ; midline pale. Scutellum yellow between wing pads; suture
between scutum and scutellum marked with wide dark reddish-brown streak
extending on either side of midline laterally and anteriorly to base of each wing
pad; large yellow spot on base of each wing pad with black spot at lateral pos-
terior corner; hind wing pads on metanotum covered by forewing pads. Thoracic
pleuron sclerites dark brown, some with black borders; unsclerotized areas
yellow. Legs paler than body, with fringe of fine, long pale hairs along postero-
lateral margins of coxae, and along outer margins of femora and tibiae; femora
and tibiae with heavy, reddish-brown spines among marginal hairs; coxae dark
brown with small black patch basally and black along posterolateral margin;
trochanters brown, paler than coxae; femur I with a gray to black spot dorso-
The Pan-Pacific Entomologist 45: 103-112. April 1969
]04
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 2
Fig. 1. Thraulodes speciosus Traver, female nymph. Fig. la. Gill
APRIL 1969] MAYO THRAULODES SPECIOSUS NYMPH
105
medially near base (figs. 20, 21) ; some specimens with light brown area medi-
ally with yellow area anterior and posterior to this spot, other specimens with
light brown area over entire femur except for pale strip along posteriorapical
border; femora II and III lacking black spot near base, with yellow strip along
anterior border extending medially near base, and yellow area along posterior
border as in femur I (figs. 19, 22, 23) ; some specimens with two more extensive
yellow areas separated by light brown medially; specimens with more extensive
brown areas most common; pale areas variable from side to side on some nymph;
tibia I yellow with black band apically, narrowly margined with reddish-brown;
tibiae II and III yellow, narrowly penciled with reddish-brown dorsobasally ; all
tarsi pale to dark brown except for pale yellow basal one-fifth, narrowly margined
with reddish brown at base; denticles and tips of all claws reddish-brown. Spines
on legs: dorsal surface of trochanters with three groups of grooved spines (figs.
8, 9) : longitudinal row on side that attaches to femur, oblique row parallel to
posterior margin that attaches to coxa, and a group near dorsal margin; all
grooved spines probably pinnate; trochanter I with from 6-10 spines in longi-
tudinal row, from 7-8 in oblique row and from 0-4 in group near dorsal margin;
trochanter II with from 4-9 in longitudinal row, from 4-7 in oblique row, and
from 1-4 in dorsal margin group (figs. 8, 9) ; trochanter III with from 4-8 in
longitudinal row, 0-1 in oblique row, 0-1 in dorsal margin group. All femora
with short, grooved fairly blunt-tipped spines (fig. 17) irregularly spaced over
dorsal surface, except femur I with spineless area near base; outer border with
long spines among hairs, some spines slightly narrowed near tip, spine tips
widened and somewhat spoon-shaped or spatulate, some long spines appear sharper
at tips, not spatulate; femur I with fewer spines along outer border than femora
II and III, with single row of pinnate spines on dorsobasal one-third near outer
border, some specimens show these among hairs; without spines on ventral sur-
face; ventral surfaces of femora II and III, between midline and inner border
with small comb-like pinnate spines slightly curved with spinelets on inner side
only (fig. 15) occurring in irregular rows about 3-5 deep, all spines curved and
pointed uniformly. Tibiae and tarsi, leg I: tibia with 3-4 irregular rows of
many coarsely pinnate spines crowded together (fig. 16) along entire inner
border, with even row of long pinnate spines (fig. 24) dorsoposteriorly, apical
inner corner of tibia with cluster of both types of spines with pinnate spines longer
than border spines and curved inward (fig. 25) ; ventral surface of tarsus near
midline with 5-7 short pinnate spines and one coarsely pinnate spine; ventral
apical margin of tarsus with blade-like spine (fig. 12), claw with 6-7 large
denticles and 4-5 minute basal ones (as in fig. 6) ; tibiae and tarsi leg II: tibia with
one long pinnate spine on ventral apical margin and eight smaller pinnate spines
along midline; tarsus with three nonpinnate spines; claws with seven large den-
ticles and six minute basal ones; (one specimen tibia with one long and one
short pinnate spine on apical margin near inner border) ; four pinnate spines
along midline, seven nonpinnate spines along inner border; tarsus with blade-like
spine (fig. 12) on apical border and five additional small nonpinnate spines;
tibiae and tarsi, leg III: tibia with five to seven pinnate spines across ventral
apical margin (fig. 10), thirteen specimens with five spines across, two with six,
and one with seven, one long nonpinnate sturdy spine in same row at one end,
one specimen with twenty eight long pinnate spines along two-thirds of shaft
from near midline to outer border on one specimen, not in rows, but irregular;
1D6
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Figs. 2-17. Thraulodes speciosus. Fig. 2. Left mandible. Fig. 3. Right man-
c ible. Fig. 4. Hypopharynx. Fig. 5. Labium. Fig. 6. Claw from leg III. Fig. 7.
Labrum. Fig. 8. Trochanter, leg I. Fig. 9. Trochanter, leg II. Fig. 10. Ventral
APRIL 1969] MAYO THRAULODES SPECIOSUS NYMPH
107
eighteen specimens with from thirteen to thirty pinnate spines along two-thirds of
shaft, tarsus with five small nonpinnate spines and one long blade-like spine at
apical ventral border (fig. 12) ; claws with average of seven large denticles and six
minute ones nearer base; of twenty seven specimens: four with five denticles,
eight with six, eleven with seven, four with eight, all with three to seven minute
denticles near base. Abdomen: few specimens with abdominal tergite 1 yellow to
pale brown, some specimens with tergites 1-3 yellow to pale brown; most specimens
with tergites brown except for tergites 4-5 and 9; tergites 4-5 of most specimens
with variable yellow triangular area at midline, with base of triangle along
posterior border, in some specimens with pale areas on either side of midline,
and absent from a few specimens; tergite 9 with yellow area at midline; tergite
10 narrowly yellow along anterior margin, rimmed with black along posterior
border. Some specimens with dark brown spots near base of gills on tergites
1-7 ; pale streak under gills extending from posterior margin to two-thirds distance
to anterior margin, very close to medial side of gill attachment. Minute spines
present on posterior borders of abdominal tergites; intersegmental areas pale
with diffuse, transverse, penciled streaks. Posterolateral spines on tergites 2-9,
those on 2-8 small; spine projects rearward laterally on tergite 9, length equal to
depth of tergum at midline. Gills double, lanceolate, widest on segments 1 and 2,
decreasing in size rearward with edges white; main tracheae dark gray with
delicate branches obscured by the gray pigmentation (fig. la). Ventral surface
of body pale yellow. Ganglia on cervix, one on prosternum, two on mesosternum
and one on abdominal sternite 7 prominent dark gray, other ganglia on abdomen
faint, on some specimens not apparent. Mesosternum lateroposteriorly with wide,
light brown longitudinal streaks. Abdominal sternites 2-8 laterally with light
brown streaks about midway between anterior and posterior borders, slightly
diagonal to pleural fold (apparent on most specimens). Caudal filaments pre-
dominantly light brown, with dark brown on alternate segments basally, every
fourth segment medially, with apical third entirely light brown; spines on pos-
terior border of each segment in basal half; apically with fine white hairs in
place of spines ; terminal filament more stout than cerci.
<-
apical margin tibia, leg III. Fig. 11. Maxilla. Fig. 12. Blade-like spine on tarsus,
leg I. Fig. 13. Coarsely pinnate spine on tarsus, leg I. Fig. 14. Coarsely pinnate
spine on tarsus, leg I. Fig. 15. Comb-like spine on ventral surface femora II, III.
Fig. 16. Coarsely pinnate spine on leg I. Fig. 17. Spine on dorsal surface femur.
Fig. 18. Oak Creek Canyon specimen, spine on dorsal surface femur. Figs. 19-25.
Thraulodes speciosus. Fig. 19. Pattern on dorsal surface femur, leg II. Fig. 20.
Pattern on dorsal surface femur, leg. I from same specimen as fig. 19. Fig. 21.
Pattern on dorsal surface femur, leg I showing variation from fig. 20. Fig. 22.
Pattern on dorsal surface femur, leg III. Fig. 23. Pattern on dorsal surface femur,
leg II. Fig. 24. Pinnate spine on ventral surface tibia, leg III. Fig. 25. Group of
pinnate spines on tibia I. Figs. 26-30. Oak Creek Canyon specimens. Fig. 26.
Pattern on dorsal surface femur I. Fig. 27. Pattern on dorsal surface femur I
showing variation from fig. 26. Fig. 28. Pattern on dorsal surface femur leg II,
left side. Fig. 29. Pattern on dorsal surface femur leg II right side same specimen
as fig. 28. Fig. 30. Pattern on dorsal surface femur leg III.
138
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Figs. 31-40. Symhiocladius sp. Fig. 31. Larva. Fig. 32. Caudal end, larva.
Fig. 33. Anterior proleg larva. Fig. 34. Caudal end, pupa. Fig. 35. Lateral view
APRIL 1969] MAYO THRAULODES SPECIOSUS NYMPH
109
Variations. — In male pale yellow of posterior border of head extends anteriorly
between dark red turbinate eyes; area between anterior border of eyes dark
brown; in female area between eyes mottled with brown near midline.
The description of the nymph was based on 220 nymphs taken by V.
K. Mayo 4 June 1959, 25 and 26 June 1960, 9 and 10 July 1960, and 10
September 1960 from a stream that runs through the American Museum
Research Station near Portal, Arizona; in entomological collection
University of Utah, Salt Lake City, and 6 nymphs in collection of Jay R.
Traver, Amherst, Mass.
Nymphs which were taken at Oak Creek Canyon near Sedona, Coco-
nino County, Arizona 23 June, 1951 were tentatively described by
Traver (1967) as belonging to the species Thraulodes arizonicus
McDunnough. These are very similar to the nymphs of T. speciosus
and the following are the differences between the two series. The only
difference between the spines of the two series seems to be between the
small grooved spines on dorsal surfaces of femora. The tips of these
spines on Oak Creek Canyon specimens (fig. 18) are more blunt than
those of T. speciosus (fig. 17) and are nearly as wide as the base and
slightly flattened across the end. On femur II of two specimens of
T. speciosus, one blunt tipped spine like those on Oak Creek Canyon
specimens was found and on Oak Creek Canyon specimens there was an
occasional spine more pointed, somewhat resembling those on T. speci-
osus. There seem to be more spatulate spines among hairs on the outer
margin of Oak Creek Canyon specimens than on T. speciosus. The
color pattern of abdominal terga differs with more yellow on Oak
Creek Canyon specimens: tergite 7 with faint yellow area close to pos-
terior margin at midline; tergite 8 with yellow area at posterior margin
extending midway to anterior margin, with a narrow brown streak along
midline. Tergites of T. speciosus 7-8 are brown. Laterally on either
side of the gill attachment, abdominal terga of Oak Creek Canyon
specimens are yellow; T. speciosus has only a pale streak under the gill.
Posterior margins of tergites 7-10 of Oak Creek Canyon specimens are
narrowly rimmed with dark brown; only tergite 10 of T. speciosus is
rimmed with black posteriorly. Caudal filaments of Oak Creek Canyon
specimens dark brown; those of T. speciosus are yellowish-brown.
A symbiotic relationship was found between a large number of
nymphs of T. speciosus and a chironomid larva belonging to the genus
labial plate of larva. Fig. 36. Labial plate of larva, dorsal view. Fig. 37. Mandible,
larva. Fig. 38. Larva on nymph of Thraulodes speciosus. Fig. 39. Antenna, larva.
Fig. 40. Pupa on nymph of T. speciosus.
10
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
S ymbiocladius Kieffer (Kieffer, 1925). The chironomid larvae and
some pupae were found attached along the sides of the abdominal tergum
0 the nymphs of T. speciosus, medial to the bases of the gills. A few
s])ecimens had two larvae attached, one on either side. On one nymph
a larva was found attached along the abdominal sternum ventral to the
bases of the gills. In some cases the chironomid larvae had prevented
the gills from developing and some nymphs were found on which the
gills were atrophied on one side. The larvae and pupae are encased in
a yellowish granular membrane, and where this is attached to the nymph
there is a deposit of silt; figs. 38, 40 show the larva and pupa on the
a bdomen of the nymph, the coarse stippling representing the silt. In
f]g. 38, a large larva is shown attached to an immature nymph: head
lies along side of wing pad between second and third legs; a thick
c umpled deposit with silt extends beyond head and is firmly attached
aiong side of wing pad close to second coxa; the silt is cemented down
s de of tergum; caudal end of larva has broken out of case and is
s ightly raised. The drawings were made from specimens that had been
i;i alcohol for some time and parts of membranous cases had been
broken. On the same specimen there is evidence that a second larva
lad been attached to the right side where part of the membrane with
S ilt is deposited near the bases of the gills on tergites 1—5. The specimen
V as collected at the Southwestern Research Station on 9 July 1960.
(ihironomid larvae were found attached to 9 additional nymphs.
Following are descriptions of pupae in relationship to nymphs: in fig.
40 a pupa is shown attached to the left side with head rearward; caudal
end is between second and third coxae and a thick membranous deposit
\dth silt extends along side of wing pad nearly to second coxa; there
is a deposit of silt along the tergum with a thickened deposit extending
leyond head to middle of ninth tergite; head of pupa lies on posterior
1 order of seventh tergite ; head has broken away from tergum and
f ppears in fig. 40 to be on gills but is not attached to them. The speci-
men was collected on 26 June 1960. On another nymph a chironomid
] upa adheres by posterior end to side of wing pad near second coxa;
lest of pupa is extended out laterally from tergum of nymph; head
extends to middle of seventh tergite over to one side above gills. One
specimen broken away from nymph collected 26 June 1960; ventral
surface of head and thorax not covered by membrane which covers
caudal end and tergum up to third segment where it is broken away;
membrane covers thoracic tergum; extends beyond head with crumpled
area and much silt for a length equal to length of head and thorax
combined. Same yellowish granular membrane as on larvae; ventrally
APRIL 1969] MAYO THRAULODES SPECIOSUS NYMPH
111
the membrane covers head capsule of larva of which the two halves
are flattened out on fourth abdominal sternite with tips bearing claws
of prolegs lying in center on unsclerotized area between fourth and
fifth sternites; membrane does not seem to cover the abdominal sternites
below fourth segment but envelopes caudal end.
A pupa in an early stage of development was collected 26 June 1960;
entirely encased in a colorless membrane except for head of larva which
is still free; there is no silt in membrane but a small patch on one side
of caudal end where: it had probably attached to nymph. Nymph on
which pupa had been attached with evidence of larvae having been on
both sides. Pupa witi wings and legs developed in early stage; abdomen
widened out. Claws of anterior and posterior prolegs are in membrane
covering pupa.
There is no hole in chitin of nymphs. The larvae have apparently
not fed on nymphs and are therefore not parasitic, but symbiotic. Pen-
nack (1953 : 650) states that ^^Symhiocladius and some species of Spani-
otoma are commensals on mayfly and stonefly nymphs.” The term
symbiotic is preferred in this paper. Other examples of symbiotic
relationships between chironomid larvae and mayfly nymphs are cited
by Johannsen (1937) as follows: larvae of Spaniotoma sp. E. group
Epoieocladius Zavrel were found clinging to the legs and gills of an
ephemerid nymph, Hexagenia recurvata. The larva of a European
species, Spaniotoma ephemerae has been described as living in symbiotic
relation with Ephemera vulgata. Spaneotoma {Dactylocladius) brevi-
palpis (Goetg.) of Europe has been found living under wing covers of
mayflies. Another symbiotic relationship is mentioned by Claassen
(1922) in which the larvae and pupae are carried under the wing pads
of mayfly nymphs belonging to the genus Rhithrogena.
Traver sent to the writer a Thraulodes nymph collected by F. Plau-
mann from Rio Jacutinga, Brazil, April 1962. This had a chironomid
pupa attached to the abdominal tergum. After being preserved in
alcohol the pupa dropped off, but it was evident that it had been attached
on the right side adjacent to the midline. It appears to have been
cemented to the abdomen with traces of cement still adhering to tergites
1—9. This does not resemble the silt deposited on the tergites of T.
speciosus where the larvae and pupae had been attached; neither does
it correspond in position on the abdomen to that of the larvae and pupae
on T. speciosus. The pupa does not appear to be encased in a membrane,
but a small amount of fine silt remains on head.
The writer is indebted to Jay R. Traver for her suggestions in the
preparation of this paper.
112
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Literature Cited
Claassen, P. W., 1922. The larva of a chironomid {Trissocladius equitans n. sp.)
which is parasitic upon a mayfly nymph (Rhithrogena sp.) Univ. Kans.
Sci. Bull., 1, 14: 395-405.
JoHANNSEN, 0. A., 1937. Aquatic Diptera III. Chironomidae : subfamilies
Tanypodinae, Diamesinae, and Orthocladiinae. Cornell Univ. Agr. Exp.
Sta. Mem., 205: 1-84, plates I-XVIII.
E lEFFER, J. J., 1925. Deux genres nouveaux et plusieurs especes nouvelles du
groupe des Ortho cladiariae {Dipteres, Chironomides) . Ann. Soc. Sci.
Bruxelles, 44: 555-566.
1 ENNAK, R. W., 1953. Freshwater invertebrates of the United States. Ronald
Press Co., New York, 1-769.
1 RAVER, J. R., AND G. F. Edmunds, Jr., 1967. A revision of the genus Thraulodes
{Ephemeroptera : Leptophlehiidae) Misc. Publ. Entomol. Soc. Amer.,
5 (8): 351-402.
Gall Wasp Infestations in Forest Trees,
Chiefly Pines, of California
(Hymenoptera : Eurytomidae)
George R. Struble
Pacif. SW. Forest and Range Exp. Sta.,
Forest Serv., U.S.D.A., Berkeley, Calif.
According to recently published reports, the phytophagous gall wasps,
Eurytoma spp. (Eurytomidae), attack introduced pines in California.
But unpublished notes^ show that these insects also attack native pines:
(Coulter pine, Pinus coulteri D. Don: Figueroa Mtn., Los Padres Natl.
Forest, 1937. Jeffrey pine, P. jeffreyi Grev. & Balf. : Mt. Laguna, Cleve-
land Natl. Forest, 1940; Lee Vining, Inyo Natl. Forest, 1958; Big Pine
Jdat, San Bernardino Natl. Forest, 1959; Prairie Forks, Angeles Natl,
l^orest, 1959. Ponderosa pine, P. ponderosa Laws.: Bogard, Lassen
Natl. Forest, 1959. Scotch pine, P. sylvestris L. : Ben Lomond, Santa
Cruz Co., 1960. Italian stone pine, P. pinea L. : Woodside, San Mateo
Co., 1968. Quaking aspen, Populus tremuloides Michx. : Lee Vining,
j-nyo Co., 1956. The first such infestations were found in Coulter pine,
E^inus coulteri D. Don in 1937. In addition, gall wasps have been found
attacking quaking aspen, Populus tremuloides Michx.
Among the introduced pines, the most recent infestations have been
1 Hopkins’ note series on file at Pacific Southwest Forest and Range Experiment Station, Forest
Service, U. S. Dep. Agric., Berkeley, Calif.
The Pan-Pacific Entomologist 45: 112-115. April 1969
APRIL 1969]
STRUBLE — GALL WASPS IN PINES
113
Fig. 1. Eurytoma sp. galls in sapwood of infested Coulter pine limb.
Fig. 2. Eurytoma sp. infestations cause swelling, pits, and gall formations in
Digger pine.
114
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Fig. 3. Larval form and setal pattern of: (A) Eurytoma tumoris (Stark and
ioehler, 1964) ; (B) Eurytoma sp. in Coulter pine (drawing by Donald DeLeon
n 1937).
•eported in Scotch pine, P. sylvestris L. (Stark and Koehler, 1964), and
Italian stone pine, P. pinea L. (Agr. Res. Serv., 1968) .
Most Eurytoma species that attack native pines have similar habits,
although their life cycles are not known. Their infestations cause galls
;o form in the living sapwood (Fig. 1). Healing tissue, recognized by
irregular swellings, surrounds the stem area holding galls. Deep pits in
the cambium area mark the presence of galls buried in the sapwood
(Fig. 2). The wood graining surrounding each gall resembles the
“bird’s-eye” configurations often observed in pines and other tree species.
APRIL 1969]
STRUBLE- — GALL WASPS IN PINES
115
The life history and damage of E. tumoris Bugbee, identified in 1962
(Stark and Koehler, 1964) in Scotch pine, provide the only source of
biological information on infestations in California pine species. The
infestations in Scotch and stone pine probably originated in Monterey
pine, suggesting a similar biology in this native host tree and in the two
introduced host trees planted nearby.
E. pachyneuron Gir., identified in 1959 from Jeffrey pine, and E.
tumoris are the only two species described as infesting pines in Cali-
fornia. Drawings of E. tumoris larvae infesting Scotch pine compared
with those of larvae infesting Coulter pine showed sharp differences in
the second through the sixth abdominal segments (Fig. 3) .
Gall-forming Eurytoma infestations found in indigenous and intro-
duced pines in California occurred in stems up to 4 inches in diameter.
Although the smaller stems in Christmas tree plantings (Stark and
Koehler, 1964) were heavily damaged, less than 10 per cent of the
trees were infested.
Thomas and Herdy (1961) reported E. calycis Bugbee attacking jack
pine, P. banksiana (Lamb.), in northwestern Ontario, Canada. Their
report is the only other published record of phytophagous Eurytoma
attacking pines in North America.
Literature Cited
Agricultural Research Service, U. S. Dep. Agr. 1968. Coop. Econ. Insect Rep.,
18(7) : 78.
Stark, R. W. and C. S. Koehler. 1964>. Biology of the gall wasp, Eurytoma
tumoris, on Scots pine. Pan-Pac. Entomol., 40(1) : 41-46.
Thomas, J. B. and H. Herdy. 1961. A note on Eurytoma calycis Bugbee
( Hymenoptera : Eurytomidae) occurring in shoots of jack pine {Pinus
banksiana Lamb.). Can. Entomol., 93(1): 34^39.
116
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
The Immature Stages of Gonomyodes tacoma Alex.
(Diptera : Tipulidae)^
C. Dennis Hynes
California State Polytechnic College, San Luis Obispo
Collection of the immature stages of Gonomyodes show them to be
s rikingly different from all other genera found within the tribe Erio-
p terini. The larvae and pupae of Gonomyodes tacoma were taken at the
type locality (Alexander, 1949), along the banks of the White River in
Mount Rainier National Park, Washington. All stages of immatures
\ ^ere found in quantity in silt impregnated leaf drift, and were especially
rumerous where the leaves were caught between the rocks through
\^hich the water moved slowly. Both larvae and pupae were found
cccasionally around rocks in sandy areas where they had been washed
c uring higher water conditions, but neither were ever found in the
swifter and deeper parts of the stream. The following descriptions
Mere based on ten last instar larvae and will serve for all but the first
iastar. The pupal description is based on one female and five male
specimens.
Gonomyodes tacoma Alexander
Last Instar Larvae. — Length 7.0-8.5 mm; dextrosinistral and dorsoventral di-
E meters both 0.5-0.6 mm. Cuticle colorless, covered with very short, pale yellow
pubescence, pencils of elongate setae not evident. Body cylindrical, slightly
tapered at anterior end, posterior end truncate. Spiracular disk with seven lobes,
those homologous with those found in other Eriopterini with thick setal pro-
tuberance at tip. Clear plate-like protuberance between ventral lobes, ventral and
lateral lobes, and lateral and dorsolateral lobes, none between dorsal and dorso-
lateral lobes; tuft of setae subequal to length of lobe forming setal fan around
inargins of outer half of lobe; each lobe with light yellowish-brown markings,
marking of dorsal lobe slightly bifurcate at base of lobe, remainder of markings
,LS indicated (Fig. 1). Spiracles very small (diameter = 0.017 mm), set widely
apart, outer rim light brown, center black. Anal lobes short, blunt, fused along
nidline to a terminal point. Head capsule typically eriopterine; length 0.332-
0.391 mm; dextrosinistral diameter at mandibular articulation 0.061 mm; dorso-
entral diameter at mandibular articulation 0.111 mm; anteclypeal fold arched
dorsad and cephalad from the anterior mandibular articulation; antenna cylin-
irical, an elongate papilla at tip curved mesad and ventrad; dorsal, lateral and
1 This investigation is supported by National Science Foundation Grant GB-4532.
->
Fig. 1. Spiracular disk of Gonomyodes tacoma. Fig. 2. Dorsal view of cauda of
male pupa of Gonomyodes tacoma. Fig. 3. Lateral view of pupa of Gonomyodes
tacoma.
The Pan-Pacific Entomologist 45: 116-119. April 1969
APRIL 1969] HYNES IMMATURE STAGES GONOMYODES
117
118
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
ventral bars curving sharply mesad at caudal ends, caudal ends spatulate; anten-
r al buttress curved mesad with no apparent break or suture between buttress and
junction of dorsal and lateral bars; clypeal region lightly sclerotized with frontal
legion membranous anteriorly; posterior fourth of frontal region sclerotized with
c ark stripe on either side of median suture, stripes pointed caudally, not joining
corsal bars; anterior margin of ventral bars not toothed, curving sharply mesad at
junction with mandibular strengthening ridge; two eye spots present on each side,
I'osterior spot nearly twice size of anterior spot; maxillae bilobed and blunt;
laandibles with standard slant for Eriopterini, strengthening ridge seen as simple
strip expanded slightly at midlength to dorsal tip, the ventral tip pointed; antennal
length 0.059 mm; width 0.017 mm.
Pupa . — Length 3.30-3.95 mm; width 0.5-0.8 mm. Body liglit yellow, darker
-when mature. Cephalic crest armed with two tubercles, each with one seta
cephalad and one ventrad. Pronotum standard; mesonotum (Fig. 3) with median
Carina sharp, mesonotal face before dorsal crest patterned with brown, whitish
vitta continuous with median carina to slightly heyond dorsal crest, face and crest
vith very small spines, more numerous medially. Mesonotal breathing horns
straight, delicate, base slightly expanded, stem directed anteriorly and laterally,
surface pitted; mesonotum patterned with brown, but pattern not constant.
Dorsum of abdominal segments two through seven unarmed, unmarked except for
ransverse row of small black spines on posterior margin of segments; sternal
areas of abdominal segment four with transverse row of spines; abdominal segments
ive through seven with transverse row of small spines along posterior margins
with middle third of each row devoid of spines; pleural region unmarked, unarmed,
spiracular opening prominent; wing pads end slightly beyond junction of abdominal
segments three and four. Eye sheath smooth; antennal sheath with scape slightly
enlarged; pedicle greatly expanded compared to flagellar sheath; flagellar
sheath crenulate on dorsal margin with crenulation ending four to five segments
before base of wing pad; marked grooves between eye sheaths, antennal sheaths,
and coronal suture. Leg sheaths ending at posterior margin of abdominal segment
four, prothoracic leg sheaths and metathoracic leg sheaths subequal, mesothoracic
leg sheaths slightly shorter. Male cauda (Fig. 2) with basistyle expanded,
slightly perceptable indentation separating outer dististyle from inner dististyle,
two sheaths of outer and inner dististyles not turned mesad as in many eriopterines.
Tergal arms with two large spines, dark at tips, directed laterally and dorsally, two
setae directed laterad at midlength of each spine of tergal arm. Area between
eighth abdominal segment and tergal arms with transverse foldings. Female cauda
with spiracular disk area similar to that pictured for male, with short anteapical
spine directed dorsally and laterally at the tip of each tergal valve.
The presence of the seven lobes of the spiracular disk of the larva
and the homologous structure of the pupa readily separate the immature
stages of this genus from the known immature stages of all other genera
of the tribe Eriopterini. The larva has several characteristics in common
with the genus Ormosia, particularly the subgenus Parormosia as repre-
sented by Ormosia {Parormosia) divergens. In both, the spiracles are
widely set apart, the surface area of the spiracles is very small, and the
setal fan at the tips of the spiracular lobes are constructed similarly.
APRIL 1969]
FENDER — NEW CALIFORNIA MALTHODES
119
Literature Cited
Alexander, C. P. 1949. Records and descriptions of North American craneflies.
(Diptera) Pt. VIII. The Tipuloidea of Washington, 1. Amer. Midland
Natur., 42(2) : 257-333.
New California Malthodes and Distribution Notes on
Other Species in California
(Coleoptera : Cantharidae) ^
Kenneth M. Fender
Linfield Research Institute, McMinnville, Oregon
The following new species and records are mainly from two large
collections of Cantharidae loaned me by Mr. Robert 0. Schuster of the
University of California at Davis and Mr. T. N. Seeno of the Bureau of
Entomology, California Department of Agriculture at Sacramento. In
addition are some records of collections of G. H. Nelson, Kansas City,
Missouri. For these loans, I am most grateful. Included, also, are
recent collection notes of mine. The distribution notes often extend the
previously known range of the species (Fender 1951, 1968) . Types of
the new species are deposited in the collections of the California Academy
of Sciences, San Francisco (CAS) or the University of California,
Davis (UCD).
Malthodes bissellarum Fender, new species
(Figs. 1, 2)
Piceous, head black, mandibles testaceous. Pronotum flavous with sides narrowly
black on anterior half, prosternal episternum flavous, seventh sternite and sixth
sternite at top of emargination flavous. Pubescence cinereous, fine, sparse, incon-
spicuous. Length 2.75 mm.
Male. — Shining. Head finely sparsely punctate in front of antennae, micro-
reticulate and finely punctured behind, eyes moderately large and prominent,
antennae of type partially concealed, second and third segments equal, fourth
evidently longer, intermediate segments about four and a half times as long as
wide. Pronotum transverse, nearly half again as wide as long, slightly narrower
than head, sides sinuately divergent from obtuse and explanate basal angles to
sharply rounded and narrowly reflexed anterior angles, basal and apical margins
finely beaded, lateral margins more widely so. Scutellum semicircular, finely
closely punctured. Elytra extending to base of sixth abdominal sternite, finely
rugose basally, becoming more coarsely so apically. Thorax beneath finely sparsely
punctured. Abdominal sternites alutaceous and finely punctured.
r This study was supported by National Science Foundation Grant GB-6283X.
The Pan-Pacific Entomologist 45: II9-I26 April 1969
120
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Figs. 1-10. Male terminal abdominal segments. Fig. 1. Malthodes bissellorum,
ventral view. Fig. 2. same, lateral view. Fig. 3. Malthodes arieticornus, ventral
view. Fig. 4. same, lateral view. Fig. 5. Malthodes bifurculus, ventral view. Fig.
6. same, lateral view. Fig. 7. Malthodes immodestus, ventral view. Fig. 8. same,
lateral view. Fig. 9. Malthodes angustifurcus, ventral view. Fig. 10. same, lateral
view.
Male Terminalia.— Ventral view: sixth abdominal sternite broadly deeply
emarginate, apex of emargination rounded; seventh sternite terete, elongate,
moderately slender, slightly expanded apically, apex deeply narrowly notched,
apex of notch rounded, tips rounded. Lateral view: last three tergites normal;
sixth sternite normal; seventh sternite elongate podiform, apically ascending;
dorsal accessory process dual, a somewhat foliate thin plate, beneath and abutting
this, a bispinose thicker appendage.
APRIL 1969]
FENDER — NEW CALIFORNIA MALTHODES
121
Holotjpe male, Kings River Canyon, California 25 June 1948,
collected by A. T. McClay (UCD) ,
This species, with the last three tergites normal, seventh sternite
terete and apical notched, keys to my group five of the genus. From the
other known members of that group it may be readily recognized by the
podiform seventh sternite. The other members do not have the seventh
sternite enlarged apically. Named for the Fred Bissell family who have
added numerous specimens to my collection.
Malthodes arieticornus Fender, new species
(Figs. 3, 4)
Piceous, head black behind antennae, mandibles testaceous apically. Pronotum
flavous, anterior margin narrowly, lateral margins and broad median longitudinal
infuscate stripe black, extending from base to apex, varying in breadth from
specimen to specimen. Prosternal episternum flavous, margins black. Abdominal
sternites through fifth with apical margins rather broadly testaceous, apical half
of seventh sternites somewhat paler, spongiose portion of ultimate and penultimate
tergites whitish. Legs piceous to piceous with apices of femora narrowly and
basal half or more of tibiae testaceous. Pubescence cinereous, very fine, sparse
and inconspicuous. Setiferous hairs of sheath of seventh sternite black. Length
3.5 to 4 mm.
Male. — Shining. Head smooth in front of antennae, finely sparsely punctured
behind, eyes moderately large and prominent, antennae extending to base of sixth
abdominal sternite, second and third segments equal, fourth segment about one
third longer, intermediate segments about three and a half times as long as wide.
Pronotum transverse, lateral margins shallowly concave, diverging from nearly
rectangular hind angles to rounded anterior angles, marginal bead fine, coarser
on sides near angles and along basal margin, disc finely sparsely punctured.
Scutellum subtriangular, apex sharply rounded, finely closely punctured. Elytra
short, extending to about middle of second abdominal sternite, finely rugose basally
becoming more coarsely so to tumid and finely punctured apices. Thorax beneath
finely, sparsely punctured. Abdomen dull, little more coarsely punctured.
Male Terminalia. — Ventral view: sixth abdominal sternite broadly deeply
emarginate, apex of emargination sharply rounded, sides of emargination concave;
seventh sternite extending beyond sixth by almost twice length of sixth, wide
basally, evenly narrowed to deeply triangularly notched tip; beneath basal half
of seventh sternite a translucent, shallowly emarginate skirt with apical margin
armed with long setigerous black hairs. Lateral view: seventh sternite evenly
arcuately ascending apically, evenly narrowed from base to tip; translucent basal
skirt curved above sternite; ultimate and penultimate tergites produced and
somewhat arched, on ventral surfaces a common spongiose area, dorsal accessory
process strongly chitinized, somewhat arched and digitate, apically pointed,
bearing on ventral surface a less strongly chitinized process narrowing to suddenly
elevated and filamentous tip, ventral accessory process strongly chitinized, thick
basally, narrowing evenly to suddenly bent up tip.
Female. — Similar to male. Eyes smaller and less prominent; antennae extending
to base of metacoxse, intermediate segments about 2.7 times as long as wide, median
122
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
broad stripe of pronotum usually narrower to occasionally absent. Elytra shorter,
pitending to middle of second visible abdominal sternite. Terminal abdominal
segments normal for the sex.
Holotype male, allotype female and twenty paratypes Ramshorn
I OREST Camp (for which species is named), Tahoe National Forest,
California, 9 June 1968, collected by K. M. and W. M. Fender (CAS).
This species should by the conformation of the seventh sternite key
to group I. The arched last tergites would place it in group V. However
t ie exposed accessory processes remove it from group I. The presence
cf a ventral skirt outside the seventh sternite and the spongiose area
common to the ventral or inner face of the ultimate and penultimate
tergite remove it from any previously known group.
Malthodes bifurculus Fender, new species
(Figs. 5, 6)
Black, shining. Mandibles testaceous. Pronotum flavous, anterior and basal
margins widely darker, lateral margins widely black, elytra becoming paler
toward apices. Apices of first three visible sternites narrowly testaceous towards
s des, seventh sternite testaceous becoming infuscate beyond middle, produced
s de pieces of penultimate tergite testaceous. Pubescence cinereous, fine sparse
a ad inconspicuous. Length 2.5 to 3.5.
Male. — Shining. Head slightly narrower than pronotum, finely sparsely
punctured, eyes moderately wide and prominent, antennae long and slender, ex-
tending to tips of wings in repose, second segment seven-tenths as long as third
and three-fifths as long as fourth, intermediate segments about four times as
long as wide. Pronotum transverse, widest near rounded anterior angles, sides
feebly sinuately convergent to obtusely rounded hind angles, sides shallowly
reflexed near each angle, apical margin very finely beaded, basal margin a little
more coarsely so, disc finely, very sparsely punctured. Scutellum wider than long,
s ubtriangular, apex rounded, finely moderately closely punctured. Elytra extending
to base of seventh sternite, a little more coarsely closely punctured basally,
becoming finely rugose apically. Thorax beneath finely sparsely punctured. Basal
abdominal sternite transversely microstrigulose. Femora of all legs with virtually
r o pubescence.
Male Terminalia. — ^Ventral view: sixth sternite broadly deeply emarginate,
e inargination shallow to near middle, abruptly deepened to truncated apex;
seventh sternite unusually elongate and slender, extending beyond produced
t srgites by nearly half its length, broadly deeply forked, apex of furcation rounded,
t ps of forks rounded. Lateral view: sixth sternite short, somewhat oblique
seventh sternite unevenly sinuately ascending towards apex, becoming broadly
f attened at middle, tips sharply rounded; sides of penultimate tergite sub-
t iangularly produced downward and apically somewhat caudad tips acute.
Female. — Similar to male, head narrower than pronotum, eyes small and less
irominent, antennae extending nearly to elytral apices, intermediate segments
about two and a half times as long as wide, apical abdominal segments normal
f^r the sex.
APRIL 1969] FENDER — NEW CALIFORNIA MALTHODES
123
Holotype male, allotype female and one male and two female para-
types; Lake Lagunitis, Marin County, California, 30 March 1961,
collected by D. Q. Cavagnaro (UCD). One male paratype, Davis,
Yolo County, California, May 11, 1964, collected by Serje Seminoff.
The paratype from Davis differs from the others in having the
pronotum flavous with sides rather broadly black at the anterior angles.
This species is most closely related to M. visceratus Fall and M.
spinicaudus Fender. It may be separated from each of these by its
larger more slender, more widely, deeply forked seventh sternite which
in lateral view is more widely flattened. Too, the side pieces of the
penultimate tergite are narrower and apically more acute. Malthodes
spinicaudus also has an acute spine medially on the dorsal surface of the
seventh sternite.
Malthodes immodestus Fender, new species
(Figs. 7, 8)
Head piceous, elytral apex paler, mandibles testaceous, underside of basal
antennal segment paler. Pronotum yellow. Scutellum and elytra dark testaceous,
prosternum testaceous, mesosternum, metasternum and legs dark brown, abdominal
sternites testaceous, segments one to five becoming infuscate towards sides, the
lateral margins flavous, apices of seventh sternite becoming infuscate beyond
middle, penultimate tergite pale yellow. Pubescence cinereous, fine, sparse and
inconspicuous. Length 3.25 to 3.5 mm.
Male. — Shining. Head finely punctured in front of antennae, rather sparsely
but coarsely pustulate behind; eyes very large and prominent (ocular index 45) ;
antennae slender, extending to base of sixth abdominal sternite, segments two
to four progressively longer, intermediate segments four and a half times as long
as wide. Pronotum narrower than head, transverse, sides nearly straight, converging
somewhat from rounded anterior angles to obtusely rounded basal angles, marginal
bead thin, a little wider near angles, disc finely sparsely punctured, scutellum
triangulcu-, apex sharply rounded, finely closely punctured. Elytra extending to
base of sixth sternite, finely rugose to tumid and finely punctured apices. Thorax
beneath smooth, and hare. Basal abdominal sternites rugose.
Male Terminalia. — Ventral view: sixth sternite widely, deeply, arcuately
emarginate, seventh sternite elongate, stout, extending heyond sixth by about one
and a half times length of sixth, forked at about apical two thirds, narrowed to
furcation, forks twisted about 45 degrees and flattened into spatulate plates;
basal half with broad, elongate, shallow oval impression. Lateral view: seventh
sternite slender basally, gradually expanding to broadly beveled beneath apex,
arcuately ascending apically. Side pieces of penultimate tergite bent down and
caudad into seemingly crumpled oval processes. Ultimate tergite small, rounded;
ventral accessory process slender, feebly sinuate, rod-like, apically ascending
above seventh sternite.
Female. — Unknown.
Holotype male. Running Springs, 15 miles north of Redlands, San
Bernardino Mountains, San Bernardino County, California, 5 June
124
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
1945, 6000 feet, on Ceanothus integerrimus Hooker and Arnott, collected
ly G. H. Nelson (CAS). One paratype, Wrightwood, San Bernardino
County, California, June 28, 1964, collected by J. S. Buckett.
The paratype is darker than the type, the head being nearly black
£nd the scutellum and elytra piceous, the dark areas towards the sides
cf the basal five sternites are more pronounced.
This species keys out to my group X, but it can’t be closely associated
\ ith any member of that or any other group. It can readily be differenti-
ated from any other known North American species by the spatulate
£nd twisted forks of the seventh stemite of the male.
Malthodes angustifurcus Fender, new species
(Figs. 9, 10)
Black, mandibles testaceous, pronotum flavous, large median X-shaped black
soot, lateral margins narrowly black, widely so just behind anterior angles; elytra
ficeous with apices black; seventh sternite, median portion of sixth sternite and
produced side-pieces of penultimate tergite testaceous.
Male. — Head shining, wider than pronotum, finely sparsely punctured, a
1 ttle more coarsely so behind antennae, eyes large and prominent, antennae slender,
extending to tips of wings in repose, segments two to four progressively longer,
intermediate segments about three times as long as wide. Pronotum shining,
t .ansverse, about one-third wider than long, anterior margin narrowly arcuate,
finely beaded, anterior angles rather sharply rounded and explanate, lateral
margins sinuately narrowed to prominent and rounded hind angles, sides beaded
i 1 apical half, unbeaded in basal half, basal margin strongly arcuate, unbeaded
medially, becoming deeply so towards angles, disc finely sparsely punctured,
elytra parallel sided, extending to about base of sixth abdominal sternite, finely
1 agose basally, becoming a little more coarsely so apically, thorax beneath rather
coarsely closely punctured. Abdominal sternites more finely so.
Male Terminalia. — Ventral view: sixth sternite widely deeply emarginate,
e inargination U-shaped. Seventh sternite elongate, slender, forked at about apical
t rird, forks narrowly separated, slightly convergent apically, tips of forks rounded,
spex of furcation rounded, sides of penultimate tergite slightly enveloping seventh
sternite near middle. Lateral view: seventh sternite straight in basal half,
lisinuately ascending to sharply rounded tips in apical half. Side pieces of
penultimate tergite abruptly descending as rather broad subovate plates, on plates
£11 elongate venose ridge arising medially at top, splitting and diverging apically.
Ultimate sternite small subtriangularly rounded, feebly produced.
Holotype male, Echo Lake, El Dorado County, California, 23 July
1955, E. I. Schlinger collector (UCD).
Most closely related to M. contortus Fender, this species can readily
1 e separated from that species by the seventh sternite being bisinuately
lent beyond middle in lateral view, the last abdominal tergites less
strongly extended, the produced side pieces with a venous forked ridges.
APRIL 1969]
FENDER NEW CALIFORNIA MALTHODES
125
In ventral view, the forks of the seventh sternite are less widely separated,
converging somewhat near tips.
California Distributional Records
Malthodes jracidus Fender. Crystal Lake, Los Angeles County.
Previously known from Donner Pass and Fallen Leaf Lake.
Malthodes evanidus Fender (?). Sagehen Creek, near Hobart Mills,
Nevada County and near Clayton, Contra Costa County. This species
was described from Duck Creek Forest Camp, Dixie National Forest,
Utah. If this is the same species, and there is little to suggest otherwise,
this is a considerable westward extension of the range of the species.
It is thus a new record for the state.
Malthodes basalis Fall. 4 miles west of Woodward, Alpine County.
Malthodes pictithorax Fender. Knight’s Landing, Yolo County;
Putah Creek, Yolo County.
Malthodes vigilans Fall. Echo Lake, El Dorado County. Previously
recorded from the San Bernardino Mountains and Dalton Creek, Fresno
County.
Malthodes pictus Fender. Previously known from the types collected
at Doner Pass, new records are: Davis, Yolo County; Mt. Diablo,
Contra Costa County; 4 miles west of Zamora, Yolo County; 5 miles
northwest of Winters, Yolo County; Michigan Bar, Sacramento County
and Rutherford, Napa County.
Malthodes vapidus Fall. Fall (1919) described this species from
Lake Tahoe and noted the pronotum as being yellow. He knew only the
type. Several examples were studied from Fallen Leaf Lake and Tallac,
El Dorado County. Whereas there was some variation in the pronotal
coloration, I found none in which I could call the pronotum yellow.
At best they could be described as pale brownish-yellow. Three examples
were recently seen from Silver Lake, Amador County, in which the
pronota were pale brownish-yellow, each with a large median black
spot. About 12 specimens from Little Greyback Mountain, Del Norte
County have the pronota pale yellowish-brown. No structural differences
could be found to separate these. This extends the range of this species
from the Lake Tahoe area to the northwest corner of the state.
Malthodes mollis Fall. New localities are: Courtland, Sacramento
County; 8 miles south of Trinidad, Humboldt County.
Malthodes williami Fender. Described from McBride Spring Forest
Camp, Mount Shasta, this species was abundant at McArthur-Burney
Falls Memorial, Shasta County on June II, 1968.
Malthodes oregonus Fender. Little Greyback Mountain, Del Norte.
County. This is the first California record for this species.
126
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Malthodes fusculus (LeConte). Previously recorded west of Novato,
]V arin County. New Records: Alameda, Alameda County; near
Clayton, Contra Costa County.
Malthodes bicurvatus Fall. Forestville, Sonoma County.
Malthodes visceratus Fall. Davis, Yolo County; Twin Peaks, San
Francisco County.
Malthodes spinicaudus Fender. New localities: Echo Summit, El
E orado County; Huntington; Silver Lake, Amador County; Yuba Pass,
Sierra County; 3 miles southeast of Quincy, Plumas County.
Malthodes blackwelderi Fender. Previously known only from the
type, collected in the Argus Mountains, Death Valley, an additional
sjiecimen has been examined from High Grove, Riverside County.
Literature Cited
Fall, H. C. 1919. The California Species of Malthodes. Ann. Entomol. Soc.
Amer., 12: 31-42.
Fender, K. M. 1951. The Malthini of North America (Coleoptera-Catharidae) .
Amer. Midland Natur., 46(3) : 513-629.
1968. New Malthodes from Western North America (Coleoptera : Can-
tharidae). Northwest Sci., 42(3) : 108-111.
A. New Species of Cordulegaster from the Great Basin Region
of the United States
(Odonata : Cordulegasteridae)
Robert William Cruden
Department of Botany, University of Iowa, Iowa City
Literature dealing with the distribution of Cordulegaster in the Great
I asin presents two quite different pictures. The first shows the distribu-
t on of C. dorsalis Hagen restricted to regions west of the crest of the
Sierra Nevada and Cascade Range (Kennedy, 1917; Fraser, 1929; La
l ivers, 1941; and Walker, 1958). Further, Fraser and La Rivers sug-
gested that C. dorsalis is replaced east of the Sierra Nevada by C.
erroneoA Hagen. This latter species was previously reported only from
tie Appalachian region of the eastern United States. In the second
picture, C. dorsalis extends from California east through Nevada to
Utah (Needham and Westfall, 1955; Musser, 1962) and Wyoming
1 The greek -gaster is feminine and the adjectival specific epithet should agree with it in gender,
hence erronea rather than erroneus.
The Pan-Pacific Entomologist 45: 126-132. April 1969
APRIL 1969]
ORUDEN A NEW CORDULEGASTER
127
(Smith and Pritchard, 1956). Needham and Westfall side step the
problem of C. erronea by attributing the western records to cases of
incorrect determination and suggest that records of C. erronea from
Nevada and Utah need verification.
The purpose of this paper is to clarify the distribution pattern of
Cordate faster in the Great Basin. Underlying the present differences
of opinion concerning distribution patterns in that region is a here-
tofore unrecognized and undescribed species. First, I will consider the
initial report of C. erronea from the Great Basin. This is followed by the
description of a new species and comparison of the new species with
the closely related C. dorsalis.
The initial steps leading to todays confusion concerning C. erronea
were formalized by Fraser (1929). Although Fraser was apparently
a victim of circumstance, he must still bear the responsibility for creating
the problem. Fraser (1929) credits Selys as the author of “C. erroneus”
and assumed that Selys had annotated particular specimens from the
western United States in the Brussels Museum. He states, “The original
description is very brief and appears to have been made from a single
specimen, although not one of the nine males in the Selysian collection
agree entirely with it.” There is suggested in Fraser’s words that he
thought one of the nine males might have been a type. Obviously
Fraser overlooked several things: (1) Selys credits the authorship of
C. erronea to Hagen (1878, p, 688-89), (2) The description is based
on a minimum of three, not one specimen, (3) the holotype and two
paratypes, including two males and one female are in the type collection
of the Museum of Comparative Zoology, Harvard University (Evans,
pers. commun.). It seems quite unlikely that Selys had either seen or
possessed material in his collection of C. erronea at the time of its
description.
The notation “(Coll. Selys, Hagen)” in the original description
would indicate that Selys expected to have at least one of the type speci-
mens for his collection and would explain why he prepared a label for
the species and presumably left a space for it in a cabinet drawer. It
has been said that when the Selysian collection was reorganized to
conserve space, some specimens were moved from one drawer or space
to another without moving the species labels (Gloyd, pers. commun.).
The supposition being that someone other than Selys moved the series
of nine males cited by Fraser (1929, p. 134) into a space labeled in
Selys’ hand “C. erroneus.’’’’
Fraser’s redescription of C. erronea based on the nine males from
western North America in the Brussels Museum and one male from
128
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
4
Figs. 1-4. Cordulegaster deserticola Cruden. Fig. 1. Holotype: male. Lateral
view of thorax and abdomen. Xl-4. Fig. 2. Holotype: male. Dorsal view of abdomen
>'1.4. Fig. 3. Allotype: female. Lateral view of thorax and abdomen. Xl-4. Fig.
4 Allotype: female. Dorsal view of abdomen. Xl-4.
California in the British Museum should he rejected. Specimens from
the Great Basin in Utah and California are specifically distinct from
loth C. erronea and C. dorsalis. A new species is proposed below to
include such specimens.
Cordulegaster deserticola Cruden, new species
Male.— total length 73-79 mm; abdomen 53-59 mm long; hindwing 43-48
r.im long.
Head: Labrum yellow with narrow brown margin; anteclypeus dark brown
to black; postclypeus yellow; frons yellow, sometimes with a smudge of brown
at top of vertical surface as in C. dorsalis; vertex black with yellow dashes above
antennal sockets; occiput yellow to brown with long pale hairs at rear; eyes
riilky blue in life; rear of head yellow.
Prothorax: brown with dorsum of front and rear lobes yellow, two L-shaped
yellow dashes on center lobe.
Thorax (Fig. 1) : black with yellow markings; antehumeral stripe parallel
sided or slightly widened or tapered downward, in some specimens nearly covering
front of thorax; lateral stripes quite wide; a small spot dorsally between ante-
APRIL 1969]
CRUDEN A NEW CORDULEGASTER
129
humeral stripe and first lateral stripe ; a narrow continuous or broken stripe
between lateral stripes. A thin dash beneath spiracle; small spots above 1st and
2nd coxa; these coxae each with a spot on outside; rest of legs dark brown to
red brown.
Wings: Costa yellow to beyond stigma, nearly to tip; triangle two celled in
both wings; anal loop with 4-5 cells (in one wing 6) ; antenodal crossveins
16-20/10-14; postnodal crossveins 10-14/10-14. Second thickened antenodal
crossvein 5-7. (One male from Zion National Park with 15 antenodal crossveins
in one fore wing and 9 postnodal crossveins in one hind wing).
Abdomen (Figs. 1, 2) : Segment 1 brown, 2-10 black, marked with yellow as
follows: 1) a horse shoe or nearly closed ring on side; 2) pale spot on oreillets,
triangular spot at rear angle, large spot on dorsum with two elongate spots near
posterior edge; 3) large spot nearly covering middle half of dorsum, elongate
spot near posterior edge, streak along ventral margins; 4-6) large oval spots
which fuse on dorsum, covering about middle % of segment, small spots along
posterior margin; 7) spots similar to 6 with narrow extension downward to
ventral margin, thus encircling segment, posterior spots greatly reduced; 9)
irregular shaped spot with projection to rearward, not meeting on dorsum; 10)
two large spots, the upper not meeting on dorsum.
Caudal appendages: black, superiors with prominent downward pointing tooth
toward base and smaller, basally directed tooth near middle; inferior appendage
% as long as superior.
Female (Figs. 3, 4). — length including ovipositor 76-81 mm; abdomen 57-61
mm; hind wing 47-49 mm.
Color pattern similar to male with spots larger. Spots on 1-8 yellow, on 9 and
10 whitish yellow. On segment two, anterior spots fused; segment 3, the spot
continues downward to meet streak along ventral margin.
Wings: Anal loops of females examined from California have 5 & 7, 9 & 9,
and 9 & 11 cells respectively, those from Utah 5 & 6, 7 & 7.
The haploid chromosome number of C. deserticola is 13. This agrees
with other species in Cordulegasteridae which have been studied cyto-
logically (Cruden 1968, and unpubL).
Holotype Male, Batchelder Spring, Inyo County, California, ca.
6,000 feet; 29 July 1967, R. W. Cruden 67-885. To be deposited at
UMMZ. Allotype female. Antelope Springs, Inyo County, California,
ca. 5,700 feet, 29 July 1967, R. W. Cruden 67-892. To be deposited
at UMMZ.
Paratypes. — CALIFORNIA: Inyo County: Antelope Springs: male and female,
15 June 1961, C. A. Toschi, (Calif. Insect Survey, Berkeley) ; male, 29 June 1961,
G. I. Stage (CIS) ; male 29 June 1961, J. Powell, (CIS) ; male, 10 July 1961,
R. M. Bohart (Univ. Cal. Davis) ; 2 males, 29 July 1967, R. W. Cruden (pers.
col.; UMMZ). Toll House Spring ( =:Batchelder Spring) 4 mi W of Westgard
Pass: female, 19 June 1961, J. Powell (CIS) ; 2 males, 29 July, R. W. Cruden
(pers. col.; UMMZ). Independence, male, 17 June 1937, (CIS). UTAH: Sevier
County: 3 males, above Clear Creek, 5 -j- miles east of Cove Fort, 7 July 1960,
G. G. Musser (UMMZ) ; 2 males and 1 female. Service Berry Creek, Monroe
Mt., elev. 6,500 feet, 15 June 1960, G. G. Musser (UMMZ). Salt Lake County:
130
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
male and female, Red Butte Canyon, stream above Ft. Douglas, reared from nymphs,
J Musser (UMMZ). Washington County: Zion Nat. Park: male, near Temple
oE Sinawava, 30 July 1933, C. J. D. Brown (UMMZ) ; 2 males, below Weeping
lock, 17 June 1968, R. W. Cruden (CIS, pers. col.).
Cordulegaster deserticola, in California, is found in springs in the
c esert regions east of the Sierra Nevada. The Inyo County localities
£re in shad-scale scrub between 4,000 and 6,000 feet. In Utah, this
species inhabits streams in mountainous areas between 4,000 and 6,500
feet. Unfortunately the specimens of C. erronea from Nevada (La Rivers,
1941) were destroyed in a fire in the early 1940’s (La Rivers, pers.
commun.). It seems probable that those specimens would now be
leferred to C. deserticola.
Two distinct forms are represented in the material studied. A majority
of the Utah specimens fall in the upper limits of size for the species.
' he antehumeral stripe of these specimens is more divergent and tends
to be slightly tapered or wedge shaped. The narrow stripe between the
two lateral stripes is usually reduced to three irregular dashes. The
dorsal, posterior spots extend to segment 6, rarely to segment 7. In
nearly half the male specimens segment 10 was unmarked. Although dis-
inct forms are found on opposite sides of the Great Basin it is premature
to propose subspecific status for the two populations. It seems likely
hat geographically intermediate populations will eventually be found.
J^ormal subspecies would obscure clinal variation if the geographically
intermediate populations were also morphologically intermediate.
Cordulegaster deserticola is easily distinguished from other western
species of Cordulegaster and the eastern C. erronea. A western species,
C. diadema Selys, and C. erronea belong to a group of species charac-
terized by narrow yellow bands which encircle each segment. Further,
he backs of the eyes of C. erronea are black, an unusual condition in
the genus. Cordulegaster erronea is known only from the Appalachian
region of the eastern United States.
Although there are no obvious structural differences between C.
dorsalis and C. deserticola they are easily separated on color pattern,
ecology and distribution. Individuals of C. deserticola are more spotted
than those of C. dorsalis. Specimens of C. deserticola have yellow on
the rear lobe of the prothorax, a continuous or broken stripe between
the two lateral stripes on the thorax, posterior spots on abdominal
segments 3 to 6 or 8, and a spot which extends downward and en-
circles segment 7. These yellow markings are usually lacking in
individuals of C. dorsalis.
The distribution ranges of the two species are allopatric. The geo-
graphical isolation is associated with ecological differences. In Cali-
APRIL 1969]
GRUDEN A NEW CORDULEGASTER
131
fornia, C. dorsalis usually inhabits streams west of the crest of the Sierra
Nevada and Cascade Range which are not subject to continuous or
repeated winter freezing. Cordulegaster deserticola is found east of
these mountains. I have collected it at two locations, both spring fed
streams which are subject to winter freezing. It is likely that the Utah
streams inhabited by C. deserticola are also subject to winter freezing.
Perhaps the differences noted by Musser (1962) between Utah ^‘deserti-
cokd’ nymphs and the C. dorsalis nymphs described and illustrated by
Walker (1958) are related to the different environmental conditions
experienced by the nymphs.
Despite the lack of structural differences in the adults, the eco-
geographical disjunction is apparently an effective isolating mechanism.
Cordulegaster dorsalis can reach the Great Basin (La Rivers, 1941).
Also in northern California, where the crest of the mountains breaks
down, C. dorsalis has successfully crossed and become established east
of the crest. This shows that the mountains are not an absolute barrier
to migration and establishment. Secondly, it emphasizes the general
failure of C. dorsalis to become established on the Great Basin side of
the Sierra Nevada and Cascade Range.
I believe there has been sufficient time and opportunity for C. dorsalis
to become established in the Great Basin if it were adaptively and com-
petitively able to do so. The species failure to successfully invade desert
habitats is strong evidence to support the idea of ecological isolation
between C. dorsalis and C. deserticola.
I suggest that the coloration differences between western and eastern
C. deserticola with respect to C. dorsalis are evidence for a past sepa-
ration and ecological divergence of a common ancestor into two popu-
lations. Later the two populations became sympatric although occupying
quite different habitats. At this time selection for behavioral and
other recognition characters would have been of great adaptive value
in maintaining genotypes already adapted to quite different habitats.
Thus in the western Great Basin selection favored amplification or
exaggeration of the C. deserticola color pattern. To the east no such
selection pressures were experienced and in Utah the color pattern is
superficially still like that of C. dorsalis.
In general, the distribution of Cordulegaster in the western United
States is as follows: C. dorsalis, with some exceptions, west of the
Sierra Nevada and Cascade Range from British Columbia south into
Baja California; C. deserticola in the Great Basin from eastern California
and western Nevada east to Utah; and C. diadema, southern California
east through Arizona to southern Utah and south into Mexico.
132
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Acknowledgments
I would like to thank Dr. George Bick, St. Mary’s College, Notre
I*ame, Indiana and Mrs. Leonora K. Gloyd, University of Michigan,
/.nn Arbor, for their critical suggestions concerning the manuscript.
The illustration was prepared by Mrs. Ann Schoner, Iowa City. This
study was supported in part by an Old Gold Summer Faculty Research
I ellowship from the University of Iowa.
Literature Cited
CAuden, R. W. 1968. Chromosome numbers of some North American dragonflies
(Odonata). Can. J. Genet. CytoL, 10 : 200-214.
Iraser, F. C. 1929. A revision of the Fissilabioidae, Pt. 1, Cordulegasteridae.
Mem. Ind. Mus., 9: 69-167.
Hagen, H. 1878. In Selys. Quatriemes additions an Synopsis des Gomphines
(suite et fin), Legions: Lindenia-Chlorogomphus — Cordulegaster et
Petalura. Bull. Acad. Belg., (2 ) 46 : 658-698.
l.ENNEDY, C. H. 1917. Notes on the life history and ecology of the dragonflies
(Odonata) of central California and Nevada. Proc. U. S. Nat. Mus.,
52: 483-635.
La Rivers, I. 1941. Additions to the list of Nevada dragonflies (Odonata).
Entomol. News, 52: 126-130.
Musser, R. J. 1962. Dragonfly nymphs of Utah (Odonata : Anisoptera) . Univ.
Utah Biol. Ser., 12(6) : 1-66.
Needham, G. and M. Westfall. 1955. A manual of the dragonflies of North
America (Anisoptera). Univ. Calif. Press, Berkeley, 615 p.
Smith, R. F. and A. E. Pritchard. 1956. Odonata — in Usinger, R. L. (ed.)
Aquatic insects of California. Univ. Calif. Press, Berkeley, 508 p.
^V^alker, E. M. 1958. The Odonata of Canada and Alaska, 11. Univ. Toronto
Press, Toronto, 318 p.
Synonymy of Dryocosmus attractans (Kinsey) and
Callirhytis uvellae Weld
(Hymenoptera : Cynipidae)
D. Charles Dailey
University of California, Davis
The ecological similarity of Dryocosmus attractans (Kinsey) (1922)
and Callirhytis uvellae Weld (1944) and the indistinct separation of
American species of these genera, based on scutum texture, suggested
they might be the same species. Type insects and galls of both were
examined and on the following evidence they are synonymized, the
name becoming Callirhytis attractans (Kinsey).
The Pan-Pacific Entomologist 45: 132-134. April 1969
APRIL 1969] DAILEY CALIFORNIA CYNIPID SYNONYMY
133
Galls. — Both descriptions indicate the monothalamous bud galls
develop in the spring on Quercus wislizetiii A. DC., often more than
one in a bud. They are marked with red and are about 2 mm in diam-
eter by 3 mm long. Kinsey collected galls on 2 April 1920 at Redding,
Shasta County, California with emergence occurring later. Weld found
his at Lakeport, Lake County, and Ukiah, Mendocino County, California
on 10 and 12 May 1922 respectively, emergence also was later. The
latter two locations are about 75 miles closer to the ocean, 500 to 1,000
feet higher in elevation, and usually cooler than Redding which helps
explain the difference in emergence dates. A series I reared from
Folsom Lake, Placer County, California emerged 13 to 21 April 1967.
The emergence time, galls and habitat agree closely with Kinsey’s
description, including the sticky secretion the galls produce which is
attractive to ants. The one D. attractans cotype gall in the California
Academy of Sciences showed double cavities when dissected, the lower
being the larval cell, similar to C. uvellae galls figured in Houard ( 1940,
1946) and Weld (1944).
Galls from Ukiah which Weld thought to be D. attractans were
figured by Houard (1935). There is one female in the U. S. National
Museum collection (Hopk. 15640*^) reared from similar galls submitted
by Weld. When he recognized this was not D. attractans he assigned
it field number 1690. Thus the apparent contradiction in Weld (1957)
stating he had not recognized D. attractans in the field and the reason
for the note on figure 194 “This is not D. attractans which secretes
copius honeydew.”
Insects. — Kinsey’s 13 males were imperfect, having issued, died, and
molded in the rearing container. The type probably is the cleanest and
most nearly whole specimen. The small carina separating the foveae of
the type is not typical for the species. The cotypes examined do not
have the carina and it is present on only one male from my series.
Weld’s notes on the D. attractans type indicate it fits in Dryocosmus
though no mention of scutum texture is made. As only males were known
and generic keys are based on females, and as sculpturing of males is
sometimes smoother, his generic assignment was speculative. The C.
uvellae description (mesoscutum microcoriaceous) would by Weld’s
own key place it in Dryocosmus though he placed it in Callirhytis and
it agrees with other species in the latter genus. His insects were not
described until 1944 and apparently had faded to brown. Disregarding
color, if both D. attractans and C. uvellae are keyed in Weld’s manu-
script species keys they run to the same couplet depending on which
generic key is used.
134
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Lyon (personal communication) has shown Dryocosmus grumatus
\7eld to be the alternate generation of Callirhytis serricornis (Kinsey)
and as more life histories are worked out, generic concepts will be
clarified. Since Lyon has taken a C. attractans female ovipositing in
tie underside of a leaf of Quercus agrifolia Nee (new host record) at
l a Canada, Los Angeles County, California, the unknown alternate
i nisexual generation probably develops in a leaf gall.
Acknovi^ledgments
The author is grateful to Dr. J. G. Rozen, Jr. of the American Museum
of Natural History for the loan of the D. attractans holotype, H. B.
l^eech of the California Academy of Sciences for the loan of D. attractans
cotypes. Dr. R. M. Bohart of the University of California at Davis for
assistance in obtaining specimens, and R. J. Lyon of Los Angeles City
College for use of the Weld collection and notes, and assistance in the
study.
Literature Cited
Houard, C. 1935. Marcellia 28(4-6) : 162, figs. 191-194.
1940. Les zoocedidies des plantes de I’Amerique du Nordi, p. 413, figs.
1641-1645. Privately printed, Paris.
1946. Les collections cecidologiques du laboratoire d’entomologie du Museum
d’Histoire Naturelle de Paris. Ann. Entomol. Soc. France, 112: 54,
figs. 298-300.
&NSEY, A. C. 1922. New Pacific Coast Cynipidae. Bull. Amer. Mus. Natur.
Hist., 46: 279.
W^ELD, L. H. 1944. New American cynipids from galls. Proc. U. S. Nat. Mus.,
95: 23.
1957. Cynipid galls of the Pacific Slope. Privately printed, Ann Arbor,
Michigan.
APRIL 1969] ZIMMERMAN FRANCIS XAVIER WILLIAMS
135
JfranrtH iXauwr HilltamH
IBB2-13fir
Fig. 1. Francis Xavier Williams, 1882-1967. Photo taken circa 1960 by Dr.
George E. Lindsay, Director, California Academy of Sciences, San Francisco.
Elwood C. Zimmerman
Bishop Museum, Honolulu
Dr. Francis Xavier Williams (Fig. 1), a distinguished son of Cali-
fornia and an entomologist and naturalist of rare ability and notable ac-
complishment died, following a stroke, at Chula Vista, California on 16
The Pan-Pacific Entomologist 45: 135-146. April 1969
]36
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
December 1967 at age 85. Three nephews and a sister, Mother Louise of
the Convent of the Sacred Heart at El Cajon, California, survived him.
He was, with C. E. Pemberton, one of the two remaining members of the
c istinguished team of biologists who did so much pioneering and success-
ful work on the biological control of insects in Hawaii, and his research
£nd publications on the faunas of the Galapagos Islands, California,
Hawaii, the Philippines and other Pacific islands and on Hymenoptera
are widely known.
Dr. Williams was born in the small town of Martinez, near the north-
eastern end of San Francisco Bay, California, on 6 August 1882. His father
was Welsh, and he died at age 89. He was a farmer who became an ac-
countant, and he served with the Board of Supervisors and as secretary
to the Board of Health in San Francisco. His mother, Georgina Mel de
Fontenay, was bom on Staten Island, New York. She was educated in
T^rance, and she sailed around Cape Horn and married in Cailfornia. She
lied at age 94.
When Dr. Williams was about five years old, the family moved to San
Francisco. There he later entered St. Ignathius College where he went
hrough the lower grades and high school and obtained an AB degree in
1903 with classical training in Latin and Greek. The devout Catholic
family hoped that he would enter the priesthood, and he gave the pro-
posal serious consideration. After St. Ignathius College, Williams went
;o Stanford University where he studied under Prof. Vernon Lyman
Kellogg, the noted zoologist, and botanist Prof. LeRoy Abrams. At Stan-
ford he associated with fellow students David T. Fullaway and Cyril E.
Pemberton with both of whom he was later to be closely associated in
Hawaii. He graduated from Stanford in 1908, and Fullaway was in his
class. Fordyce Grinnell was also a student friend at Stanford.
Williams very early became associated with the California Academy of
Sciences. He was introduced to the Academy by Dr. Edwin C. Van Dyke
(later Professor of Entomology at the University of California, Berkeley)
who was a practicing physician and then a skilled amateur entomologist
with an extraordinary knowledge of the insects of California. Williams
used to visit Dr. Van Dyke in his office where they would discuss ento-
mology, and Dr. Van Dyke gave him much assistance and encourage-
ment as he did to many young entomologists of western North America,
including the biographer. Van Dyke’s enthusiasm and intense interest in
insects was contagious, and no serious student of entomology could avoid
gaining greatly from his association with Dr. Van Dkye.
Williams’ interest in natural history began at an early age, and in later
APRIL 1969] ZIMMERMAN FRANCIS XAVIER WILLIAMS
137
life he would recall with pleasure his collecting butterflies at Martinez
between the ages of four and five years. He began serious collecting and
study as a small boy, and his father gave him great encouragement. He
had insect boxes made for young Francis, and he had an attic room
modified as a laboratory where the budding scientist could house his col-
lections, study and carry on his experiments. The family lived within
eight blocks of Lone Mountain in San Francisco, where Francis collected
repeatedly and extensively. He also did much collecting at Land’s End,
and he collected aquatic insects in Mountain Lake (Marine Hospital
Lake) and in the stream and pools in the Presidio. He became interested
in aquatic insects as a school boy, and that interest stayed with him all
of his life. His extensive collections of San Francisco insects were made
at a time when there was much open space and much natural vegetation
in areas now covered by roads and buildings, and his valuable collec-
tions were given to the California Academy of Sciences. J. A. Powell
(Univ. California Pubs, in Entomology 32:5, 1964) has said “In 1908
and 1909, F. X. Williams made extensive collections [of Tortricidae] in
San Francisco, together wih a few others from central California locali-
ties. His material, which is the best sample of the now essentially extinct
Lepidoptera fauna of the San Francisco sand dunes, is at the California
Academy of Sciences.”
In 1905-1906, when a student at Stanford, Williams joined the Cali-
fornia Academy of Sciences’ Galapagos Island Expedition as entomolo-
gist. About a year was spent in the Archipelago, and all of the islands were
visited. Williams’ work on the insects of the Galapagos was outstanding,
and other workers have profited greatly from his extensive collections.
An account of the expedition was published by J. R. Slevin in 1931 in
his Log of the Schooner “^Academy’’ on a Voyage of Scientific Research
to the Galapagos Islands, 1905-1906 (Occas. Pap. Calif. Acad. Sci., 17:
1-162, pis. 1-16). Williams published The Butterflies and Hawk-Moths
of the Galapagos Islands in 1911 (Proc. Calif. Acad. Sci., (IV) 1:289-
322, pis. 20, 21), and in that worthy paper he gave an excellent discus-
sion of the Islands. In 1926, he published The Bees and Aculeate Wasps
of the Galapagos Islands (Proc. Calif. Acad. Sci., (IV) 2(2) : 347-
357). In 1907, Williams published a short report on the Expedition
(Entomol. News, 18: 260-261) in which he mentioned having collected
about 150 species of Coleoptera, but it was not until the publication of
Prof. Van Dyke’s posthumous The Coleoptera of the Galapagos Islands
(Occas. Pap. Calif. Acad. Sci. 22: 1-181, pis. 1-7) in 1953, nearly half
a century after their collection, that the Williams’ Galapagos Coleoptera
138
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
were reported upon. Van Dyke noted that the Williams’ collection of
C oleoptera was the largest ever made in the Galapagos.
After graduating from Stanford University in 1908, Williams worked
until 1910 as a plant inspector at the Ferry Building in San Francisco
under E. M. Ehrhorn, who later also went to Hawaii where he served as
a Hawaiian Government entomologist for many years. Williams then
went to the University of Kansas at Lawrence as an Assistant Curator of
tl e Snow Entomological Collection, and he there conducted with grad-
uate students three summer expeditions to western Kansas. The late
Prof. H. B. Hungerford, the noted heteropterist, was one of the graduate
students on the first field trip in 1910. Williams and Hungerford then
gathered details for their Notes on C oleoptera from Western Kansas (En-
tomol. News, 25; 1-7, pis. 1, 2, 1914) in which they discussed Ambly-
c.iila cylindriformis Say, a tiger beetle, and Hornia gigantea Wellman,
a meloid beetle predator of Anthophora bees. Williams obtained his MS
degree at Kansas, and his thesis was his classical work on the Larridae
of Kansas (Kansas Univ. Sci. Bull., 8(4) : 121-213, figs. 1—120, 1913).
1 1 that thesis work his characteristic method of lucid illustration was
already well developed. It was at Kansas that he took up his life-long
studies of Hymenoptera, and before that period he favored the Lepidop-
tera more than any other group of insects.
In 1910 (Entomol. News, 21(1): 30^1), Williams published The
hiitterflies of San Francisco, California. He listed 43 species of butter-
flies that were known at that time to inhabit or to have inhabited the
area, and he said that perhaps not more than 30 species could be found
there in 1910. He also said “Before a decade has passed away there will
b e little left of the insect fauna of our city, and therefore the importance
and necessity of making local insect lists while the fauna is still extant
and in an undiluted condition cannot be too much emphasized.” His
valuable comments on the vegetation and conditions prevailing in San
I rancisco in 1910 are in striking contrast to what one can now say of the
area.
In 1913, Williams left Kansas to accept an assistantship at Harvard
Ifniversity under Prof. William Morton Wheeler in the Bussey Institu-
tion, and he also served as an assistant in the beginning course in general
Zoology under G. H. Parker. Also while at Harvard, he had a part-time
j oh as a teacher of entomology in the Lothrop School of Landscape Ar-
cliitecture for women at Groton, Massachusetts. Williams received his
D.Sc. under Prof. Wheeler in 1915, and his thesis was the outstanding
and often quoted work on the embryology, development and habits of
APRIL 1969] ZIMMERMAN — FRANCIS XAVIER WILLIAMS
139
fireflies. This work, Photogenic Organs and Embryology of Lampyrids
(J. Morphol., 28(1) : 145-186, 1916) resulted from two years of study
on Photuris pennsylvanica (DeGeer) and Photinus consanguineus Le-
Conte. One of his close Harvard associates was W. W. Mann, the distin-
guished student of ants who later became the popular director of the Na-
tional Zoological Gardens at Washington, D.C. Williams’ association
with Prof. Wheeler meant much to him and greatly influenced his later
career. Prof. Wheeler conducted numerous field trips for his students
to the Blue Hills Reservation and other areas near Boston, and these
field trips were a particular joy to Williams. Prof. Wheeler considered
Williams to he one of his most promising students, and his faith was
not misplaced.
After receiving his doctorate, Williams found it difficult to find em-
ployment, but L. 0. Howard assisted him in obtaining a position, at a
very low salary, under A. F. Burgess in the United States Department
of Agriculture’s Gypsy Moth Laboratory in Massachusetts, in 1915.
Within a year, however, Frederick Muir, Entomologist at the Hawaiian
Sugar Planters’ Experiment Station in Honolulu (and the noted English
entomologist, David Sharp’s son-in-law), wrote to Prof. Wheeler asking
him to recommend a student who could be assigned to foreign travel to
search for parasites and predators that might be introduced to Hawaii to
assist in the control of insect pests of sugarcane. Prof. Wheeler recom-
mended Williams, and Williams was appointed Assistant Entomologist
and arrived in Honolulu in May, 1916, at age 34.
When Williams arrived in Hawaii, Muir and H. T. Osborn were search-
ing for parasites and predators in the Philippines, and 0. H. Swezey in-
troduced him to Hawaii and the insect problems of sugarcane and took
him into the plantations to demonstrate the damage being done by
Anomala orientalis beetles. Williams stayed in Honolulu only about 10
days, and then he sailed for the Philippines where he stayed from June
1916 to September 1917. In the Philippines he worked mostly with
Tiphia and Scolia wasp parasites of Anomala scarabs, and most of his
work was done in the Los Banos area. Williams sent Tiphia^ Dolichurus
and N otogonidia wasps to Honolulu, and they became established
and remain today familiar members of the introduced Hawaiian
fauna. It was during this work in the Philippines that he gathered
the data for his well known Philippine IV asp Studies (Experi-
ment Station, Hawaiian Sugar Planters’ Assoc., (Entomol. Ser.) Bull.,
14: 1-186, figs. 1-106, 1919). In that paper, S. A. Rohwer described
many of the new species discovered by Williams. Williams collected
140
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
riore than 180 species of aculeate wasps in the Philippines, and he in-
cluded accounts of the biologies of 52 of the species in the monograph.
Much of what he had to report was new and original, and it makes
the paper one of the more important contributions to the study of acule-
ate wasps. Howard Evans, the noted Harvard authority on wasps, told
r ie that Philippine Wasp Studies includes much that has never been re-
peated or surpassed. The section on Methocha, the tiphiids that attack
tiger beetle larvae, has been added to little (Williams had earlier pub-
1 shed a report on the north eastern United States species, in 1916, as a
lesult of studies he made while at Harvard). The section on mud-using
p ompilids is unrivalled, even today, as is the section on ampulicids. The
studies of Stenogaster were, until quite recently, the only information
available concerning the behavior of these wasps, and even now little
more is known than what Williams reported. Prof. Wheeler used William’s
information and illustrations of Stenogaster in his book The Social In-
sects ”
In 1917, Williams returned to Honolulu, and in early 1918 he was sent
to Australia to search for insect enemies of sugarcane leafhoppers and
aphids. He sent a number of parasites and predators back to Honolulu, and
a mongst them were the now familiar and widely spread brown lacewing,
Micromus navigatorum Brauer, and the ladybird beetle, Coccinela ar-
cuata Fabricius. He was in Queensland about six months, and he spent
1 inch of his time there studying wasps.
After returning to Honolulu from Queensland, he went to Pahala,
Hawaii, for about three months to work on the sugarcane leafhopper. In
1920, he went again to the Philippines where he searched unsuccess-
fully for wireworm parasites, and he studied fig-pollinating insects. He
discovered a new speices of larrid wasp (now Larra luzonensis Rohwer)
c Hacking the mole cricket, Gryllotalpa africana (Palisot de Beauvois),
ivhich was later introduced to Hawaii where it continues to be a valued
fid in the control of the destructive mole crickets. Williams visited about
10 Philippine islands and climbed many mountains to study altitudinal
faunal changes. On this trip he gathered material which he used for his
delightful and highly informative paper on The Natural History of A
Philippine Nipa House With Descriptions of New Wasps (Philippine J.
Sci., 35(1) : 53-118, pis. 1-8, 1928). This monograph reveals Williams’
ccute powers of observation and his very broad interests in natural his-
tory. Williams said of his nipa palm hut that “The building, to receive
the full benefit of such occupancy [by animals], must be an open nipa
l ouse . . ., preferably of some antiquity. No doubt the modernized
APRIL 1969] ZIMMERMAN — FRANCIS XAVIER WILLIAMS
141
screened-in structures with ceilings and tight wooden floors are more
durable and in some ways excel in comfort, but they exclude such inter-
esting household pets as the larger lizards, the banana frog, and the
several bats, as well as the greater bulk of winged insects that are at-
tracted to lights.” Williams told me that one night when he was quietly
writing notes by lamp light that he heard a slight noise, and, looking up,
he saw, just over his head, an enormous cobra searching for rats in the
the thatch of his hut.
Williams returned to Hawaii in 1922 and then went to the U.S. Na-
tional Museum to study the collections, and from there he went to
South America to search for wireworm parasites. He first went to
Ecuador, he crossed the Andes to the Amazon Basin, and he went to
Trinidad for about 10 days, to British Guiana for three or four months,
thence to Bardbados and to Rio de Janeiro, Sao Paulo, Pernambuco and
to Belem where he studied mole cricket and cockroach parasites. He re-
turned to Honolulu via New York and San Francisco in 1924 with co-
coons of various parasites, including larrid wasps and a Podium (a good
cockroach parasite) , but the parasites did not travel well during the long,
slow trip, and they were in poor condition when they arrived in Hono-
lulu and nothing became established from the attempted introductions.
Between all of his overseas assignments and other duties, Williams
found time to prepare his unsurpassed Notes on the Habits of the Bees
and Wasps of the Hawaiian Islands (Proc. Hawaiian Entomol. Soc., 6
(3) : 425-464, figs. 1-15, pis. 13-17, 1927) .
In 1928, Williams outstanding monograph Studies of Tropical Wasps
— Their Hosts and Associates {With Descriptions of New Species) ap-
peared (Experiment Station, Hawaiian Sugar Planters’ Assoc. (Entomol.
Ser.) Bull., 19: 1-179, figs. 1-14, pis. 1-33). This work (a collection of
eight high-quality essays on a variety of wasps ranging from fig wasps to
the Larridae of the Philippines to parasites of tiger beetles and other sub-
jects in various places from the Philippines to Queensland to South
America) is a mine of information and includes an abundance of highly
instructive illustrations. Howard Evans has told me that “Williams treat-
ment of the biology of Larra remains the standard reference (these wasps
are among the most primitive sphecids in nesting behavior). He also
treated the taxonomy of the South American Larra and all of the Philip-
pine Larrinae. The section on Podium remains very valuable. The paper
includes about all that is known about Notocyphus, a parasite of avicu-
larid spiders, all that is known of the nests of Priochilus (placed by
142
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
^J'^illiams in Pseudagenia) , and there is an extended, never repeated ac-
count of Aporus hirsutus at San Francisco.”
In 1928, Williams went to Webster Groves, Missouri, to study pred-
ators and parasites with A. F. Satterthwait. He introduced several spe-
cies, including Anaphoidea calendrae Gahan, to Hawaii, but they failed
to become established.
In 1934, Williams went to Guatemala in an attempt to discover pos-
s ble parasties and predators that could be used against Anomala scarabs.
I here he bred an Elis scoliid wasp which appeared to be a highly promis-
ing parasite. It was sent to Honolulu in numbers, but it failed to become
established. Williams also studied Pyrophorus elaterid beetles in the
1 ope that they might be used for wireworm control.
In 1939, Williams married Louisa Lewis Clark, of Honolulu. In July,
1940, they went to New Caledonia to study sugarcane insects, because
the establishment of the trans-Pacific air service posed a threat to the
Hawaiian sugar industry. In November, 1940, the Williams returned to
Honolulu with a valuable general collection of insects and living speci-
riens of Ampulex compressa (Fabricius), a large, beautiful, brilliantly
colored wasp which he found parasitizing large cockroaches in New Cal-
edonia. This fine parasite soon became established and spread widely in
Hawaii where it is now one of the most conspicuous insects of the low-
land fauna and is very common about houses and gardens in Honolulu.
His field work resulted in his paper The Aculeate Wasps of New Cale-
donia (Proc. Hawaiian Entomol. Soc., 12(2) : 407—452, figs. 1-8, 1945),
the only work of its kind on the New Caledonia fauna.
In October, 1947, at the request of the National Research Council and
the United States Navy, the Williams went to East Africa to study the
giant African snail, Achatina fulica, and its parasites and predators. The
pest snail was introduced by the Japanese to Micronesia where it was
found widely spread at the end of the second world war, and it became
established in Hawaii where it soon became a garden pest. Early in 1948,
the Williams returned to Honolulu.
Later in 1948, having retired from the Experiment Station, Hawaiian
Sugar Planters’ Association, the Williams moved to Danville, California,
near the base of Mt. Diablo and east of San Francisco Bay. Williams
vished to be near the California Academy of Sciences, to which he had
such a long and close attachment, and he wished to resume his studies of
California Hymenoptera. Danville is not far from Martinez, where
iV^illiams was born. The climate did not appear to agree with Mrs.
Williams, and they leased out their house for two years and bought a
APRIL 1969] ZIMMERMAN FRANCIS XAVIER WILLIAMS
143
house in Mill Valley, Marin County, and closer to San Francisco. This
location also proved unsuitable to Mrs. Williams, and they sold the Mill
Valley house and moved into a flat nearby. Williams then accepted an
an appointment from the Peruvian sugar growers to go to Peru for two
years to study methods of biological control of the serious sugarcane pest
Diatraea. However, Mrs. Williams had an eye operation and then devel-
oped arthritis, and Williams had to cancel his plans for the Peruvian
work. The Williams returned to the home in Danville, and then moved
to La Mesa, near San Diego, California, in search of a climate that ap-
pealed to Mrs. Williams. After several years at La Mesa, Mrs. Williams
health and eyesight continued to deteriorate, and they moved into a re-
tirement home in Chula Vista, also near San Diego. Tragically, Mrs.
Williams lost her eyesight and became bedridden with arthritis and was
confined to the hospital for a very long time. She died in September, 1965.
After the move to southern California, Williams associated closely with
the San Diego Museum of Natural History and continued his studies of
wasps, particularly his favorite larrids. By 1963, Williams had lost the
sight of one eye, but he continued collecting and his studies, and he
usually visited the Museum two days a week. His intense interest in nat-
ural history never faltered, and his observations and study continued
with keen devotion as long as he was physically able to work. His latter
California researches led to the publishing of The Wasps of the Genus
Solierella In California (Proc. Calif. Acad. Sci., (IV) 26(11) : 355-417,
pis. 11—21, figs. 1-3, 1950) ; Life History Studies of Pepsis And Hemi-
pepsis Wasps In California (Ann. Entomol. Soc. Amer., 49(5) : 447-466,
figs. 1-26, 1956) (this is a summary of his studies in 1937 and 1951-54,
and it is an excellent example of his work and his ability to convey im-
portant details accurately and lucidly to his readers) ; and The Wasps of
the Genus Plenoculus (Proc. Calif. Acad. Sci., (IV) 31(1): 1-49, 90
figs., 1960) .
I first met Williams upon my return from Bishop Museum’s Mangare-
van Expedition to southeastern Polynesia in the autumn of 1934. I found
him to be a storehouse of biological information, and I drew heavily
upon this source of knowledge during all of our years together in Hawaii.
I considered Williams to be the best informed, most broadly based and
his field work the most carefully done of any of his Hawaiian contem-
poraries. He was an extraordinarily keen observer and a most diligent
and persistent worker. He was a shy, very quiet, extremely modest and
retiring person. He had a gaunt appearance that may have resulted from
attacks of malaria. He was a lover of classical music, and he played the
144
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
flute. It was almost unknown of him to speak a harsh word or to reveal
any feeling of anger. He was a devout catholic, and in some ways, no
doubt because of his training and early consideration given to entering
the priesthood, he retained various qualities generally associated with
priestly ideals. Mrs. Williams was a protestant, and it appeared to be
with a great feeling of satisfaction to Williams that she was converted to
Catholicism just before her death.
Williams’s long years of bachelorhood and his being unburdened by
care of property or family responsibilities made possible his spending an
unusual amount of time in the pursuit of entomological investigations,
h e made many trips into the Hawaiian fields and mountains, particularly
during his long researches on the aquatic insects. Most weekends found
\/illiams in the field. On Sundays, before he was married, he attended
early Mass and then he frequently went alone into the mountains. He was
often also in the field on Saturdays. Scores of collecting trips during a
long period of years were made with Otto H. Swezey, his colleague at
the Experiment Station and very close friend and for whom Williams
4 ad the greatest admiration and respect. I accompanied Williams at var-
ious times, and I learned how careful and thorough were his methods of
study. I have considered his discovery of the highly unusual terrestrial
larva of the damselfly Megalagrion oahuense (Blackburn) to be the
crowning achievement of his Hawaiian field studies. Williams had been
collecting data for his classic series Biological Studies in Hawaiian Wa-
ter-Loving Insects (Proc. Hawaiian Entomol. Soc., 9(2) : 235-349, figs.
1-10, pis., 1-18, 1936; 10(1): 85-119, fig. 1, pis. 1-9, 1938; 11(3):
313-338, figs. 1-8, pis. 8-13, 1943; 12(1) : 186-196, figs. 8-16, 1944),
and he had studied the life histories of all of the Odonata on the island
cf Oahu with the exception of this fine, large, endemic damselfly. Al-
though the adults could be found without great difficulty, Williams was
i nable to discover larvae in any of the known or suspected odonate habi-
tats. Over and over again he searched for larvae in the areas where the
z dults flew, but all his attempts were failures. He noted that the adults
\/ere often to be seen on steep mountainsides far from water, and week
cfter week he studied their movements. He found that the females were
sometimes seen on dense thickets of the pestiferous staghorn fern, Gleiche-
ida linearis, which forms tangled masses in openings in the lower and
raiddle forests. By persistent observation he found that some females de-
scended into the masses of ferns. Armed with this information, he began
£. search of the damp trash and leaf mold beneath the fern thickets. With
extraordinary patience and after repeated and laborious searches that
APRIL 1969] ZIMMERMAN FRANCIS XAVIER WILLIAMS
145
would have defeated a lesser man, Williams finally succeeded in discov-
ering that the nymphs of this damselfly are highly unusual because they
are terrestrial and are capable of living far from water. The nymphs
are very hairy, and their gills are shortened, thickened and hairy and
are adapted for a life in the damp ground litter beneath dense masses of
vegetation. Most of his other studies on the Hawaiian water-loving in-
sects are the first and only of their kind in Hawaii. His studies are of
great value and have added much to our knowledge of the Hawaiian
fauna.
In spite of all of his travels, other duties and many other research proj-
ects, Williams wrote the excellent and basic Handbook of the Insects and
Other Invertebrates of Hawaiian Sugar Cane Fields (Hawaiian Sugar
Planters’ Association Experiment Station, pp. 1-400, figs. 1-190, pis. 1-
41, 1931) . This book has been the standard text on the common insects
of Hawaii since its publication, and it is an invaluable source of informa-
tion on the Hawaiian fauna.
After their marriage, the Williams built a small cottage near the
Hawaiian Sugar Planters’ Experiment Station in Honolulu and sur-
rounded it with flowers, shrubs and trees. Here they fed and attracted
to their garden many birds, and they planted various flowers (and some
weeds ! ) to attract insects. They raised an orphan mynah bird as a pet,
and this led to a detailed study of the mynah {Acridotheres tristis) in
Honolulu. They printed privately their entertaining and informative book
Mike The Mynah in 1946. In their garden they built a small pool to at-
tract birds and invertebrates. The introduced tropical American toad,
Bufo marinus would frequently swarm into the pool, and Williams would
periodically carry away for release elsewhere numbers of the toads. He
noted that one of the toads he had carried to the grounds of the Experi-
ment Station (about a half mile away and across several roads and a
busy highway) had a deformed foot. Much to his astonishment, he found
the toad again in his pool, and he thus determined that the toads have a
homing instinct and although carried a considerable distance away,
some of them found their ways back through many hazards to his pool.
Williams name is fixed firmly in the annals of hymenopterology, and
many of his studies include original and unique discoveries and observa-
tions. Howard Evans told me that Williams often sent him wasp larvae,
which Evans described in his outstanding series on larvae, and Williams
was always “exceedingly generous” in sending him specimens for study
and took special care to write to him to tell him how much he enjoyed
reading his papers. This was characteristic of Williams. Evans also told
146
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
me that “Williams paper on Epyris extraneous is one of the few detailed
s udies of a bethylid wasp and the only one for that genus. He described
t VO unusual genera of pompilids from Fiji, N esopompilus and Dendro-
pompilus, and, as usual, his descriptions and abundant illustrations make
it unnecessary to see the types to be able to place the genera. He dis-
CDvered a striking new genus of sphecids {Xenosphex) in southern Cali-
fornia which still cannot be placed satisfactorily in the classification of
t ie family. His last paper is an excellent report on an otherwise un-
studied spider wasp that nested in milkweed stems in the garden of his
retirement home at Chula Vista.”
Williams developed a highly distinctive artistic style early in his ca-
1 eer, and his hundreds of published illustrations are mostly recognizable
cS his at a glance. They are clear, crisp, highly instructive and as per-
sonal as his signature.
Williams was such an excellent, broadly based researcher that it is
perhaps regretable that his employment was not in a museum or a uni-
versity instead of a commerical agricultural experiment station. Although
he was employed as an economic entomologist assigned to devote his
labors to the protection of the sugarcane crop, he was able to make more
original observations, make more high quality illustrations and publish
more excellent monographs on ethology, ecology and taxonomy than
many persons accomplish when employed to concentrate on such sub-
jects. All who have to work with his many publications have reason to
thank that exceedingly modest, sensitive, highly competent, devoted nat-
uralist for what he has left to posterity. A bibliography of his published
vritings and a list of the new taxa he described has been prepared by Dr.
Paul H. Arnaud, Jr. and will appear in the Occasional Papers of the
California Academy of Sciences for 1969.
APRIL 1969]
PHILIP NEW NEOTROPICAL TABANIDAE
147
New or Little-Known Neotropical Tabanidae
(Diptera)
Cornelius B. Philip
National Institutes of Health, Pub. Health Serv.
U.S. Dept. H.E.W., Hamilton, Montana
Descriptions and comments on Neotropical Tabanidae are provided
below.
Esenbeckia (Ricardoa) potrix Philip, new species
(Fig. 1)
Robust, blackish species with beard, chest and narrow abdominal incisures, all
whitish-haired, Irons with tall, narrow blackish keel, antennae and palpi red, latter
short and pointed, about one-fourth as long as proboscis, proboscis subequal to
height of head ; legs dark brown to black with black hairs.
Female. — (Holotype, 19 mm). Frons yellow pollinose, slightly widened above
and below, keel narrow with small irregular expansion near base but widely sep-
arated from the upper margin of the yellow pollinose subcallus; 3 ocelli at vertex.
Face and cheeks buff-gray pollinose with red quadrate bare area above mouth
parts. Shapes of antennae and palpi as figured, scapes and pedicels mostly yellow-
haired changing to black distally; basal annuli of flagellum somewhat fused; apical
palpal segments mostly black-haired intermixed with yellow hairs dorsally and
around ovoid, deep lateral grooves. Labella almost completely sclerotized.
Notum and scutellum dull black with narrow brown margins around former,
and covered with short black and yellow hairs. Pleura sooty-brown, with
mostly pale yellow pile which extends onto blackish coxae. Femora and hind tibiae
black with concolorous hairs ; fore and midtibiae dark brown with short black hairs.
Wings suffused with yellow, intensified in costal cells. Halteres reddish.
Abdomen subshiny black with indefinite red-brown shades widely on sides of
tergite 2, incisures above and below with narrow pale-haired bands which could
disappear with wear.
Holotype Female, Sinaloa, Mexico, 37 miles east of Concordia, 2
September 1957, R. E. Widdows. In collection of L. L Pechuman.
This species would run to couplet 15 in my key to North American
species (1954) . There E. incisuralis var. tinkhami Phil, has considerable
resemblance, but, in addition to being smaller, the palpi are longer, the
knees and tibiae are reddish and all appendages predominantly yellow-
haired.
Esenbeckia filipalpis divergens Philip, new subspecies
(Fig. 2)
Though Philip (1954) included both sexes of E. filipalpis (Will.) in
keys to North American species, and Hine (1925) included reference to
a specimen from Costa Rica, Fairchild does not include it in several
The Pan-Pacific Entomologist 45: 147-152. April 1969
148
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Fig. 1. Frons, antenna and palp of Esenbeckia {Ricardoa) potrix Philip, female.
Fig. 2. Antenna and palp of Esenbeckia filipalpis divergens Philip, male. Fig. 3.
Frons of Catachlorops (Psalidia) medemi Philip, female, a. wing; h. antenna and
palp. Fig. 4. Frons, antenna and palp of Stypommisa (Styphocera) antennina,
Philip, female.
publications dealing with other species in the better known fauna of
Panama. Until confirmatory specimens are studied, occurrence of the
typical, more southern form in Central America and Mexico remains
doubtful.
Further collecting of both sexes is also needed to determine whether
2 males from Nayarit, Mexico, received from L. L. Pechuman, which
pear close resemblance, are distinct specifically; for the present, they
are described below as subspecies.
Medium-sized fly with brown notum; bicolored legs with bright yellow tibiae;
palpi slender, spatulate and blunt apically and half or more as long as proboscis;
abdomen yellow on basal 3 segments with narrow middorsal black triangles, re-
inaining segments black.
Male. — (Holotype, 15 mm). Head but little wider than thorax, eyes bare with
uniformly small facets; 3 prominent ocelli at occipital notch. Face and cheeks buff-
gray with yellow hairs. Antennae and palpi red with black hairs, shaped as figured,
palpi bare laterally for about two-thirds their lengths.
Notum and scutellum brownish-red with sparse short yellow and black hairs;
pleura dark red, mostly dark brown-haired. Coxae and femora dark brown, black-
haired; tibiae contrasting bright yellow with short concolorous hairs. Wings with
customary venation, except cells M^^ pointed at base, cross-vein at apex of discal cell
thus eliminated; spur veins present and cells Rg closed; suffused with brown, inten-
sified costally and hasally. Halteres red.
APRIL 1969]
PHILIP NEW NEOTROPICAL TABANIDAE
149
First tergite and first 2 sternites yellow, predominantly yellow-haired inverted
dark triangle with black hairs on first not as wide as scutellum. Second and third
tergites yellow, mostly black-haired with tall median inverted black triangles, not
quite crossing tergites, black spot on each outer upper corner. Sternite 3 yellow,
mostly black-haired. Remaining segments black with black hairs and small tufts
of yellow hairs on outer corners of tergites.
Holotype Male, San Blas, Nayarit, Mexico, 2 July 1956, R. & K.
Driesbach. In the collection of L. L. Pechuman.
Paratype male, same data, in close agreement; in collection of the
author. The median spot on tergite 3 is not narrowed behind and the
outer dark spots on tergite 2 are evanescent.
The resemblance to typical E. filipalpis is striking, but the brown not
blackish notum, broader blunt palpi, and bicolored legs of this subspecies
are distinctive and may ultimately justify specific differentiation. A
typical male from Formosa, northern Argentina, is available for com-
parison.
Cataclilorops (Psalidia) medemi Philip, new species
(Fig. 3)
Rather large, predominantly brown fly with reddish-brown appendages, pictured
Avings with lighter areas more reduced than usual in this group, and antennal spine
reaching beyond base of style.
Female. — (Holotype, 19 mm). Eyes bare, no bands revived. Front tall and nar-
row, parallel-sided, buff-gray pollinose with strong median, red keel reaching almost
to vertex and not touching eyes; index 1:5.8. No ocelli. Subcallus small, buff-pol-
linose laterally, and sharply bare in middle. Face and cheeks buff -yellow pollinose,
brown pilose below. Antennae shaped as figured, red, annuli dark brown, black
hairs on 2 basal segments and tip of spine. Palpi long and slender, blunt, red with
black hairs on apical segment, and reaching base of sclerotized black labella.
Thorax reddish-brown with sparse, short, mixed black and yellow hairs dorsally,
light brown pile on pleura. Forelegs brown with black hairs, basal half of foretibiae
pale yellow. Two hind pairs red with concolorous hairs, darkening on tibiae with
black hairs. Wing mottled brown, more yellow hasocostally, irregular paler but not
clear, fenestrae across cross-veins in middle and centers of cells Rg, Rg and M^;
apex beyond tip of vein R 4 + 5 , and anal area similarly suffused, pale. Cell Rg nar-
rowed at margin to about half its maximum Avidth; no spur veins. Halteres yellow.
Basicostas bare.
Abdomen uniformly reddish-brown, predominantly black-haired above, yellow on
venter.
Holotype Female, La Pedrera (Amazonas) Lower Rio Caqueta,
Colombia, 29 September 1968, F. Medem. “In a hut at night.” In col-
lection of the author through courtesy of the collector, a student of cai-
mans from which he has taken T. fervens Linn, and T . modestus Wied.
while they were biting these reptiles.
150
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Paratype females (on loan from G. B. Fairchild) ; 16-19 mm. 3, Rio
de Janeiro, 12 October 1917, E. M. Miller. In collections of Stuttgart
J 'Natural History Museum and of G. B. Fairchild, who had also decided
lheir distinctness. 1, Brazil, in Lutz Coll.; label indistinct but inter-
preted by G. B. F. as “Estrada Ferro Madeira Mamore” (between Porto
Yelho and Guajara Mirim, Rondonia) . In good agreement with the holo-
type. The vestiture of body and appendages may be entirely orange-
laired, and the bare area in middle of the subcallus may be reduced
and shield-shaped but is not separated from the frontal carina in any.
Dr. Fairchild has suggested this species may not have been noticed
joefore because of nocturnal (or crepuscular) habits, indicated by data
on the holotype.
The closest relative appears to be C. (P.) scurra (Fchld.) from Pan-
ama which, however, has the basal cells entirely dark to the apices with
sharper substigmal, subapical and apical fenestrae, as well as closed
cells Rf,, and the bare median area of the subcallus is separated from the
median keel by an upper pollinose margin.
Tabanus (Neotabanus) xerodes Philip. — ^This was described from a
single worn female from Arica (Philip, 1967) . A better-preserved speci-
men was loaned me by Dr. Sixto Coscaron, collected at Azapa, Cata-
raarka Prov., by Etcheverry (no date) which permits augmenting the
description.
The callosity is cherry red, more quadrate than in the type and with
the median extension obscured by sparse pollen. Palpi a little less
swollen than figured. Notum with 3 gray notal lines prominent, vesti-
ture with brassy appressed hairs among gray pile. No spur veins. Hind-
tibial fringes whitish, darker hairs only distally. Median abdominal
line with gray pollinosity ; tergite 1 predominantly black, sides of tergites
2 and 3 widely red with diagonal dashes of yellow hairs, the trivittate
pattern evident from tergite 4 caudad. The Neotabanus eye pattern was
confirmed.
Tabanus monotaxis Philip. — This likewise was based on a unique
female with broken antennae from Antofagasta Prov. A well-preserved
specimen, also from Antofagasta Prov., Calama, 7 December 1959, Inst.
Pedagogico, was also received from Dr. Coscaron, which permits figur-
ing of the complete antennae and some additional comments. Length
15 mm. Single eye-band confirmed, and lower border darkened. Neither
the figure nor this specimen confirm the original statement of front “. . .
strongly convergent below . . .” though there is mild convergence. Cal-
losity a little narrower, more conical. Palpi more drawn out into an
APRIL 1969]
PHILIP — NEW NEOTROPICAL TABANIDAE
151
attenuated point than figured and entirely pale-haired. Abdominal pub-
escence pale yellow; sternite 7 not inconspicuous and noticeably, but
not excessively narrower than 6.
Stypommisa Enderlein
Styphocera Philip, new subgenus
Antennae shorter than height of fronts, with triannulate styles (named in refer-
ence to the contracted flagellum), fronts wider than typical Stypommisa, about
twice taller than wide, small but distinct ocellar tubercle, eyes sparsely pilose, un-
handed, wings tinted with isolated clouds and spur veins at bases of veins R^, basi-
costas with few setae and abdomen without patterns other than pale incisures.
The triannulate antennal styles and relatively broad fronts set this sub-
genus apart among other Stypommisa species, analogous to separation
of subgenera Brachytahanus Fchld. and novum A Fchld. in Steno-
hanus which otherwise have clear wings without spur veins and com-
pletely bare basicostas and eyes. Stypommisa subgenus novum B
Fchld. also has broad fronts and basicostas often with some setae, but
the eyes are bare, wings clear without spur veins, styles quadriannulate,
and abdomens with median stripes.
Type Species. — Stypommisa [Styphocera] antemiina Philip.
Stypommisa (Styphocera) antennina Philip, new species
(Fig. 4)
Small, dark brownish-black species with narrowly pale-banded abdomen, suffused
wings with intensified clouds on cross-veins, short spur veins, basicostas with few
setae, sparsely hairy eyes, and short all-red antennae.
Female. — (Holotype, 8 mm; wings 8.5 mm) . Head small, hardly as wide as notum
between wings; eyes with short brown sparse hairs; no bands revived. Frons buff-
gray to brown pollinose mesally; slightly widened below, index 1:2.2; indistinct
tubercle near vertex; callosity black triangular, not touching eyes and attenuated
about midway to vertex. Subcallus dull brown pollinose. Face dull sooty-gray to
brown pollinose, the apodemal pits unusually large and sunken but pollinose;
cheeks mostly brown-haired. Antennae bright orange-reddish, small apical annulus
black; scapes sub cylindrical, nearly twice longer than thick and subequal to plates
in length, covered with shaggy black hairs; pedicels somewhat bead-like with low
dorsoapical points, also coarsely black-haired; flagellums robust, not excised dor-
sally, style chunky and indistinctly 3-annulate. Palpi brown, shaggy-black-haired,
falcate and tapered to blunt point nearly to tip of short fleshy black proboscis.
Thorax blackish, unpatterned (apparently discolored), pronotol lobes yellow,
prealar ones and scutellum black; vestiture sparse yellow and black-haired. Legs
brownish-black, the tibiae more dull reddish; mostly black-haired. Wings with
normal venation, suffusion intensified anteriorly, costal cells and isolated clouds
brown. Basicostas pointed with few black setae apically. Halteres brown.
Abdomen above and below dark brown to black with narrow yellow incisures,
narrowest caudally; sternite 2 with yellow crossing it sublaterally ; tergite 1 divided
152
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
nesally by deep sulcus which forks peculiarly on anterior third; scattered yellow
and black hairs ; tergite 7 broad and extruded as long as 6.
Holotype Female, Huanuco, Peru. 17 September 1954. 7,500 ft. In
c Dllection of L. L. Pechuman.
The eyes are more plainly hairy than most species now assigned to
Stypommisa, while the cheeks, palpi and antennae basally are much more
shaggy-haired. The shapes of the antennae diverge from those known in
ether species in the more compact red triannulate flagellums with dorsal
Hunt angles near the middle of the plates, and styles short and robust.
Change of name. — As called to my attention by both Drs. Alan Stone
end G. B. Fairchild, my recently described T. attenuatus from Honduras
is preoccupied by Walker (1848, List 1: 159). Therefore T. abattenuis
new name is proposed for the former.
Literature Cited
Hine, J. S. 1925. Tabanidae of Mexico, Central America and the West Indies. Univ.
Mich., Decas. Paper Mus. Zool. No. 162, 35 pp.
Philip, C. B. 1954. New North American Tabanidae. VIII. Notes on and keys to
the genera and species of P angoniinae exclusive of Chrysops. Rev. Brasil.
Entomologia, 2: 13-60.
1967. Species of Tabanus related to T. xanthogaster Philippi in Chile. J. Med.
Entomol., 4: 463.
Two new species of Sepedon separated
from S. armipes Loew in Western North America
(Diptera: Sciomyzidae)
T. W. Fisher and R. E. Orth^
University of California, Riverside
Sepedon armipes Loew, a nearctic species, is widely distributed in the
Jnited States and Canada (Steyskal 1951; Foote 1951; Neff and Berg
1966). Neff and Berg, however, expressed caution regarding the total
acceptance of this concept in a footnote to a distribution map (p. 112)
for the species which states: “The recent discovery of a sibling species
in Utah and southern Idaho casts some doubt on armipes records in that
area.” Although our records of species of Sepedon collected in California
( — a result of a survey in progress^ of the marsh flies of California to
I Specialist and Laboratory Technician, respectively.
“Univ. Calif. Agric. Exp. Sta. Project No. 2037.
The Pan-Pacific Entomologist 45: 152-164. April 1969
APRIL 1969]
FISHER & ORTH — TWO NEW SEPEDON
153
be reported in a forthcoming issue of the California Insect Survey Bul-
letin — ) had been forwarded to Neff and Berg for inclusion in their
paper, we were suspicious that our California “armipe^,” too, was dis-
tinct from eastern United States representatives of the species (Fisher
1966). Subsequent correspondence and exchange of material and draw-
ings with L. V. Knutson confirmed the existence of three forms of
“armipes,” the nominal typical species and the two distinct species de-
scribed herein.
The terminalia and/or hind femur and tibia of S. armipes have been
figured by Cresson (1920, fig. 39), Steyskal (1951, fig. 1; 1956, fig.
17), and Foote (1961, fig. 16). These workers and Melander (1920)
also provided keys to the species. Loew (1859) described the species
from a male taken by Osten Sacken at Washington, D. C. and it is as-
sumed the specimens used by Steyskal and Cresson for illustration were
from eastern United States and are conspecific with Loew’s type. In
order to standardize our basis of comparison with the two new species,
S. armipes is refigured in this paper in the hope that characters hereto-
fore not, or minimally, used will facilitate future comparative studies in
the Armipes group.
Characteristics of the Genus Sepedon. — Propleuron without strong-
bristle; first segment of foretarsus not of a color strongly contrasting
with remaining segments. Vallar bristles absent. Ocellar and humeral
bristles absent. Post ocellar bristles well-developed. Frons with one pair
of fronto-orbital bristles. One pair of scutellar bristles. Posterior cross-
vein arcuate, or slightly so. The three species of concern here run to 5.
armipes in existing keys.
Characteristics of the Armipes Group. — Supraspiracular convex-
ity with black hairs. Medifacies with scattered black hairs. Second an-
tennal segment slender, nearly round in cross section. Body less than
six mm. long. Approximately midway on ventral surface of hind femur
of the male is a deep indentation (notch) whose basal margin is de-
marcated proximally by a heavy process (basimedian prong) (Steyskal
1951) which may be clearly bifid, as in S. bifida Steyskal, or nearly
simple as in S. melaaderi Steyskal. A single simple process (basal
prong) is likewise located on the ventral surface of the hind femur and
is situated between the basimedian prong and the base of the femur. The
basal prong is lacking in S. haplobasis Steyskal. Hind femur of female
simple. Hind tibia of both sexes distinctly curved forward in distal
third. A freshly pinned specimen of S. capellei (Fig. 1) illustrates well
the hind leg characteristic of the Armipes group.
154
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Fig. 1. Sepedon capellei Fisher & Orth { $ ) Calif., Mono Co., NW. comer Mono
Lake, 11 July 1968, (T. W. Fisher and R. E. Orth), AS-715. Dextral appendages
removed.
Species of Sepedon placed in the Armipes group are S. anchista Stey-
skal, S. armipes, S. bifida, S. capellei, S. haplobasis Steyskal, S. meland-
tri, and 5. pseudarmipes. Some of the males can he separated by
differences in the hind femur. All can be separated by morphological
differences in the genitalia.
Figs. 2-7. Ventral and sinistral views of male terminalia: Figs. 2-3. Sepedon
crmipes Loew, New York, Tompkins Co., 29 June 1958 (no coll.). Figs. 4-5. Sepe-
don pseudarmipes Fisher & Orth, paratopotype, Washington, Grant Co., O’Sullivan
Dam, 18 July 1954 (H. G. Davis). Figs. 6-7. Sepedon capellei Fisher & Orth, para-
topotype California, Inyo Co., 2.5 mi. W. Bishop, 1 August 1965 (T. W. Fisher and R.
E. Orth) AS-344. aed, aedeagus; ap, anterior process of surstylus; ce, cercus; ep,
epandrium; s, surstylus; vp, ventral process of hypandrium.
APRIL 1969] FISHER & ORTH TWO NEW SEPEDON
155
156
THE PAN-PACIFIC ENTOMOLOGIST
[voL. 45,
NO
2
d
/
10
11
APRIL 1969] FISHER & ORTH — TWO NEW SEPEDON
157
All are considered to be nearctic species, with the possible exception
of S. haplobasis which Steyskal (1960) described from material taken 15
km. east of Mexico City, Mexico, a locality which we estimate to be ap-
proximately 7,500 feet in elevation.
Morphological Differences Between S. armipes, S. capellei, and
S. PSEUDARMIPES. — Size differences between these three species are vir-
tually nil. The average wing length of 20 males of S. armipes from Utah
and Idaho was 4.37 mm. Male paratypes of S. capellei and S. pseudar-
mipes had average wing lengths of 4.43 and 4.34 mm, respectively. Stey-
skal (correspondence) earlier had suggested to Knutson that the distance
from the base of the hind femur to the distal portion of the notch relative
to the total length of the femur might be useful in the separation of S.
armipes from S. capellei. However, at that time, S. pseudarmipes was un-
known. On the other hand differences in the terminalia (Figs. 2-7) , with
particular emphasis on the aedeagi (Figs. 8-11), provide a high level of
confidence for postive placement of the three species. By these criteria
all but 4 of 1,357 males were placed without hesitancy.
Characteristics of the aedeagus used to distinguish the three species
are (1) angle a b d and (2) the distance between points e (gonopore)
and / (basal anterior process) relative to the distance between points b
and d. In S. armipes angle ab d approximates 85 °;c/< bd; and point /
is short, blunt, and directed ventrally. Point c varies in its position be-
tween b and d in different specimens in all three species, and is of no
diagnostic value.
We found no reliable morphological differences to distinguish females
of the three species.
Sepedon capellei Fisher and Orth, new species
Holotype male. — Postabdomen as in Figs. 6, 7. Aedeagus as in Fig. 11; angle
ah d approximately 137°; ej'^hd; point massively dentate, directed anteroven-
trally. Wing length 4.4 mm. External morphology as in S. armipes, except uniformly
darker color.
Allotype female. — Terminalia as in Fig. 13. Wing length 5.2 mm. Coloration
same as male.
<-
Figs. 8-11. Aedeagi, sinistral views: Fig. 8. Sepedon armipes Loew, Utah, Cache
Co. (no locality), 13 September 1964 (K. Capelle). Fig. 9. S. pseudarmipes Fisher
& Orth, holotype, Washington, Grant Co., O’Sullivan Dam, 11 October 1953, (M. T.
James). Fig. 10. S. capellei Fisher & Orth, Utah, Cache Co. (no locality), 13 Sep-
tember 1964, (K. Capelle). Fig. 11. S. capellei Fisher & Orth, holotype, Calif.,
Inyo Co., 2.5 mi. W. Bishop, 1 August 1965, (T. W. Fisher and R. E. Orth), AS-
344. (See text for significance of lettered points of reference.)
158
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Holotype male, 2.5 miles west of Bishop, Inyo County, California,
Line Street, Elevation, 4,250 feet; Latitude, 37° 20' North; Longi-
ti de, 118° 22' West approximately; 1 August 1965; (T. W. Fisher and
R. E. Orth) ; field notes Accession No. AS-344. Allotype female same
data. The type locality is a roadside ditch with shallow, flowing clear
water, and bordered by low sedges and grasses.
Paratypes. — 35, all from California. Wing length 22 $ , 4.1^.7 mm, average
4.43. Wing length 13 2, 4. 7-5.1 mm, average 4.96. Inyo Co.: 1 , 12 mi. W. Big-
pine, Glacier lodge, 7,800'; 12 August 1966 (T. W. Fisher), MA-118; 2 same
locality, (T. W. Fisher and R. E. Orth) (hereafter stated F & 0), AS-640; 2 2
(Paratopotypes) , 2.5 mi. W. Bishop, 4,250', 1 August 1965, AS-344, (F & 0).
]V. ODOC Co.: 2 ^,2 2,1 mi. S. Alturas, 4,300', 21 September 1966, (F & 0), AS-
533. Mono Co.: 3 $,2 2, Hot Cr., 2 mi. E. Hatchery, 6,940', 20 September 1966,
( " & 0), AS-522; 2 ^,1 2, NW. corner Mono Lake, 6,440', 2 August 1965, (F &
0), AS-347 ; 2 i5, 1 2, same locality, 20 September 1966, (F & 0), AS-523; 4 $ ,
12,2 mi. W. Bridgeport, 6,500', 6 June 1966, (F & 0), AS-437. Plumas Co.: 2 $ ,
1 2, Rock Creek, Hwy 36, 4,900', 4 August 1965, (F & 0), AS-359. Shasta Co.:
1 2,3 mi. E. Anderson, 430', 12 June 1965 (T. W. Fisher), AS-313; 2 $ , Red-
ding, 560', 12 June 1965, (T. W. Fisher), AS-312; Sierra Co.: 2 ^,3.5 mi. NW.
Sierraville, 4,935', 22 September 1966, (F & 0), AS-538; 2 2 , 1.6 mi. W. Sierra-
ville, 4,940', 23 August 1967, (F & 0), AS-627.
Deposition of Type Material. — Holotype and Allotype, California Academy of
Sciences, CAS 10166 — also, 6 paratypes; 8 paratypes, USNM; 4 paratypes, Cornell
Lniversity; 16 paratypes. Museum of Department of Biological Control, University
0 California, Riverside.
Material was also examined from the following localities and in the number in-
dicated in parenthesis: — California (1464 total). Alpine Co. (79) : Heenan Lake,
1 mi. E. Luther Pass, Markleeville, Sorensen’s, 2 mi. S. Woodfords, Silver
Creek. Inyo Co. (34) : Big Pine, 2.5 mi. W. Bishop, 1.7 mi. N. Cartago, Buclchorn
Spring (Deep Springs Lake), 12 mi. W. Big Pine. Lassen Co. (32): 2 mi. W.
h allelujah Jet., Smith Flat Reservoir, 16 mi. N. Susanville, 6 mi. E. Bieber, 12 mi.
S Ravendale. Modoc Co. (107) : Alturas, Cedar Pass Camp Gnd, S. Cedarville, 3
n i. N. Eagleville, 8 mi. SW. Eagleville, 1.5 mi. E. Buck Creek Ranger Sta., Lake
City, Lily Lake Rd., Likely, So. Fork Pit River, Willow Ranch. Mono Co. (534) :
Bridgeport, 2.5 mi. S. Coleville, Fish Slough, Hot Creek, 3 mi. NW. Topaz, Topaz,
^ W. corner Mono Lake, E. Sonora Pass. Nevada Co. (20) : 1 mi. SW. Boca Spr.,
h obart Mills, Sagehen. Placer Co. (1): 3.5 mi. SE. Truckee (Martis Cr.).
Plumas Co. (45) : Chester, Crescent Mills, 3 mi. S. Graeagle, Meadow Valley,
(Quincy, Rock Cr. (Hwy 36). Shasta Co. (10) : Anderson, 5 mi. SW. Redding, Red-
ding, 2 mi. S. Pondosa, Cassel. Sierra Co. (154) : Sierraville. Siskiyou Co. (20) :
Pray (Butte Cr.), 20 mi. SW. Gazelle, Grass Lake, Grenada (Shasta Riv.), Meiss
lake, Orr Lake. Tehama Co. (1) : 4 mi. SE. Vina. Tuolumne Co. (5) : E. Doug-
Ifss. Plus 77 from inexact localities.
IDAHO (23). Blaine Co.: Picabo (Silver Cr.), Gannett. Canyon Co.: Succor
Cr. (?). Caribou Co.: Bancroft. Cassia Co.: Elba Basin Pass. Elmore Co.: Lake
Camas Res., Dixie. Franklin Co.: Cub River Canyon. Gooding Co.: 2 mi. S.
P agerman. Hailey Co.: 16 mi. S. Hailey (Wood River). Lake Co.: Fish Haven.
APRIL 1969] FISHER & ORTH TWO NEW SEPEDON
159
12
0.3mm
Figs. 12-13. Ventral views of female terminalia: Fig. 12. Sepedon armipes Loew,
New York, Tompkins Co., McLean, 24 June 1958, (no coll.). Fig. 13. S. capellei
Fisher & Orth, allotype, Calif., Inyo Co., 2.5 mi. W. Bishop, 1 August 1965, (T. W.
Fisher and R. E. Orth), AS-344. The characteristics of tergites VIII and X vary
significantly between- and within species and geographic areas.
NEVADA (17). Douglass Co.: 4 mi. E. Stateline. Elko Co.: Deeth, 10.7 mi. S.
Elko. Ormsby Co.: Carson City. Washoe Co.: Wadsworth.
OREGON (18). Crook Co.: Ochoco Cr. Deschutes Co.: Bend, 20 mi. S. Bend.
Grant Co.: 8-10 mi. N. Seneca. Hood River Co.: Hood River. Josephine Co.:
1/2 mi. S. Cave Jet. Klamath Co.: Fort Klamath (Crooked Creek), Harpold Dam,
Klamath Marsh, 4 mi. E. Sprague River. Lake Co.: 9 mi. S. Lakeview. Malheur
Co.: Malheur River Canyon. Union Co.: Anthony Lake, 10 mi. N. Powder (?),
North Powder. Wallowa Co.: Enterprise, Minam. Wheeler Co.: 5 mi. SE. Rich-
mond.
UTAH (83). Box Elder Co.: Brigham Canyon. Cache Co.: Blacksmith Fork and
Curtis Creek, Logan, Logan Canyon, Providence. Emery Co.: Moab. Iron Co.:
Cedar City. Millard Co.: Delta. Rich Co.: Garden City. Sevier Co.: Salina. Utah
Co.: Spanish Fork. Washington Co.: Leeds. Wayne Co.: Lyman Milleville (no
county determined) .
The patronym is after Mr. Kenneth J. Capelle, Wildlife Research Biol-
ogist, Bear River Research Station, Brigham City, Utah. He provided ex-
cellent series of specimens which materially broadened this study.
Sepedon pseudarmipes Fisher and Orth, new species
Holotype male. — Postabdomen as in Figs. 4, and 5. Aedeagus as in Fig. 9;
angle a b d approximately 100°; e j — h d; point sharply pointed, directed
anteriorly. Wing length 4.6 mm. External morphology and color as in S. armipes.
Allotype female. — Not designated.
150
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Holotype male, O’SuLLivAN, Dam, Station #2, Grant County Wash-
inton, 11 October 1953; (M. T. James) ; (elev. ca. 1,100'; Latitude 47°
^ ., Longitude 119° W., approximately).
Paratypes. — 28. Wing length 27 $, 4.1-4.7 mm, average 4.34. One $, 4.9 mm.
IDAHO. Bonner Co.: 2 ^,8 mi. N. Sandpoint, 8 July 1959, (B. A. Foote).
OREGON. Umatilla Co.: 9 $, Cold Springs, 11 April 1954, (M. T. James and J.
J Davis). WASHINGTON. Grant Co.: O’Sullivan Dam, — 2 $, 11 October 1953,
m. T. James) ; 1 2 April 1954, (MTJ) ; 1 $, 22 May 1954, (MTJ) : 2 $,22
J rly 1954, (MTJ) ; 1 <^ , 18 August 1954, (H. G. Davis) ; 1 $ , 6 September 1954;
(HGD) ; 1 ^,28 May 1955, (HGD). 1 $, Willow Cr., LaCrosse-Hooper, 13 April
1951, (J. J. Davis) ; 1 $ , \ $ (common mount). Coulee, 16 September 1940, (F .C.
F armston). CANADA. British Columbia: 1 $, Oliver, 5 August 1959, (G. G. E.
Scudder) ; Robson, — 1 $, 6 May 1958, (H. R. Foxlee) ; 1 $,28 May 1958,
( HRF) ; 1 5 , 4 September 1958; 1^,4 October 1958, (HRF) .
Deposition of Type Material. — H olotype, California Academy of Sciences, CAS
#10167 — also, 4 paratypes; 8 paratypes, USNM; 4 paratypes, Cornell University;
2 paratypes. University of British Columbia; 10 paratypes, Museum of Department of
Eiological Control, University of California, Riverside.
Material was also examined from the following localities: — IDAHO. Kootenae
Co.: 2 $, Hauser Lake, 22 May 1959, (H. W. Homer). OREGON. Umatilla Co.:
16 $ , Cold Springs, 11 April 1954, (M. T. James and J. J. Davis). WASHINGTON.
G rant Co.: O’Sulbvan Dam, (M. T. James) — 4 $, 11 October 1953; 1 ^ , 22 July
1954. Same locality, — 5 $, 8 July 1954; 1 i5, 11 May 1954; 1 $,29 July 1954;
1 15 July 1955; 2^,6 September 1954, 2 ^ , 12 August 1956, (H. G. Davis).
Sepedon armipes Loew
Males we examined from west of the Continental Divide include: COLORADO
(1). Gunnison Co.: Gunnison. IDAHO (36). Blaine Co.: 10 mi. S. Galena, Gan-
nett. Boundary Co.: Bonners Ferry. Caribou Co.: Bancroft. Gem Co.: Emmett.
C coding Co.: Bliss, 5 mi. S. Hagerman. Jerome Co.: Hazelton. Lemki Co.: 2 mi
ri. Salmon (nr. pseudarmipes) . Lincoln Co.: 4 mi. E. Shoshone. MONTANA (2).
G allatin Co.: Bozeman. Granite Co.; Bennett ( 2 ). UTAH (50). Box Elder Co.:
Irigham City (Bear Riv. Ref.). Cache Co.: Clarkston, Logan, Logan Canyon,
Ilacksmith Fork & Curtis Creek. Duchesne Co.: Duchesne, Myton. Emery Co.:
Moab. Millard Co. : Delta. Salt Lake Co. : Midvale. Summit Co. : Coalville.
Uinta Co.: Leeton, Verval. Wasatch Co.: Current Creek. Weber Co.: Hooper,
Ogden, Ogden Canyon, Plain City. MEXICO. Sonora (I) : Imuris. CANADA.
British Columbia (1) : Merritt.
Males we examined from east of the Continental Divide include: COLORADO
(5). Boulder Co.: Valmont. El Paso Co.: Manitou, Colorado Springs. Larimer
(! o.: Fort Collins. Weld Co.: Roggen. MONTANA (6). Dawson Co.; Beaver
Creek. Gallatin Co.: Three Forks. Jefferson Co.; Whitehall. NEW MEXICO.
Rio Arbiba Co. (8): (Males and females). Sandoval Co. (16): (Males and fe-
rrales). WYOMING (13). Niobrara Co.: 12 mi. N. Lusk. Also, we examined an
I dditional 151 males from the states of Georgia, Illinois, Iowa, Kansas, Kentucky,
Maine, Maryland, Michigan, Minnesota, Nebraska, New Hampshire, New Jersey,
New York, Ohio, Pennsylvania, South Dakota, Tennessee, Texas, Vermont, Wiscon-
APRIL 1969] FISHER & ORTH TWO NEW SEPEDON
161
sin. CANADA (45). — New Brunswick, Nova Scotia, Ontario, Manitoba, Northwest
Territory (Good Hope) (plus 3 females).
Discussion
Geographic Distribution. — Sepedon capellei occurs in a horseshoe
shaped pattern from mideastern California northward, including western
Nevada, through southern and eastern Oregon, into Idaho, thence south
through Utah and eastern Nevada. Its occurrence is always west of the
Continental Divide.
Sepedon pseudarmipes occurs in the general vicinity of the Columbia
River basin in extreme northern Oregon and south central Washington,
thence easterly and northerly into northern Idaho and British Columbia.
Its occurrence, too, is always west of the Continental Divide.
Sepedon armipes is widely distributed throughout North America east
of and a shoit distance west of the Continental Divide. The northern-
most specimens we examined were 5 females and 3 males labeled Good
Hope, NWT., Canada, 27 August 1929, (0. Bryant) (CAS). This loca-
tion is approximately 66° N., 129° W., approximately 20 miles south of
the Arctic Circle and probably along the MacKenzie River. The south-
ernmost specimens we examined from North America were labeled Im-
uris, Sonora, Mexico, 11 May 1953, (R. C. Bechtel and E. I. Schlinger)
(UCB) — approximately 31° N. Latitude, and a male labeled Cameron
Co., Texas, 3 August 1928 (J. G. Shaw) (UK) — approximately 26° N.
Latitude. West of the Continental Divide S. armipes is sympatric with S.
capellei in Utah and southern Idaho. Both S. armipes and S. pseud-
armipes occur in northern Idaho and in British Columbia, but ac-
tual overlap of populations has not been established. Sepedon armipes
does not occur in California but it is probably present in extreme eastern
Oregon and Washington and northeastern Nevada. The distribution of
S. armipes as reported by Foote (1961) and Neff and Berg (1966) must
now be tempered to include S. capellei and S. pseudarmipes west of the
Continental Divide.
Of these three species only S. capellei occurs in California. The type
material was restricted to that state because, if it included specimens
from throughout the entire range of the species, the possibility would be
greater that discrete populations could be represented, thus complicating
future studies on subspeciation.
Seasonal Distribution. — Collecting dates, other than those for para-
types, have not been indicated because most occurred between April and
September, the expected spring and summer collecting period, and
hence probably do not reflect a true picture of seasonal spread. On 13
L62
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Fig. 14. Representative western north American distribution of the three species.
Good Hope, N.W.T., and Merritt, B. C., localities for S. armipes not shown. The
Continental Divide is indicated by:
September 1968, adult S. armipes and S. capellei were collected simul-
taneously (in the same aerial-sweep net) by R. E. Orth when sampling a
habitat near Logan, Utah. This record may indicate that the two spe-
cies can coexist in a rather restrictive habitat. It is assumed, therefore,
that species-separative mechanisms are functioning at adult and possibly
at subimago levels.
Atypical Specimens. — From a total of 1,357 males of the three spe-
cies examined, only four could not be placed definitely. These were from
APRIL 1969] FISHER & ORTH TWO NEW SEPEDON
163
areas of distributional overlap. One male, the only specimen from the
area, labeled 2 mi. N. Salmon, Idaho, 25 May 1959, 3,995' (B. A. Foote) ,
had point / (refer to Figs. 8-11) on the aedeagus closer to e than was
typical for pseudarmipes, yet / was too distant from e for typical S. arm-
ipes. Two males from a total of 25 taken at Cold Springs, Oregon, Uma-
tilla Co., 11 April 1954, (M. T. James and J. J. Davis), tended from S.
pseudarmipes toward S. capellei in that angle a h d oi the aedeagus was
greater than that in typical S. armipes and S. pseudarmipes. In these two
males, point / was situated characteristically as in S. pseudarmipes.
Another male, the only specimen from the area, labeled Troy, Idaho,
Latah Co., 5 September 1958, (B. A. Foote) also showed aedeagal
characteristics of S. capellei and S. pseudarmipes. Northerly from Cold
Springs and Troy only S. pseudarmipes occurred, and southerly only S.
capellei.
Acknowledgments
Without the generosity and cooperation of L. V. Knutson, Systematic
Entomology Laboratory, United States Department of Agriculture, this
paper would be noticeably narrower in scope. He discovered the form we
call S. pseudarmipes and independently recognized the form we call S.
capellei. We are grateful for the many specimens of the three taxa
provided through him for this comparative study from the following
sources; Canadian National Collection (Ottawa), K. J. Capelle, Cornell
University, Kent State University (Ohio), Oregon State College, Univer-
sity of Idaho, University of Minnesota, University of Wisconsin, Utah
State University, and Washington State University. Institutions from
which we were able to obtain additional material directly were Univer-
sity of California (Berkeley, Davis), California Academy of Sciences,
Kansas State University, University of Kansas, and University of British
Columbia.
Not only was the manuscript reviewed by L. V. Knutson, and J. C.
Hall but they gave freely of their counsel during its preparation. Others
who have either offered helpful taxonomic or editorial suggestions are
E. 1. Schlinger, J. W. MacSwain, E. R. Oatman, and G. C. Steyskal.
Literature Cited
Cresson, E. T. 1920. A revision of the Nearctic Sciomyzidae (Diptera: Acalyp-
tratae) . Trans. Amer. Entomol. Soc., 46(799) : 27-89. 3 plates. March, p. 86.
Fisher, T. W. 1966. A preliminary list of the Sciomyzidae of California (Diptera).
Pan-Pac. Entomol., 42(4) : 318-20.
Foote, B. A. 1961. The marsh flies of Idaho and adjoining areas (Diptera: Sciomy-
zidae). Amer. Midland Natur., 65(1) : 144-67.
164
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 2
Loew, H. 1859. Die nordamerikanisclien Arten der Gattungen Tetanocera und Sepe-
don. Wien. Entomol. Monatsschr. Nr. 10. Ill Band. October, pp. 289-300.
Helander, a. L. 1920. Review of the nearctic Tetanoceridae. Ann. Entomol. Soc.
Amer., 13(3) : 305-332. 1 plate. September.
Neff, S. E. and C. 0. Berg. 1966. Biology and immature stages of malacophagous
Diptera of the genus Sepedon (Sciomyzidae) . Agr. Exp. Sta., Blacksburg,
Va., Virginia Polytechnic Inst. Bull. 566, pp. 1-113.
Steyskal, G. C. 1951 (1950). The genus Sepedon in the Americas (Diptera:
Sciomyzidae). Wasmann J. Biol., 8(3) : 271-97.
1956. New species and taxonomic notes in the family Sciomyzidae (Diptera:
Acalyptratae) . Pap. Mich. Acad. Sci., Arts, Lett., vol. 41, part II. pp. 73-
87. (1955 meeting).
1960. New North and Central American species of Sciomyzidae (Diptera:
Acalyptratae). Proc. Entomol. Soc. Wash., 62(1) : 33-43.
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ENTOMOLOGICAL EXCHANGES AND SALES
This page is available to members who wish to advertise entomological exchanges,
sales, announcements, etc. We can offer Society members an ad in four issues for
^ 1.50 per line. Ads will be arranged in order of their receipt. Contact the Advertis-
ing Manager, W. H. Lange, Department of Entomology, University of California,
Davis, California 95616.
Research Request. — Am nearing completion a study of the Microlepidoptera as-
sociated with oaks in California. Would appreciate the opportunity to borrow any
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I
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,
and distribution. Excess pagination must he approved and may be charged to the author.
Papers are published in approximately the order that they are received. Immediate publi-
cation may be arranged after acceptance by paying publication costs. Papers of less than
a printed page may be published as space is available, in Scientific Notes.
Manuscripts for publication, proof, and all editorial matters should be addressed to the
Editor, Robbin W. Thorp, Department of Entomology, University of California, Davis,
California 95616.
Typing. — All parts of manuscripts must be typed on bond paper with double-spacing and
ample margins. Carbon copies or copies on paper larger than 8^ X H inches are not
accepted. Do not use all capitals for any purpose. Underscore only where italics are
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Essig, E. 0. 1926. A butterfly migration. Pan-Pac. Entomol.,
2 : 211 - 212 .
1958. Insects and mites of western North America. Rev.
ed. The Macmillan Co., New York. 1050 pp.
Abbreviations for titles of journals should follow the list of Biological Abstracts, 1966,
47(21) : 8585-8601.
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All articles are accepted for publication only on the basis of scientific merit
and without regard to the financial support of the author.
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Vol. 45 July 1969 No. 3
THE
Pan-Pacific Entomologist
In Memoriam
ROBERT LESLIE USINGER
1912-1968
LINSLEY — Robert Leslie Usinger, 1912-1968 167
ASHLOCK — Robert L. Usinger bibliography and list of names proposed 185
BUCKETT AND GARDNER — Revision of the chordeumid milliped genus
Tynomma Loomis from California (Chordeumida: Lysiopetalidea:
Lysiopetalidae) 204
BOHART AND GRISSELL — -New species of Psenini (Hymenoptera: Spheci-
dae) . 216
FISHER AND ORTH — A new Dictya in California, with biological notes
(Diptera: Sciomyzidae) 222
HYNES — The immature stages of the genus Rhabdomastix (Diptera: Tipuli-
dae) 229
SCIENTIFIC NOTES .. 237
ZOOLOGICAL NOMENCLATURE 240
BOOK REVIEWS 241
RECENT LITERATURE 243
SOCIETY NOTICES 244
SAN FRANCISCO, CALIFORNIA • 1969
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
THE PAN-PACIFIC ENTOMOLOGIST
EDITORIAL BOARD
R. W. Thorp, Editor
L. G. Linsley R. O. Schuster, Asst. Editor E. S. Ross
I . D. Hurd, Jr. D. L. Briggs, Editorial Assistant H. B. Leech
P. H. Arnaud, Jr. Treasurer W. H. Lange, Jr. Advertising
Published quarterly in January, April, July, and October with Society Proceed-
i igs appearing in the January number. All communications regarding nonreceipt
cf numbers, requests for sample copies, and financial communications should be
i ldressed to the Treasurer, Dr. Paul H. Arnaud, Jr., California Academy of
J ciences. Golden Gate Park, San Francisco, California 94118.
Application for membership in the Society and changes of address should be
cddressed to the Secretary, Dr. Marius S. Wasbauer, Bureau of Entomology, Cali-
fornia Department of Agriculture, 1220 N. St., Sacramento, California 95814.
The annual dues, paid in advance, are $5.00 for regular members of the Society
cr $6.00 for subscriptions only. Single copies are $1.50 each or $6.00 a volume.
I lake checks payable to Pan-Pacific Entomologist.
The Pacific Coast Entomological Society
Officers for 1969
K. S. Hagen, President Paul H. Arnaud, Jr., Treasurer
R. W. Thorp, President-elect Marius S. Wasbauer, Secretary
Statement of Ownership
I'itle of Publication: The Pan-Pacific Entomologist,
requency of Issue: Quarterly (January, April, July, October).
Location of Office of Publication, Business Office of Publisher, Publisher and Owner: Pacific Coast
Entomological Society, California Academy of Sciences, Golden Gate Park, San Francisco, California
94118.
Editor: Dr. Bobbin W. Thorp. Department of Entomology, University of California, Davis, California
95616.
Managing Editor and Known Bondholders or other Security Holders: None.
This issue mailed 14 October 1969.
Second Class Postage Paid at Lawrence, Kansas, U.S.A. 66044.
ALLEN PRESS, INC. LAWRENCE, KANSAS
Robert Leslie Usinger, 1912-1968
The Pan-Pacific Entomologist
VoL. 45 July 1969 No. 3
Uobtrt ILeslit flsingcr
E. Gorton Linsley
University of California, Berkeley
Robert Leslie Usinger, Professor of Entomology and Entomologist
in the Agricultural Experiment Station, University of California, Berk-
eley, died in San Francisco, California, on 1 October 1968 after an
extended illness. In the latter months of his life, while his health was
being undermined by cancer, he completed a term as Chairman of the
Division of Entomology and Acarology, Department of Entomology and
Parasitology, University of California, Berkeley, served brilliantly as
President of both the Entomological Society of America and the Society
of Systematic Zoology, dictated a fascinating accoimt of his life,^ and,
if his health had permitted, would have attended the Xlllth Interna-
tional Congress of Entomology in Moscow, 2-9 August 1968, as a guest
of the Academy of Sciences, USSR, and spent part of 1969 collecting
and studying the Naucoridae of New Guinea with support from the
National Science Foundation. To have collected at Wau, New Guinea,
would have been the realization of a life-time dream.
In addition to his University position, at the time of his death he was
affiliated with the California Academy of Sciences in San Francisco as
Research Associate, and, in the same capacity with the Bernice P. Bishop
Museum in Honolulu and the American Museum of Natural History in
New York. With his untimely death at the age of 55 years, the scientific
world has lost an outstanding biologist, an author of numerous zoolog-
ical books, entomological monographs, and writings on the systematics,
bionomics, biogeography and evolution of various groups of Hemiptera;
the academic world has lost a highly respected and inspiring teacher;
and the world in general a great humanitarian and leader.
Robert Usinger was born on 24 October 1912 at Fort Bragg, Mendo-
cino County, California, the eldest of two sons of Edith Johnson and
Henry Clay Usinger. His mother was of Scandinavian origin; the
Johnson family having come from Denmark in the early 1800’s. His
father was of German extraction; the Usinger family having originally
^ This document, written at my suggestion and with the constant encouragement of his wife, Martha,
will he published in the near future as Memoir Number 5 of the Pacific Coast Entomological Society.
The Pan-Pacific Entomologist 45 : 167 - 184 . July 1969
1()8 THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
ccme from Usingen in the Taunus Mts. Robert Usinger was one
o: the relatively few Californians who was a “native son of a native
sen.” When he was a few years old, the family moved from Fort Bragg
tc San Anselmo for a brief period and then to Oakland, where young
Rabert, or “Bob” as he was affectionately known to his many friends
all through his life, attended grammar school and high school before
entering the University of California, where he received the Bachelor
of Science degree in 1935 and the Ph.D. degree in 1939. His early
rremories in Fort Bragg centered around the days in the redwood
country, where grandfather Johnson was superintendent of the Union
L;imber Mill. He spoke of watching the loading of the big logs onto
Selling vessels for their trip to the port of San Francisco, and of standing
on “tip-toe” to reach the table to help pound out the whale steaks.
During his college years he managed to be active not only in matters
eiitomological, but was a long-time member of the Glee Club, and active
ii fraternity and campus affairs as well. It was in the Glee Club that he
first became acquainted with Richard M. Eakin, now Professor of
Zoology, University of California, Berkeley, who became a life-long
fiiend and who delivered the Memorial Address held in his honor at
ti e First Congregational Church of Berkeley on 14 October 1968.
In 1938, Usinger married Martha Boone Putnam, a college classmate,
who filled his life with companionship, understanding, and inspiration.
Ttieir two children, Roberta Christine Usinger (Mrs. Ronald Manuto),
Richard Putnam Usinger, and a grandchild, Benjamin Robert Manuto,
also survive him, besides a brother, Russell Usinger.
Among his many activities and honors, some of which will be placed
ii context in the discussion which follows, it should be mentioned
that he served in the Department of Entomology, Bernice Paui Bishop
IV.useum, Honolulu, during 1935 and 1936, the Department of Ento-
n ology, California Academy of Sciences, from 1936 to 1939, and
joined the faculty of the University of California as Instructor in
Entomology and Junior Entomologist in the Agricultural Experiment
S cation on the Davis Campus in 1939, moving to Berkeley (after service
ill World War II as Major in the Sanitary Corps, United States Public
health Service), where he rose to the ranks of Professor and Entomol-
ogist in 1953. From 1964 to 1968 he served as Chairman, Division
o : Entomology and Acarology and prior to his death, he was given
recognition by the University Administration as one of the most dis-
tinguished members of the Berkeley faculty. In 1948-49 he held a
National Institute of Health Special Research Fellowship at the British
IVfuseum of Natural History. He participated in the Marshall Islands
JULY 1969]
LINSLEY ROBERT LESLIE USINGER
169
Coral Atoll Study sponsored by the Pacific Science Board, National
Academy of Sciences in 1950, and the Laysan Expedition of 1961. From
1961 to 1963, he was chairman of the Board. In 1959 he participated
in the Congo Expedition of the Institut pour la Recherche Scientifique
en Afrique Centrale, and in 1964 he served as Director of the Galapagos
International Scientific Project. He was a member of the Permanent
Committee of the International Congress of Entomology and the Inter-
national Biological Union from 1953 until his death. He was decorated
by the King of Denmark (gold medal) in 1956 for his part in the
Galatea Expedition, and by the Government of Ecuador (medal and
award of merit) as a member of the Galapagos International Scientific
Project, and was named an honorary citizen of Guayaquil in 1964. He
also received keys to the City of New Orleans at the time of the National
meetings of the Entomological Society of America in that city. He
organized the Systematics Colloquium at the Missouri Botanical
Gardens in St. Louis in 1958 and Biological Sciences Program at the
Pacific Science Congress in Honolulu in 1961. He edited the Pan-
Pacific Entomologist from 1939-49 and was a member of various
scientific editorial boards, including that of “Pacific Insects,” of which
Volume H, no. 1, was dedicated to his memory in the following words:
“One-time staff-member of Bishop Museum, Member of Editorial Com-
mittee of PACIFIC INSECTS, Valued Advisor to the Entomology Depart-
ment, Bishop Museum, Authority on Hawaiian and Pacific Hemiptera,
Honored Teacher of many entomologists.” Usinger authored, co-au-
thored, and/ or edited Methods and Principles of Systematic Zoology
(1953), Aquatic Insects of California (1956), Classification of the
Aradidae (1959), Sierra Nevada Natural History (1964), General Zool-
ogy (4th edition, 1965, printed in many languages throughout the
world) , Monograph of the Cimicidae (1966) , Life in Rivers and Streams
(1967), and Elements of Zoology (3rd edition, 1968), and more than
200 other scientific publications (see appended bibliography by Ash-
lock) . In such an array of writings it is difficult to single out one for
special mention, but certainly his monograph of the “bedbugs” is a
classic in modern systematic biology and has been characterized by
Gressitt (1967, J. Med. EntomoL, 4: 24-25) as the most thorough study
of a family of insects for the world as a whole. Hoogstraal (1967, J.
ParasitoL, 53: 222) comments that “Everyone knows bedbugs but few
have adventured with them. Professor Usinger, who has known and
adventured around the world and in the laboratory with bedbugs inti-
mately, intellectually, and excitingly for almost 20 years, has presented
one of the finest books ever written on a family of parasitic insects.
170
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO, 3
Much of the information concerning the bedbug family derives from
th e results of Usinger’s six expeditions to Europe, South America, Cen-
tral America, Africa, and the Orient, and from his superb library, his
st mulating leadership of students, and his close collaboration with
numerous field collectors and outstanding foreign and American re-
searchers.”
He was a member of many scientific societies in addition to the
Entomological Society of America and the Society for Systematic Zool-
ogy, including the Pacific Coast Entomological Society (President,
1952, Honored Mlember, 1968), Linnaean Society of London (Fellow),
Royal Entomological Society of London (Fellow), American Associa-
tion for the Advancement of Science (Fellow), American Entomological
Society (Corresponding Member), Sociedad Entomologia Peruana
(Honorary Member) , Hawaiian Entomological Society and the Sierra
Club to give a partial list.
As a teacher. Professor Usinger’s influence was profound. Through
his courses in systematic entomology, aquatic insects and graduate sem-
ir ars, he touched the lives of generations of undergraduate and graduate
stidents leaving an impact that none can forget. Among those who
ec rned the Master of Science degree under his direction may be men-
tioned Paul D. Gerhardt, Alexander A. Hubert, Robert D. Lee, Tsuyoshi
Yamaguchi, Stephen W. Hitchcock, Norihiro Ueshima, Nualsri Sakda-
polrak, and Perry Turner; the Doctor of Philosophy degree, Ira La-
R vers, Willis W. Worth, Larry W, Quate, Toshiyuki Nishida, William
R Kellen, Jon L. Hering, Raymond E. Ryckman, Peter D, Ashlock, John
D Lattin, and Charles W. O’Brien. At the time of his death James
Haddock and Wayne Gagne were actively working toward their doc-
torate degrees.
As I write these words, inadequate as they are to characterize so
significant a life, the sun is rising over a beautiful coral-bounded lagoon
01 one of the many tropical oceanic islands that Usinger loved. As I
watch this magnificent sight, my mind goes back to the time when we
first met, in pre-college days, and the events that a few years later
b] ought him into the vast Pacific Ocean area and made such a profound
impression on his life and evolving scientific philosophy, as well as the
e\ents which entwined our personal lives from the moment of a fortu-
itous meeting while studying nature in the hills back of Oakland, Cali-
fcrnia, in the mid 1920’s.
When we first met, Usinger and I lived in different parts of the city
ar d were attending different high schools. Bob going to old “Uni” —
tbe then experimental and student teaching high school for the Univer-
JULY 1969]
LINSLEY ROBERT LESLIE USINGER
171
sity of California. However, each of us had a strong interest in natural
history. In his case this was nurtured by his Mother, and her profound
influence prompted Bob to comment in later years of his Mother —
“She was a wonderful person. Through her I knew only encourage-
ment.” It was Mrs. Usinger who first accompanied Bob at age 12 to
what was then Agricultural Hall to first talk with Professor Herms.
As Boy Scouts we had recently come under the influence of Brighton
(“Bugs”) C. Cain, a graduate of Stanford University, who was then
Naturalist for the Oakland Council, Boy Scouts of America. We
quickly became fast friends, and when Mr. Cain recognized that our
interest in insects was serious, he arranged for us to meet Dr. Edwin
C. Van Dyke, Professor of Entomology at the University of California,
and through Van Dyke we were introduced to E. P. Van Duzee at the
California Academy of Sciences. He also introduced us to other Bay
Area entomologists, amateur and professional, through the Pacific
Coast Entomological Society, including E. 0. Essig, Frank E. Blaisdell,
G. F. Ferris, E. R. Leach, J. 0. Martin and others. The collective impact
of these individuals on our lives at that stage was tremendous and the
influence of Van Dyke and later Mr. Van Duzee, upon the two of us
would be difficult to overestimate. The annual field trips of the Pacific
Coast Entomological Society were highlights of this period, and two of
these inspired us to write semi-popular articles on insect collecting in
California (items 12 and 20 in the Ashlock bibliography) and these
introduced us to another entomologist, T. D. A. Cockerell (through
whom Usinger developed an interest in fossil insects).
By mid high school days, Usinger’s taxonomic interests had focused
on the Hemiptera, prompted by E. P. Van Duzee and early contacts
with H. G. Barber and C. J. Drake. Before graduating from high
school he wrote his first paper on the subject, a revision of the genus
V anduzeeina in which he described two new species.
Between high school and college, Usinger had a truly marvelous
outdoor experience as a member of a select group of Boy Scouts from
the San Francisco Bay Area who spent a summer on a nature expedi-
tion in the southwest and Rocky Mountains under the direction of Mr.
Ansel Hall, then Chief Naturalist of the National Park Service. The
details of this experience cannot be recounted here (they are recorded
in part in “Scout Naturalists in the Rocky Mountains” by Fast, Kaiser
and Kelly ) , but they broadened his scientific horizons in a manner never
to be forgotten, and incidentally, greatly enhanced his collection of
insects, which eventually reached the number of 61,919 when presented
to the Entomological Museum (California Insect Survey) of the Univer-
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THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO.
3
sily of California, Berkeley. The first museum of Mesa Verde was
stf rted on this trip — with archaeological specimens found by this group
ard placed in the museum collection.
Usinger’s college years were eventful, as was every other experience of
his life, and I was fortunate to share many of them. However, space
will permit reference to only one or two important episodes. In 1933,
Howard Hinton, an undergraduate student interested in the Coleoptera,
pr oposed to Usinger that the two of them make a summer collecting trip
to Real de Arriba, Temascaltepec, Mexico, where his father, a mining
engineer and amateur botanist, lived. Since neither had any
money in those days of the “Great Depression,” Hinton proposed that
they finance the trip through the sale of specimens. By taking orders
from students and faculty they managed to scrape together one hundred
arid ten dollars. The investors were fully repaid by the collections
which Hinton and Usinger brought back, but the remarkable aspect of
the whole affair was that they actually made the trip with this amount
of money — by hitchhiking and riding in empty freight cars and enduring
the most uncomfortable, dirty, and sometimes dangerous kinds of cheap
trcnsportation and food. Bob said, however, “We always travelled in
the best company during those depression days — the bankers ! ” The
Hinton home itself was most comfortable and a cabin which he had
bi ilt for them was an ideal base from which to make collecting sorties.
Also, Mr. Hinton’s herbarium, identified at Kew Gardens in England,
Wc s most helpful as a source of host plant information. Usinger and
H;nton were especially excited about the aquatic collecting, Usinger
br nging back a fabulous collection of naucorids, a group in which he
maintained a lifelong interest, and Hinton an impressive collection of
D; yopdidae. Aside from the vicinity of Real de Arriba, significant gen-
eral insect collections were made at Tejupilco and in the tropical
lowlands along the Balsas River at Bejucos. He and Hinton, together,
collected 10,000 insects on this trip.
Another very important event occurred in 1935 when Usinger was
in /ited to spend a year at the Bishop Museum in Honolulu, while Elwood
Zinmerman, then Entomologist on the museum staff, returned to Berk-
eley to graduate. This was an exciting year, with weekend field trips
with 0. H. Swezey, F. X. Williams, F. Ray Fosberg and others who
in roduced him to the natural history of the Hawaiian Islands and
taught him how and where to find the native insects in the difficult
highland environment. Among the many highlights of his stay in the
islands was an expedition to Mauna Kea, Hawaii, sponsored by the
Hawaiian Academy of Sciences; graduate study in marine biology at
JULY 1969]
LINSLEY ROBERT LESLIE USINGER
173
the Waikiki Laboratory of the University of Hawaii during which he
discovered a new species of the marine water strider, Halobates, which
stimulated a lifelong interest in this group (later monographed by his
student Jon Hering) ; visits to Lanai, Kauai, Molokai and the other
islands, mostly on miserable cattle boats; joining the Hawaiian Ento-
mological Society to the Proceedings of which he contributed many
important papers; and selecting a little known but highly significant
group of Hemiptera, the Hawaiian Orsillini, for biosystematic and evolu-
tionary analysis for his doctoral dissertation at the University of
California. By means of almost superhuman effort and determination
he devoted his energy to field collecting and studies of these interesting
mirid bugs and his 167-page monograph of the Hawaiian Nysius,
published in 1942 (item 70, Ashlock bibliography) is eloquent testimony
of the success of his effort.
While in Hawaii, he also had the unique experience of spending a
month collecting on Guam with 0. H. Swezey, the “dean” of Pacific
Island Entomologists, and his wife under the joint sponsorship of the
Hawaiian Sugar Planters’ Association and the Bishop Museum. In the
Islands the party was hosted by the United States Navy and otherwise
unavailable facilities and resources were placed at their disposal. This
opportunity, as always, was used to its fullest by Usinger and advanced
his scientific interest in the Pacific even further.
On the return trip to Honolulu, he visited the Philippines, where he
studied important collections and made a number of highly productive
field trips; Lingnan University, Canton, China where he met W. E.
Hoffmann and examined his extensive collections ; and travelled by train
from Kobe, Japan to Tokyo, where he contacted entomologists at the
University. When he returned to California to accept a position at the
California Academy of Sciences in San Francisco and complete his
studies for the doctorate at Berkeley, Usinger had had an inspirational
infusion of oceanic natural history and evolution that remained a part
of him and colored his philosophy and interests throughout his life.
One final experience of student days at the University should be
mentioned, although many must be omitted. This was a 1937 trip to
the museums and University collections of eastern United States and
Canada, jointly planned by Usinger, Charles D. Michener and myself,
all three of whom were working on taxonomic problems requiring
examination of types in various institutions and also yearning to meet
the leading entomological systematists of that part of the country. We
borrowed my father’s car and gasoline credit card and with little
money, lots of enthusiasm and boxes of critical specimens we started
174
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
0 ;f. There were many humorous and not so humorous aspects to the
t] ip, but all in all it was a great success. Space will not permit recording
dstails, but highlights for Usinger included a memorable visit with
1 r. and Mrs. T, D. A. Cockerell in Boulder, Colorado and a collecting
t] ip to the fossil beds at Florissant, Colorado, stops at Lawrence, Kansas,
to see C. J. Drake and H. M. Harris, the United States National Museum,
tlie American Museum of Natural History, and the Provincial Museum
a Quebec in search of Provancher types.
A year before receiving the Ph.D. degree, the newly married Martha
Usinger, who was then teaching school in San Leandro, California,
a ong with all her other responsibilities typed his thesis, and shortly
a ter it was approved he was invited to join the faculty of the University
of California at Davis as a member of the Department of Entomology,
b e accepted the invitation in spite of a tempting alternative to succeed
E P. Van Duzee as Curator of Insects at the California Academy of
Sciences. These years were filled with demanding activities on a young
n an establishing a home in a new area, organizing courses, and work-
ing up material from his Pacific Island collections. While at Davis, the
L singers made many lifelong friends and that University campus
a ways held a special feeling for him. S. B. Freeborn, Professor of
Entomology (to mention only one), influenced the nature of Usinger ’s
serviee during World War H and provided loyal friendship during the
years of return to Berkeley. The faith and kindness of Stanley Freeborn
and his wife Mary was never forgotten by him.
The stay at Davis was broken by several short trips in western North
-America. These trips usually were planned to coincide with entomology
n eetings and did much to broaden Usinger’s horizons. They were
b ought to a close shortly after Pearl Harbor, and in February, 1943
( lis first child having been born in January, 1943), he was commis-
sioned as an officer in the U. S. Public Health Service, based at
A tlanta, Georgia, where his first assignment was to organize a mosquito
control program in the Savannah area followed immediately by assign-
n ent to Hawaii to develop means of coping with a dengue epidemic.
These years were filled with a variety of programs involving mosquitos
a id other arthropod vectors of disease as well as entomological activities
o' broader implications. He remained in service until early 1946, and
like other periods in Usinger ’s life the experience was a significant one.
4he years of “Malaria Control in War Areas” brought him back not
o ily to his beloved Pacific, but through the Caribbean as well.
At the close of Usinger ’s World War H service, he returned to the
L'niversity of California, this time, to the Berkeley campus, where I was
JULY 1969]
LINSLEY ROBERT LESLIE USINGER
175
teaching systematic entomology, forest entomology, and for a while,
insect morphology. Usinger’s first assignment was to offer a course in
aquatic insect biology and then, together, we set out to reorganize and ex-
pand the Berkeley program in entomological systematics and to lay the
groundwork for developing a center in this field. Usinger’s academic
career at Berkeley was brilliant, and will be chronicled elsewhere in
the official University of California publication “In Memoriam.” Aside
from his outstanding contributions as a teacher, there was scarcely a
significant area of University life to which he did not contribute,
whether cultural, policy-making, or quasi-administrative. Throughout
this aspect of his career, he remained an irrepressible, enthusiastic inno-
vator and promoter of worthwhile academic activities and programs.
Shortly after World War II, the National Academy of Sciences orga-
nized a Pacific Science Board, under the directorship of Harold J.
Coolidge, to sponsor research programs which could profitably make
use of naval facilities throughout the Pacific area. Usinger
was involved in the initial planning, served as chairman. West
Coast Advisory Committee, and from 1961 to 1963 as Chairman of
the Board. Perhaps the most ambitious of many projects to receive
early sponsorship was an anthropological survey of Micronesia.
During the academic year 1948—49, Usinger took sabbatical
leave from the University, and with support from the U. S. Public Health
Service, set up headquarters in London at the British Museum, to under-
take to develop a world classification of the Reduvioidea, a group
with which he had long been intrigued. He had long been interested
also in meeting and working with Dr. William China of the British
Museum, who, on seeing this boyish person introduce himself, simply
said: “I don’t believe it!”. His work carried him to the major
centers of hemipterology in Europe and permitted him to attend the
International Congress of Zoology in Paris and make a lifetime friend
of Dr. Jacques Carayon of the museum, and participate in the Inter-
national Congress of Entomology at Stockholm. At the former he
served as an Alternate Member of the International Commission on
Zoological Nomenclature which precipitated him into the stormy and
controversial period of Mr. Francis Hemming’s secretaryship, which
had started at the Lisbon Congress in 1935 and picked up momentum
at Paris (1948), Copenhagen (1953), and finally London (1958), be-
fore enough of the controversy was resolved to utimately permit
production of a new International Code of Zoological Nomenclature.
After the Vlllth International Congress of Entomology in Stockholm
(1948), which was a delightful experience, Usinger remained on to
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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
work at the Naturhistoriska Ricksmuseum, and it was at this time that
he began to assemble his remarkable library of Linnaeana. His hobby
o Linnaean biography, bibliography and anecdotes was shared in
n any ways with his students and friends and resulted in a number of
scholarly publications and in his becoming a Fellow of the Linnaean
Society. This Linnaean Library is now said to be second to only one
ii the world — that of a collector in London.
Upon Usinger’s return from Europe, he and I began sporadic but
hard work on a modern textbook of principles and methods of systematic
zoology. Progress however, was too slow for our liking and finally,
ir space made available to us by my father at the Chabot Observatory
ir Oakland, we went into complete seclusion and worked, almost, day
a id night to try to put together a complete first draft. When we were
well along on the project, we learned that Ernst Mayr, then at the
American Museum of Natural History in New York, was working on
a similar book, I flew back to his home and we exchanged drafts of
cliapters and concluded that we should pool our efforts, which we did,
a: id the book was published in 1953 and remained a standard textbook,
being translated into Russian and Arabic, until 1968, when a new
edition by Professor Mayr replaced it. Dr. Mayr kindly dedicated this
new work to Linsley and Usinger and sent the page proof of the dedi-
cation to Bob so he could know of it before he died.
During the next few years and up until his death, Usinger co-authored
a series of highly useful and very successful publishing ventures in-
volving textbooks, laboratory workbooks and laboratory manuals in
ti e field of general zoology which have been referred to by title
p eviously and in the Ashlock bibliography. He also co-authored Sierra
I\evada Natural History, an illustrated handbook highly popular with
nature lovers of all levels of background, published by the University
Press, 2nd ed., 1968.
More recently, Usinger wrote The Life of Rivers and Streams, an
outgrowth of years of interest in stream ecology, reflected not only in
h s technical publications, but dating back to his Aquatic Insects of
California (1956). This book is a volume of McGraw-Hill’s “Nature
Library.” In this series he also served as consultant on other nature
volumes.
In 1950, at the request of the U. S. Navy, the Pacific Science Board
w as asked to undertake an anthropological and ecological study of
coral atolls. Usinger agreed to serve as director of the first part of
tl e project, involving Arno Atoll in the Marshall Islands. The logistics
o’ the expedition were challenging and the experience fascinating.
JULY 1969]
LINSLEY ROBERT LESLIE USINGEiR
177
particularly from the anthropological standpoint, Usinger collected
insects, including the few aquatics, and subsequently published papers
on Heteroptera of the Marshall Islands (1951, 1952) and “Suggestions
for Collecting Terrestrial Invertebrates on Pacific Islands” in the
Handbook for Atoll Research (1953).
Usinger, like a number of his colleagues on the Berkeley campus,
had always been interested in the fossil record of insects, especially
as it related to the more modern floras and fauna. He had briefly
collected in the Florissant shales of Colorado (referred to above) and
had examined insects in Baltic amber. All of these, however, were of
Tertiary age and he longed to see Cretaceous material. In 1955, he
and Professor Ray F. Smith, with the sponsorship of the Office of Naval
Research, arranged to explore the Arctic slopes of the Brooks Range
for Cretaceous amber, of which there were fragmentary published and
word of mouth accounts. With the complicated logistics worked out
as they went along, and with the resources of the Arctic Research
Laboratory at their disposal, they made lonely and at times dangerous
trips down the Colville River above Umiat and the Kuk River back of
Wainwright. After much initial disappointment (due to the elusiveness
of the amber and to conflicting opinions as to where it could be found)
samples were found in quantity at several localities and before they
left Pt. Barrow Usinger and Smith had detected by hand lens perfect
examples of insect remains in this amber of Cretaceous age. The amber
collections are still being worked upon by specialists.
By 1957, Usinger’s studies of Cimicidae had progressed to the point
where some intensive field work in South America was indicated. With
his wife, Martha, accompanying him for part of the time, he undertook
a three months’ trip to that continent, collecting briefly in Panama
and northwestern South America, but concentrating primarily in Brazil,
Argentina and Trinidad. With logistic support, advice and assistance
from many resident entomologists, he managed to find in their natural
environments, the cimicids and blood-sucking reduviids he was search-
ing for and at the same time increase his general collection of Neo-
tropical Hemiptera. Shortly after his return he left for the Pacific
Science Congress in Thailand, where he not only participated actively
in the Congress but availed himself of field trips which contributed to
his knowledge of aquatic insects and Heteroptera in yet another part
of the world. His journey homeward took him once more to Japan
where he visited various entomologists and examined important Hemip-
tera collections.
The year 1959 was another banner one for Professor Usinger and
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THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 3
his family. He took sabbatical leave from the University, bought a car
hi England, and set up home in Paris, where he was provided a head-
qaarters, laboratory and office space, and the necessary equipment
to carry out systematic and biological studies of cimicids for his
n onograph, by Jacques Carayon at the Musee National d’Histoire
h aturelle with whom he collaborated on some phases of the project.
4 iter five months in France, Usinger and his family moved their head-
qaarters to Cairo, where Harry Hoogstraal had provided space and
logistical support for field studies through the facilities of the Navy
IMedical Research Unit laboratory stationed there. With leads provided
by Egyptian collectors affiliated with HoogstraaPs research program,
hs successfully explored in caves and other habitats of cimicids (in-
c uding pyramids) finding remarkable new species of these as well as
reduviids and other Heteroptera.
While headquartered at Cairo, the Usingers visited the Belgian Congo
a id set up a local base at the Institute pour la Recherche Scientifique
/ frique Centrale at Lwiro. Working out from Lwiro, Usinger visited
die various field stations maintained by the Institute, collected in caves
01 the shores of Lake Tanganyika, where he also collected aquatic
insects extensively. At Mont Hoyo he made additional finds of scien-
tific import and briefly visited Uganda. (Incidentally, be established
a friendship with pigmies in the Ituri Forest who also became friends
V ith Roberta and Richard Usinger, teaching the latter to shoot with
their primitive bows and arrows.) After returning to Cairo, the
I singers resumed their travels, coming back to the United States by
w ay of India, Hong Kong and Japan, thereby having encircled the
V orld.
In 1961, Usinger served as Organizer for Biology for the Pacific
Science Congress in Honolulu. Prior to the Congress, he and Professor
Paul R. Needham of the Department of Zoology, University of Cali-
fornia, Berkeley, with whom he and his students had collaborated for
niany years on limnological projects at the University’s Sagehen Creek
Experiment Station, attempted to introduce two species of mayflies
from California into streams around Kokee, Kauai, as food for rainbow
t; out which had been planted there much earlier hut were not doing
v eil. Although the mayflies apparently did not become established
during the project Usinger made a survey of potential local mayfly
predators and discovered, among other things, two new species belong-
ing to wingless genera of emesine reduviids.
Following the Congress, Usinger joined a group of biologists, with
C eorge Butler as the other entomologist, in a collecting trip to Laysan,
JULY 1969]
LINSLEY ROBERT LESLIE USINGER
179
one of the unique islands of the Hawaiian leeward chain, and to Kure
Atoll. As usual, the results were fruitful, adding in particular to the
knowledge of insular speciation in Nysius.
Early in 1962, once again pursuing his cimicid project, Usinger
traveled to southern Chile, where with the assistance of Luis Pena, he
traveled extensively in Auricaria and Nothofagus country in search of
primitive cimicids. After numerous disappointments and frustrations,
in a hollow opening in a Nothofagus tree, he discovered a remarkable
new genus of bat bugs, later called Bucimex, which proved to be inter-
mediate in its characters between the primitive Primicimex and the
more typical bedbugs of the genus Cimex. He considered this the most
significant discovery of the entire cimicid project. While in Chile, he
availed himself of the opportunity to make other collections of Heter-
optera of special significance. These included primitive lygaeid-like
forms, and subantarctic Aradidae and Teloridiidae. Before leaving
for home he also visited Punta Arena on the Straits of Magellan.
Possibly the greatest highlight of Usinger’s scientific career was his
participation in the 1964 Galapagos International Scientific Project, of
which he was officially Director and Professor Robert I. Bowman of
San Francisco State College, Co-Director. This conclusion is not only
attested by the fact that he devotes nearly 100 pages of his autobiog-
raphy to this event, but having had the privilege of traveling to and
from the Galapagos with him, sharing quarters at the Darwin Research
Station on Santa Cruz Island, and making together many local field
trips to various islands in the Archipelago, I know from personal ex-
perience that he attached great significance to this event. The following
paragraphs, quoted from his autobiography, not only record some of
the background of the project, hut reveal some of his feelings about it:
“The GISP, as planned by the University of California Extension
Division in collaboration with faculty members, was to bring together
scientists representing all the special fields of Galapagos research. A
symposium on existing knowledge of the Galapagos was held in Berkeley
and discussions of new lines of investigation occupied the hours while
sailing to the islands. Leading scientists from all over the world were
invited to participate and there was a group of young scientists to be
initiated into this natural laboratory of evolution. Important objectives
were the furtherance of conservation of the vanishing species in this
unique archipelago and support of the newly established laboratory of
the UNESCO sponsored Charles Darwin Foundation.
“In 1962 the project was still in the dream stage, since we had no
ship and no money. Then, through the vision of Capt. H. E. Richter,
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THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
Fig. 1. Usinger receiving certificate from the “Mayor” of Guayaquil in
recognition for his participation in the 1964 Galapagos Symposium.
S aperintendent of the California Maratime Academy, and Capt. C. B.
I owman it seemed possible that the 400 ft. training ship “Golden
I ear” might be fitted with extra bunks to carry sixty scientists to and
f ;om the islands as part of the annual training cruise for future
merchant marine officers. The second key decision that made the trip
f ossible was a grant from the National Science Foundation of $121,650.
(4her crucial sources of support were the Shell Oil Company (fuel oil
f Dr the Golden Bear through Dr. Harold J. Coolidge) and the Belvedere
Foundation of M. Kenneth Bechtel (funds for equipment and trans-
fortation). With such help we were definitely in business. Then came
rnexpected support that made an ordinary expedition into an experi-
eice unique in the annals of Galapagos exploration. The U. S. Navy
arranged for the USS Pine Island, with its sea planes and helicopters
to assist us, a U. S. Air Force C-130 plane transported over 100 par-
t cipants from Latin America, including members of the diplomatic
corps, destined for the dedication of the Darwin Laboratory. And the
/irmy contributed C-rations that made feasible the establishment of
field parties in the most remote and inaccessible places. The Pacific
^ cience Board through Dr. Harold J. Coolidge played a key role in all
JULY 1969]
LINSLEY ROBERT LESLIE USINGEiR
181
Fig. 2. Usinger in characteristic collecting attitude.
of these arrangements and also in final permission in Washington to
use the “Golden Bear.” The Government of Ecuador gave all-out sup-
port including air force planes, an LST to help unload the “Golden
Bear,” and the navy patrol boat that provided much interisland trans-
portation.
“Early in January, 1964, the invited scientists began to arrive from all
parts of the world. Sir Julian Huxley, evolutionist and close student of
Darwin’s work, was the keynote speaker at the Berkeley symposium. He
told of the crucial importance of the Galapagos experience in Darwin’s
career. Dr. Victor Van Straelen, Director of the Darwin Foundation,
who died three weeks after participating in the project, spoke of the
role of the Foundation in Galapagos research and the hope for imple-
menting conservation laws to preserve the unique plants and animals
of the Galapagos. After other more technical symposia the sailing
date, January 10, 1964, arrived. With banners flying and television
cameras turning, the “Golden Bear” cast her moorings and the Gala-
pagos International Scientific Project was launched. Accommodations
were described as “somewhat less than steerage” with temporary four-
132
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
Fig. 3. Usinger (right) and Linsley (left) amid scientific equipment after
landing of the 1964 Galapagos International Project at the Darwin Research
Station, Isla Santa Cruz.
decker canvas bunks crowded into a ward room, but the scientists were
net bothered by such trivia and the voyage proved to be a pleasure for
all. Meetings were held constantly to outline research plans, decide on
joint projects, and discuss controversial theories. Dr. R. I. Bowman,
co-director and old Galapagos hand, drew up the plans for field parties
a id Dr. Nathan Cohen, coordinator for University Extension, organized
tl e complicated logistics.
“On January 19 the “Golden Bear” and the USS Pine Island con-
verged on Academy Bay on the south Shore of Santa Cruz Island. The
Pine Island carried scientists, diplomats, and official representatives of
S( ientific agencies throughout the world who, earlier in the day, had
flown from Guayaquil to the airstrip on Baltra Island. Probably each
JULY 1969]
LINSLEY — ROBERT LESLIE USINGER
183
person had a different sensation on first sighting the Galapagos. My
own was colored by reading everything on the subject as a boy in
anticipation of joining an expedition that never materialized. Landing
on Baltra, I stepped out into a new world. To my eyes the south slope
of Santa Cruz Island was beautiful. From the shore-line mangroves to
the deep green highland forests it fulfilled my expectations completely.
But my eyes kept returning to the most unique Galapagos scene, the
weird cactus forest in the foreground. No pictures or words had fully
prepared me for this experience and, curiously, the islands never lost
their fascination in later weeks as I landed on one after another. Per-
haps islands appeal to the imagination as something neat and sharply
delimited, a clean break from the work-a-day world. But to the natu-
ralist each island is something more — a microcosm or world of its own.
It makes no difference how large or how small, there is a challenge to
discover how the various animals and plants have worked out their
destinies together. Usually the patterns of life are simpler than on the
mainland with more individuals and fewer species. In an attempt to
understand these intricate interrelations, each of us of whatever age
would ‘hit the beaches running’ during the next few weeks.”
Usinger did just that, whether wading ashore through the surf or
dismounting from a helicopter and thereby managing to collect on
every island in the group. It will be many years before the results of
his personal efforts are known and evaluated. A few statistics are already
indicative of the magnitude of his contribution to the ultimate knowl-
edge of Galapagos Hemiptera. Prior to the Expedition, he and I
assembled jointly as much information about the Galapagos insect fauna
as possible. We were able to accumulate a list of 618 species (Linsley
and Usinger, 1966) , of which 41 were Heteroptera, 38 Homoptera. As
of this writing, five post-expedition publications on these two groups
have come to my attention, reporting 34 new species and subspecies of
Homoptera and 26 of Hemiptera, most of which were collected by
Usinger or his protege, Peter Ashlock, with help from other members of
the Expedition.
In the summer of 1964, the Usingers went to Europe primarily to
attend the Entomological Congress in London, but also as a graduation
gilt to their daughter, Roberta who graduated from the University of
California, Berkeley, in June of 1964. They took advantage of this
opportunity to visit parts of the continent that they had not been to
previously (plus Spain, Gibraltar, and Tangier) and before and after
the Congress, Usinger studied cimicid types as well as those of certain
Galapagos Hemiptera. During the following summer, the Usingers
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
v)sited the Hawaiian Islands, largely for relaxation, but also to help
plan, as chairman, the Pacific Area International Biological Program.
However, he did find time for some collecting on Maui with Peter
Ashlock, J. W. Beardsley and D. Elmo Hardy.
During 1967, Usinger became aware of the first symptom of what
ti rned out to be his fatal illness. On 10 November 1967, he underwent
najor surgery from which he was told he would recover completely
and was encouraged to attend the meetings of the Society of Systematic
Zoology in late December in New York, over which he presided as
P resident. However, in April of 1968 the pain returned, and he under-
went another operation. It was found that all of the growth had not
been successfully removed after all, but had proliferated. From that
point on his health progressively deteriorated but his spirit remained
undaunted. With the constant attention and encouragement of his wife,
]V artha, he managed to cram a truly remarkable amount of activity into
tl e last few months of his life. Although the loss of Robert L. Usinger
will be keenly felt for years to come, few have lived such a full, exciting
a id rewarding life, and the impact of his warm personality, high level of
spirit, enthusiasm and personal concern for people will always be
rtmembered by those who knew him, no matter how casually. At the
suggestion of Martha Usinger, a memorial fund has been established
ir his memory at the University of California, Berkeley.
JULY 1969] ASHLOCK — USINGER BIBLIOGRAPHY 185
Robert L. Usinger Bibliography and List of Names Proposed
Peter D. Ashlock
University of Kansas, Lawrence
BIBLIOGRAPHY
1. 1930. Two new species of Vanduzeeina from California (Scutelleridae,
Hemipt.). Pan-Pac. EntomoL, 6(3) : 131-133.
2. 1931. A new species of Platylygus (Miridae, Hemiptera). Pan-Pac. Entomol.,
7(3): 129-130.
3. 1932. Miscellaneous studies in the Henicocephalidae (Hemiptera). Pan-Pac.
Entomol., 8(4) : 145-156.
4. 1933a. The male of Vanduzeeina slevini Usng. (Scutelleridae, Hemiptera).
Pan-Pac. Entomol., 9(1) : 30.
5. 1933b. Observations on the flight and length of life of drone bees. Ann.
Entomol. Soc. Amer., 26(2) : 239-246. D. E. Howell and R. L. U.
6. 1933c. A new record of Arizona Hemiptera. Pan-Pac. Entomol., 9(3) : 99.
7. 1933d. A new species of Gastrodes from California (Lygaeidae-Hemiptera) .
Pan-Pac. Entomol., 9 (3) : 127-128.
8. 1934a. New distributional and host plant records of Heteroptera for Cali-
fornia. 1. Pan-Pac. Entomol., 9(4) : 171-172.
9. 1934b. Two new records of Arizona Hemiptera. Pan-Pac. Entomol., 9(4) :
182.
10. 1934c. Blood sucking among phytophagous Hemiptera. Can. Entomol.,
66(5): 97-100.
11. 1934d. A simple device and method for blowing insect larvae. Bull. Brooklyn
Entomol. Soc., 29(4) : 168-170.
12. 1934e. Insect collecting in California. Pan-Pac. Entomol., 10(3) : 102-106.
E. G. Linsley and R. L. U.
13. 1935a. A second American species of the naucorid subfamily Laccorinae
(Hemiptera). Rev. Entomol., Brazil, 5(2) : 133-136.
14. 1935b. Stingless bees from central Mexico. Gleanings Bee Cult., 63(11) :
666-667.
15. 1935c. A new T ermatophylidea horn Mexico (Hemiptera: Termatophylinae) .
Entomol. News, 46(10) : 270-273.
16. 1936a. New distributional records of Hawaiian Heteroptera. Proc. Hawaiian
Entomol. Soc., 9(2) : 209-210.
17. 1936b. Book reviews. Wigglesworth, Insect physiology; Snodgrass, Insect
morphology. Proc. Hawaiian Entomol. Soc., 9 (2) : 210-212.
18. 1936c. The genus Geacoris in the Hawaiian Islands (Lygaeidae, Hemiptera).
Proc. Hawaiian Entomol. Soc., 9(2) : 212-215.
19. 1936d. Insect collecting on Lanai. Proc. Hawaiian Entomol. Soc., 9(2) :
216-218.
20. 1936e. Insect collecting in California — H. Foothills regions. Pan-Pac. En-
tomol., 12(2) : 49-55. E. G. Linsley and R. L. U.
21. 1936f. Studies in the North American Aradidae with descriptions of new
species (Hemiptera). Ann. Entomol. Soc. Amer., 29(3): 490-516.
22. 1937a. A new species of Aphelocheirus from Australia (Hemiptera, Naucori-
dae) . Aust. Zook, 8 (4) : 341-342.
The Pan-Pacific Entomologist 45: 185-203. July 1969
136
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
53. 1937b. Book review. Sweetman, The biological control of insects. Pan-Pac.
Entomol., 13(1 & 2) : 14.
24. 1937c. Notes on the biology of Hydrotrephes balnearius (Helotrephidae,
Hemiptera-Heteroptera) . Entomol. Mon. Mag., 73(879): 179-180 [3rd
series, 23 (272) ].
25. 1937d. A new species of Koanoa from the Hawaiian Islands (Miridae,
Hemiptera) . Proc. Hawaiian Entomol. Soc., 9(3): 427-439.
56. 1937e. Two new Pacific island species of Nysius (Lygaeidae, Hemiptera) .
Proc. Hawaiian Entomol. Soc., 9(3) : 439-442.
27. 1937f. A new name for Nysius monticola Kirkaldy (Lygaeidae, Hemiptera) .
Proc. Hawaiian Entomol. Soc., 9 (3) : 443.
28. 1937g. The Naucoridae of the Philippine Islands (Hemiptera). Philippine
J. Sci., 64(3) : 299^311.
29. 1938a. Book review. Jaques, How to know the insects. Pan-Pac. Entomol.,
14(1): 9.
30. 1938b. Dorsal abdominal scent glands in nymphs of Lygaeidae. Pan-Pac.
Entomol., 14(2) : 83.
31. 1938c. Biological notes on the pelagic water striders (Halobates) of the Ha-
waiian Islands, with a description of a new species from Waikiki (Gerridae,
Hemiptera). Proc. Hawaiian Entomol. Soc., 10(1): 77-84.
32. 1938d. A new genus and species of Orsillini from China (Hemiptera, Ly-
gaeidae). Pan-Pac. Entomol., 14(3) : 140-142.
33. 1938e. Review of the genus Gastrodes (Lygaeidae, Hemiptera). Proc. Calif.
Acad. Sci., (4)23(20): 289-301.
>4. 1939a. The family Polyctenidae (Hemiptera: Heteroptera) . Microentomol-
ogy, 4(1) : 1-50. G. F. Ferris and R. L. U.
35. 1939b. Key to North American Cimicidae, p. 92-93 ; Key to some predaceous
Reduviidae likely to be of medical importance, p. 100 ; Key to the principal
families of Hemiptera-Heteroptera of North America which contain preda-
ceous species, p. 101-102, In W. B. Herms, Medical Entomology. The Mac-
millan Company, New York, 3rd ed., xix + 582 p. [Also in 4th ed., 1950;
modified by others in 5th ed., 1961.]
36. 1939c. Book review. Duncan, A contribution to the biology of North Ameri-
can vespine wasps. Pan-Pac. Entomol., 15(2) : 75.
37. 1939d. Book review. DuToit, Our wandering continents. Pan-Pac. Entomol.,
15(2) : 88.
38. 1939e. Obituary. B. Preston Clark. Pan-Pac. Entomol., 15 (2) : 90.
39. 1939f. Descriptions of new Triatominae with a key to genera (Hemiptera,
Reduviidae). Univ. Calif. Publ. Entomol., 7 (3) : 33-56.
‘'^0. 1939g. Protepiptera, a new genus of Achilidae from Baltic amber (Hemip-
tera, Fulgoroidea) . Psyche, 46 (2-3) : 65-67.
41. 1939h. Van Dyke retirement. Pan-Pac. Entomol., 15 (4) : 187.
4'2. 19391. A new genus of Pacific island Enicocephalidae with new species from
the Hawaiian and Philippine islands (Hemiptera). Proc. Hawaiian En-
tomol. Soc., 10(2) : 267-270.
43. 1939] . Distribution and host relationships of Cyrtorhinus (Hemiptera; Miri-
dae). Proc. Hawaiian Entomol. Soc., 10(2) : 271-273.
^'4. 1940a. Fossil Lygaeidae (Hemiptera) from Florissant. J. Paleontol., 14 (1) :
79-80.
JULY 1969]
ASHLOCK — USINGER BIBLIOGRAPHY
187
45. 1940b. Obituary notices. Pan-Pac. EntomoL, 16 (1) : 12.
46. 1940c. A new Triatoma from Lower California (Hemiptera, Reduviidac) .
Pan-Pac. EntomoL, 16 (2) : 73-74.
47. 1940d. Obituary. Edward Payson Van Duzee. Pan-Pac. EntomoL, 16(3) :
123.
48. 1940e. The life and works of Edward Payson Van Duzee. Pan-Pac. EntomoL,
16 (4) : 145-177. E. 0. Essig and R. L. U.
49. 1940f. A new species of Aradus from Brazil (Hemiptera, Aradidae). Rev.
EntomoL, Brazil, 11 (3) : 639-642.
50. 1941a. Book reviews. Comstock, The spider book (revised by Gertsch) ; Com-
stock, An introduction to entomology. Pan-Pac. EntomoL, 17(1) : 36.
51. 1941b. Key to the subfamilies of Naucoridae with a generic synopsis of the
new subfamily Ambrysinae (Hemiptera). Ann. EntomoL Soc. Amer., 34
(1) : 5-16.
52. 1941c. Notes and descriptions of Neotropical Triatominae (Hempitera, Redu-
viidae). Pan-Pac. EntomoL, 17 (2) : 49-57.
53. 1941d. Obituaries. Pan-Pac. EntomoL, 17 (2) : 84.
54. 1941e. Problems of insect speciation in the Hawaiian Islands. Amer. Natur.,
75(758): 251-263. [Reprinted, “Symposium on the species concept,” In
Biological Symposia 4(4) : 281-293, Jaques Cattell Press, Lancaster,
Penna.]
55. 1941f. The present status and synonymy of some orsilline species (Hemip-
tera, Lygaeidae). Bull. Brooklyn EntomoL Soc., 36(3) : 129-132.
56. 1941g. The genus Oechalia (Pentatomidae, Hemiptera). Proc. Hawaiian
EntomoL Soc., 11 (1) : 59-93.
57. 1941h. A remarkable immigrant leptopodid in California. Bull. Brooklyn
EntomoL Soc., 36(4) : 164-165.
58. 1941i. Two new species of Aradidae from Baltic amber (Hemiptera) .
Psyche, 48(2-3) : 95-100.
59. 1941 j. Distribution of the Heteroptera of Oceania. Proc. Sixth Pac. Sci.
Congr., 4: 311-315.
60. 1941k. Distribution and host relationships of North and Central American
Triatominae. Proc. Sixth Pac. Sci. Congr., 4 : 459^61.
61. 1941/. Three new genera of apterous Aradidae (Hemiptera). Pan-Pac. En-
tomoL, 17(4) : 169-181.
62. 1941m. Rediscovery of Emesaya hrevicoxa and its occurrence in the webs of
spiders (Hemiptera, Reduviidae). Bull. Brooklyn EntomoL Soc., 36(5):
206-208.
63. 1942a. Key to the families of Heteroptera of North America, p. 269-273, In
E. 0. Essig, College Entomology. The Macmillan Company, New York, vii
+ 900 p.
64. 1942b. Notes on some flat bugs from the vicinity of Mt. Lassen, California
(Hemiptera, Aradidae). Pan-Pac. EntomoL, 18(2): 83-86. E. G. Linsley
and R. L. U.
65. 1942c. The Orsillini of New Zealand (Hemiptera, Lygaeidae). Trans. Roy.
Soc. N. Z., 72(1) : 41-52.
66. 1942d. A brachypterous Reduvius from Lower California (Heteroptera:
Reduviidae). EntomoL News, 53 (7) : 198-200.
188
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
il. 1942e. A new species of Oechalia from. Oaliu (Pentatomidae, Hemiptera).
Proc. Hawaiian Entomol. Soc., 11 (2) : 217-218.
C8. 1942f. A new Philippine leptopodid with remarks on the classification of the
family (Hemiptera). Bull. Brooklyn Entomol. Soc., 37(3): 103-106.
(9. 1942g. Notes and descriptions of East Asian Lygaeidae (Hemiptera). Lin-
gnan Sci. J., 20(2-4) : 249-250.
10. 1942h. The genus Nysius and its allies in the Hawaiian Islands (Hemiptera,
Lygaeidae, Orsillini) . Bishop Mus. Bull., 173. 167 p., 12 pis.
71. 1942i. Revision of the Termitaphididae (Hemiptera). Pan-Pac. Entomol.,
18(4): 155-159.
72. 1942j. The genus llyocoris StM (Hem., Naucoridae). Entomol. Mon. Mag.,
78: 241-242. [Reprinted J. Kansas Entomol. Soc., 17(2) : 76-77.]
13. 1942k. Key to the genera and species of Vesciinae with descriptions of two
new species (Hemiptera, Reduviidae). Rev. Entomol. Brazil, 13(3) : 290-296.
14. 1943a. Notes on the variation and distribution of Mesovelia mulsanti White
(Mesoveliidae, Hemiptera). Bull. Brooklyn Entomol. Soc., 37(5): 177-
178.
15. 1943b. An annectent genus of Cimicoidea from Baltic amber (Hemiptera).
Psyche, 49(3-4) : 41-46.
76. 1943c. Types of apterous Aradidae. Pan-Pac. Entomol., 19(2) : 60. [See
paper 127, fn. on p. 174.]
77. 1943d. Book review. Essig, College entomology. Pan-Pac. Entomol., 19(2) :
62.
78. 1943e. A new name for a New Zealand aradid. Pan-Pac. Entomol., 19(2) :
74.
79. 1943f. Book review. Ferris, Atlas of the scale insects of North America.
Pan-Pac. Entomol., 29 (2) : 79-80.
fO. 1943g. A new species of Campylomma from the Hawaiian Islands (Hemip-
tera: Miridae). Proc. Hawaiian Entomol. Soc., 11(3) : 287-288.
11. 1943h. A taxonomic note on Aradus depictus Van Duzee. Pan-Pac. Entomol.,
19(4): 138.
f 2. 1944a. A revised classification of the Reduvioidea with a new subfamily from
South America (Hemiptera). Ann. Entomol. Soc. Amer., 36(4): 602-
618.
13. 1944b. Entomological phases of the recent dengue epidemic in Honolulu.
Public Health Rep., 59(13) : 432-420.
£4. 1944c. Literature review. Soper and Wilson, Anopheles gamhiae in Brazil,
1930-1940. Monthly Report, Malaria Control in War Areas, U. S. Public
Health Service, Atlanta, Ga., March 1944, p. 12. [Reprinted, Mosquito
News, 4(4): 111-114.]
£5. 1944d. Host plants of western Lygus (Hemiptera, Miridae). Pan-Pac. En-
tomol., 20 (2) : 78.
£5. 1944e. Review of “A symposium on a national program for the control of
malaria.” Mosquito News, 4(2) : 44-A7.
£7. 1944f. Further notes on the habits of some flat bugs with a description of the
male of Aradus patibulus Van Duzee. Pan-Pac. Entomol., 20(3) : 111-114.
E .G. Linsley and R. L. Lf.
83. 1944g. Dengue control in the Hawaiian Islands. Malaria Control in War
JULY 1969]
ASHLOCK USINGER BIBLIOGRAPHY
189
Areas, U. S. Public Health Service Field Bulletin, Atlanta, Ga., p. 2-5.
W. E. Gilbertson and R. L. U.
89. 1944h. Heteroptera of Canton Island. Proc. Hawaiian Entomol. Soc., 12
(1): 147-148.
90. 1944i. Nomenclature of the human malaria parasites. Science, 100 (2692) :
190-192. C. W. Sabrosky and R. L. U.
91. 1944j. Yellow fever from the viewpoint of Savannah. Ga. Hist. Quart., 28(3) :
143-156.
92. 1944k. The Triatominae of North and Central America and the West Indies
and their public health significance. Public Health Bull., No. 288, iv +
83 p. [12 pis.].
93. 19441. Nomenclature of the genus Nysius and its allies (Lygaeidae: Heterop-
tera). Proc. Entomol. Soc. Wash. 46(9): 260-262. R. L. U. and R. I.
Sailer.
94. 1945a. Distribution of Icteronysius with descripition of a new subspecies
from Haleakala (Hemiptera, Lygaeidae). Proc. Hawaiian Entomol. Soc.,
12(2): 405-406.
95. 1945b. Classification of the Enicocephalidae (Hemiptera, Reduvioidea). Ann.
Entomol. Soc. Amer., 38(3) : 321-342 .
96. 1945c. Notes and descriptions of America Polyctenidae (Hemiptera). Pan-
Pac. Entomol., 21(4) : 121-124. G. F. Ferris and R. L. U.
97. 1945d. Biology and control of ash plant bugs in California. J. Econ.
Entomol., 38(5) : 585-591.
98. 1945e. Notes on the genus Cryptostemma with a new record for Georgia and
a new species from Puerto Rico (Hemiptera: Cryptostemmatidae). Entomol.
News, 56(9): 238-241.
99. 1946a. Review of the genus Saldoida with new records for Georgia and Vir-
ginia (Hemiptera, Saldidae). Bull. Brooklyn Entomol. Soc., 40(4) : 116-
118.
100. 1946b. Notes on Cuban Triatominae (Hemiptera, Reduviidae). Pan-Pac.
Entomol., 22(1) : 19-20. Reprinted in English and Spanish, 1948, Rev.
Kuba Med. Trop. ParasitoL, 4(8) : 163-164.
101. 1946c. Notes and descriptions of Ambrysus Stal with an account of the life
history of Ambrysus morman Montd. (Hemiptera, Naucoridae) . Univ.
Kans. Sci. Bulk, 31, Pt. 1(10) : 185-210.
102. 1946d. Notes and descriptions of Ceratocomhus (Hemiptera: Cryptostem-
matidae). Proc. Hawaiian Entomol. Soc., 12 (3) : 633-636.
103. 1946e. Notes on the synonymy and classification of the Enicocephalidae.
Ann. Entomol. Soc. Amer., 39(2) : 170.
104. 1946f. Biology and control of the Ash lace bug Leptoypha minor. J. Econ.
Entomol., 39(3) : 286-289.
105. 1946g. Fascicle V, Polyctenidae, p. 1-18, In W. E. China, ed.. General Cata-
logue of the Hemiptera. Smith College, Northampton, Mass.
106. 1946h. Household bugs. Pest Control and Sanitation., 1(12): 19.
107. 19461. Hemiptera Heteroptera of Guam, In Insects of Guam, II, Bishop Mus.
Bull. No. 189: 11-103.
108. 1946] . Educational activities as related to the extended malaria control
program: A progress report. J. Nat. Malaria Soc., 5(4): 245-251. W.
190
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
S. Boyd and staff [Staff; L. B. Hall, F. M. Hemphill, R. L. U., H. C.
Knutson, J. C. Terrill, Jr.].
109. 1947a. Entomological technique. Pest Control and Sanitation, 2(3) : 14-17.
110. 1947b. Notes on Graptostethus in Hawaii (Hemiptera: Lygaeidae). Proc.
Hawaiian Entomol. Soc., 13(1) : 107-108.
111. 1947c. Classification of the Cryphocricinae (Hemiptera: Naucoridae). Ann.
Entomol. Soc. Amer., 40(2) : 329-343.
112. 1947d. Native hosts of the Mexican chicken bug, Haemato siphon inodor a
(Duges) (Hemiptera, Cimicidae) . Pan-Pac. Entomol., 23(3): 140.
113. 1948a. Ash bug control. Calif. Agr., 2(3) : 13.
114. 1948b. Biology of aquatic and littoral insects. University of California Sylla-
bus Series, Syllabus SS, 244 p. R. L. U., I. LaRivers, H. P. Chandler, and
W. W. Wirth.
115. 1948c. Book review. Torre-Bueno, A synopsis of the Hemiptera-Heteroptera
of America north of Mexico. Bull. Brooklyn Entomol. Soc., 42(5) : 173-
175.
116. 1948d. A new genus and species of Aradidae from Fiji (Hemiptera). Proc.
Hawaiian Entomol. Soc., 13(2) ; 261-264.
1 17. 1948e. An all-purpose insect net made of nylon. Bull. Brooklyn Entomol.
Soc., 43(2) : 67-68.
118. 1949a. A new species of Chopardita Villiers from the Anglo-Egyptian Sudan
with notes on the subfamily Vesciinae (Hemiptera-Reduviidae) . Ann.
Mag. Natur. Hist., (12) 1; 598-604. W. E. China and R. L. U.
119. 1949b. War-time dispersal of Pacific island Nysius (Hemiptera: Lygaeidae).
Proc. Hawaiian Entomol. Soc., 13(3) : 447.
120. 1949c. A new genus of Hydrometridae from the Belgian Congo, with a new
subfamily and a key to the genera. Rev. Zool. Bot. Afr., 41(4) : 314-319.
W. E. China and R. L. U.
121. 1949d. Book review. Zimmerman, Insects of Hawaii. Entomologist, 82
(April): 95-96.
122. 1949e. The generic identification of Nearctic reduviid nymphs (Hemiptera).
Ann. Entomol. Soc. Amer., 42(3) : 273-278. S. B. Fracker and R. L. U.
123. 1949f. Classification of the Veliidae (Hemiptera) with a new genus from
South Africa. Ann. Mag. Natur. Hist., (12)2; 243-254. W. E. China and
R. L. U.
124. 1949g. A new genus of Tribelocephalinae from Fernando Poo (Hemiptera
Reduviidae). Ann. Mus. Civico Storia Natur. Genova, 64: 43-47. W. E.
China and R. L. U.
125. 1949h. Aquatic Hemiptera from Wicken Fen. Entomol. Mon. Mag., 85:311-
312. R. L. U. and E. S. Brown.
126. 1950a. Hermatobates haddonii Carpenter from the Marquesas Islands (He-
miptera: Gerridae). Proc. Hawaiian Entomol. Soc., 14(1): 53. W. E.
China and R. L. U.
127. 1950b. The origin and distribution of apterous Aradidae. Proc. Eighth Int.
Congr. Entomol. Stockholm, 1948: 174-179.
128. 1950c. On the identity of Heterocleptes Villiers 1948 and Hydrohatodes
China and Usinger 1949 (Hemiptera-Heteroptera Families Reduviidae and
Hydrometridae). Rev. Zool. Bot. Afr., 43(3) : 336-344. W. E. China, R. L.
U., and A. Villiers.
JULY 1969]
ASHLOCK USINGER BIBLIOGRAPHY
191
129. 1950d. In memoriam. William Brodbeck Herms, 1876-1949. Univ. Calif.
Faculty Bull., 19(12) : 119-120. S. B. Freeborn, H. F. Grey, R. T. Legge,
and R. L. U.
130. 1951a. Heteroptera of the Marshall Islands. Proc. Hawaiian Entomol. Soc.,
14(2): 315-321.
131. 1951h. A revised classification of Pacific Island Cyrtorhinus with a new spe-
cies from Fiji (Hemiptera, Miridae) . Comment. Biol., 12(8): 1-5.
132. 1951c. The history of the Pacific Coast Entomological Society. Pan-Pac.
Entomol., 27(3): 97-119. E. 0. Essig, P. D. Hurd, D. D. Jensen, H. B.
Leach, E. G. Linsley, J. W. MacSwain, R. C. Miller, R. L. U., and E. C.
Van Dyke.
133. 1951d. Proposed use of the plenary powers to designate type species of the
genera ‘Wysms” Dallas, 1852 and ‘‘‘'Artheneis” Spinola, 1837. Bull. Zool.
Nomencl. 2(11) : 313-314. R. L. U. and R. I. Sailer.
134. 1951e. Book review. Ferris, The sucking lice. Pac. Discovery 4(6) : 28-29.
135. 1952a. New species and additional records of Heteroptera from the Marshall
Islands. Proc. Hawaiian Entomol. Soc., 14(3) : 519-524.
136. 1952b. A new genus of Chryxinae from Brazil (Hemiptera: Reduviidae).
Pan-Pac. Entomol., 28(1) : 55-56.
137. 1952c. Obituary. Brighton Clark Cain. Pan-Pac. Entomol., 28(3) : 125.
138. 1952d. A new species of Carayonia from Ceylon (Hemiptera, Reduviidae,
Saicinae). Entomologist, 85 (Sept.) : 212-213.
139. 1952e. Two new Orsillini from Kilimanjaro (Hemiptera: Lygaeidae) . Proc.
Roy. Entomol. Soc. London, (B) 21 (9-10) : 140-146.
140. 1952f. Neotypes. Syst. Zook, 1 (4) : 171-173.
141. 1953a. Methods and Principles of Systematic Zoology. McGraw-Hill Book
Company, New York, ix -(- 328 p. E. Mayr, E. G. Linsley, and R. L. U.
142. 1953b. The insect life of Arno [Marshall Islands]. Atoll Res. Bull. (Pac.
Sci. Bd., Nat. Res. Council), No. 15: 1-28.
143. 1953c. Suggestions for collecting terrestrial invertebrates on Pacific Islands,
In Sachet and Fosberg, Handbook for Atoll Research (2nd prelim, ed.).
Atoll Res. Bull. (Pac. Sci. Bd., Nat. Res. Council), No. 17: 69-72.
144. 1953d. A new species of Aphraniola from Cambodia (Hemiptera, Cimici-
dae) . Rev. Fr. Entomol., 20(2) : 138-139. G. F. Ferris and R. L. U.
145. 1953e. Notes on the genus Metrobates in California with a description of a
new species (Hemiptera: Gerridae). Pan-Pac. Entomol., 29(3): 178-179.
146. 1953f. Book review. Pennak, Fresh-water invertebrates of the United States.
Pan-Pac. Entomol., 29(4) : 221-222.
147. 1954a. Class Insecta (part of superclass Labiata) , p. 189-195, In S. F. Light
et ah. Intertidal Invertebrates of the Central California Coast. Univ. Calif.
Press, Berkeley and Los Angeles, xiv -|- 446 p.
148. 1954b. Revision of the genus Chelonocoris Miller (Hemiptera, Aradidae).
Zool. Meded., 32(22): 259-278.
149. 1954c. Obituary. Howard Madison Parshley (1884-1953) . Pan-Pac. Entomol.,
39(1): 1-4.
150. 1954d. A new genus of Aradidae from the Belgian Congo, with notes on
stridulatory mechanisms in the family. Ann. Mus. Congo Tervuren, in-d”,
Zool., 1, 1954-Miscellanea Zoologica, H. Schoutenden, p. 540-543.
192
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
151. 1955a. The role of insects in sewage beds. Hilgardia, 23(10) : 263-321. R.
L. U. and W. R. Kellen.
152. 1955b. Elements of Zoology. McGraw-Hill Book Company, Inc., New York,
vi -j- 522 p. T. I. Storer and R. L. U.
153. 1955c. Appendix: Changes in the international rules of zoological nomen-
clature, p. 285-292 [supplement to No. 141, 1953a], Methods and Principles
of Systematic Zoology. McGraw-Hill Book Company, Inc., New York, vii -j-
366 [1955 printing].
1 )4. 1955d. Hemiptera, p. 534-536, In A Century of Progress in the Natural Sci-
ences. California Academy of Sciences, San Francisco, vii -|- 807 p.
155. 1956a. A drag-type riffle-bottom sampler. Progr. Fish-Cult., 18(1) : 42-44.
R. L. U. and P. R. Needham.
1( 6. 1956b. The stability of scientific names. Ann. Rev. Entomol., 1 : 59-70.
157. 1956c. Variability in the macrofauna of a single riffle in Prosser Creek,
California, as indicated by the Surber sampler. Hilgardia, 24(14) : 383-
409. P. R. Needham and R. L. U.
1)8. 1956d. Aquatic Insects of California. Univ. Calif. Press, Berkeley and Los
Angeles, x -|- 508 p. [Ed., author of preface, p. vii-viii, introduction, p.
3-A9, and chapter on Hemiptera, p. 182-228.]
If 9. 1957a. Arctic amber. Pac. Discovery, 10(2) : 15-19. R. L. U. and R. F. Smith.
1()0. 1957b. Notes on and descriptions of Cimicidae (Hemiptera). Microentomol-
ogy, 22(1) : 1-37. G. F. Ferris and R. L. U.
1()1. 1957c. Obituary. Edwin Cooper Van Dyke, 1869-1952, p. 180-182, In In
Memoriam. University of California. R. L. U., E. 0. Essig, and E. G.
Linsley.
1(2. 1957d. Notes on marine water striders of the Hawaiian Islands (Hemiptera:
Gerridae). Proc. Hawaiian Entomol. Soc., 16(2) : 281-283. R. L. U. and
J. L. Herring.
1( 3. 1957e. A new genus and two new species of myrmecomorphic Miridae from
North America (Hemiptera). Wasmann J. Biol., 15(1): 1-13. J. C. M.
Carvalho and R. L. U.
1(4. 1957f. General Zoology. McGraw-Hill Book Company, Inc., New York, 3rd
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and Russian.]
1(5. 1957g. Hemiptera Heteroptera Cimicidae. A new genus and species from
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L. U.
1(6. 1957h. Heteroptera: Aradidae. Insects Micronesia, 7(3): 117-172. R.
Matsuda and R. L. U.
1(7. 1957i. Marine insects. In Treatise on Marine Ecology and Paleoecology. Ecol-
ogy. Mem. Geol. Soc. Amer., 1(67): 1177-1182.
1(8. 1958a. Harzwanzen or “resin bugs” in Thailand. Pan-Pac. Entomol., 34(1) :
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1(9. 1958h. Laboratory Manual for Zoology, McGraw-Hill Book Company, Inc.,
New York, 3rd ed., x -(- 181 p. T. 1. Storer and R. L. U.
IIO. 1958c. A new genus of Reduviidae from the Hawaiian Islands (Hemiptera).
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171. 1958d. Cretaceous and Tertiary insects in Arctic and Mexican amber. Proc.
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Tenth Int. Congr. EntomoL, 1 [1956] : 851. P. D. Hurd, R. F. Smith, and
R. L. U.
172. 1958e. Notes on the higher classification of the Reduviidae, with the descrip-
tion of a new tribe of the Phymatinae. Rev. Zool. Bot. Afr., 57 (3-4) : 256-
281. J. Carayon, R. L. U., and P. Wygodzinsky.
173. 1958f. Book review. Borradaile and Potts, The Invertebrata, A manual for
the use of students. Pan-Pac. EntomoL, 34(4) : 232.
174. 1959a. Classification of the Aradidae (Hemiptera-Heteroptera) . British Mu-
seum, London. 410 p.
175. 1959b. A new species of Stricticimex from Kenya (Hemiptera: Cimicidae) .
Ann. EntomoL Soc. Amer., 52(1) : 81-82. G. F. Ferris and R. L. U.
176. 1959c. Obituary. Gordon Floyd Ferris, 1893-1958. Pan-Pac. EntomoL, 35
( 1 ): 1 - 12 .
177. 1959d. The identity of Nysius coenosulus Stal. Proc. Hawaiian EntomoL
Soc., 17(1): 92.
178. 1959e. Revision of the Metrargini (Hemiptera, Lygaeidae). Proc. Hawaiian
EntomoL Soc., 17 (1) : 93-116. R. L. U. and P. D. Ashlock.
179. 1959f. Laboratory Workbook for Zoology. McGraw-Hill Book Company, Inc.,
New York, 188 p. T. 1. Storer and R. L. U.
180. 1959g. A new species of Stricticimex from Ruanda-Urandi (Hemiptera:
Cimicidae). Rev. Zool. Bot. Afr., 60(1-2) : 61-64.
181. 1959h. New species of Cimicidae (Hemiptera). Entomologist, 92(1157):
218-222.
182. 1959i. The significance of ecologically equivalent species of Hemiptera that
coexist in nature [abstract only]. Tenth Int. Congr. EntomoL, 1: 101.
183. 1959j. Linnaeus and the development of the International Code of Zoologi-
cal Nomenclature. Syst. Zool., 8(1) : 39^7. E. G. Linsley and R. L. U.
184. 1960a. Book review. Southwood and Leston, Land and water bugs of the
British Isles. Ann. EntomoL Soc. Amer., 53(3) : 442.
185. 1960b. Enicocephalidae. Heteroptera. Insects Micronesia, 7(5): 219-230.
R. L. U. and P. Wygodzinsky.
186. 1960c. Reduviidae. Heteroptera. Insects Micronesia, 7(5): 231-283. P.
Wygodzinsky and R. L. U.
187. 1960d. Cimicidae. Heteroptera. Insects Micronesia, 7(5): 285-286. R. L. U.
and G. F. Ferris.
188. 1960e. Obituary. Otto Herman Swezey. Pan-Pac. EntomoL, 36(3) : 151-153.
R. L. U. and E. C. Zimmerman.
189. 1960f. New species of Cyrtopeltis from the Hawaiian Islands with a revised
key (Hemiptera: Miridae). Proc. Hawaiian EntomoL Soc., 17(2): 249-
254. J. C. M. Carvalho and R. L. U.
190. 1960g. The Cimicidae of Egypt (Hemiptera). J. Egypt. Public Health Ass.,
35(2): 81-89.
191. 1960h. Hemiptera, p. 400-410, In Encyclopedia of Science and Technology.
McGraw-Hill Book Company, Inc., New York.
192. 1960i. Observations on the biology of Chiloxanthus stellatus (Curtis) and
C. arcticus (Salberg) (Hemiptera: Saldidae) . Pan-Pac. EntomoL, 36(4):
189-190.
193. 1961a. New species of bed bugs from Africa. Calif. Agr., 15 (4) : 2.
]94
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
194. 1961b. Elements of Zoology, McGraw-Hill Book Company, Inc., New York,
2nd ed., viii -|- 464 p. T. I. Storer and R. L. U.
195. 1961c. Obituary. Stanley Barron Freeborn, 1891-1960, In In Memoriam,
p. 35-37. University of California. E. C. Voorhies, T. I. Storer, and R. L. U.
196. 1961d. Taxonomy, p. 992-997, In Encyclopedia of the Biological Sciences.
Reinhold Publishing Corp., New York. E. G. Linsley and R. L. U.
197. 1961e. Lygus Hahn, 1833 (Insecta, Hemiptera) ; Proposed designation under
the plenary powers of a type species in harmony with accustomed usage.
Z. N. (S.) 1062. Bull. Zool. Nomencl. 18(4) : 281-284. J. C. M. Carvalho,
H. H. Knight, and R. L. U.
198 1961f. Obituary. Stanley Barron Freeborn, 1891-1960. J. Econ. Entomoh,
54(5): 1069-1070.
199. 1961g. Book review. Gressitt, Problems of the zoogeography of Pacific and
Antarctic insects. Ann Entomoh Soc. Amer., 54(6) : 928.
210. 1962a. Foreward, p. v-viii. In Henry Walter Bates, The Naturalist on the
River Amasons Univ. Calif. Press, Berkeley and Los Angeles.
201. 1962b. Book review. The international code of zoological nomenclature. Bull.
Entomoh Soc. Amer., 8(4) : 207-208.
202. 1963a. A new bat bug from southern Chile (Hemiptera: Cimicidae). Pan-
Pac. Entomoh, 39(1) : 51-55.
203. 1963b. Sierra Nevada Natural History, An illustrated handbook. Univ.
Calif. Press, Berkeley and Los Angeles. 374 p., 24 col. ph T. 1. Storer and
R. L. U.
2)4. 1963c. Classification of the Holoptilinae and description of the first repre-
sentative from the New World (Hemiptera: Reduviidae) . Proc. Roy.
Entomoh Soc. London, (B) 32(3-4) : 47-52. P. Wygodzinsky and R. L. U.
205. 1963d. Insects and other arthropods from Kure Island. Proc. Hawaiian
Entomoh Soc., 18(2) : 237-244. G. D. Butler, Jr., and R. L. U.
2 )6. 1963e. Insects and other invertebrates from Laysan Island. Atoll Research
Bull. 98 (Pac. Sci. Bd., Nat. Res. Council), 30 p. G. D. Butler, Jr., and
R. L. U.
217. 1963f. Animal distribution patterns in the tropical Pacific, p. 255-261, In J.
L. Gressitt, ed.. Pacific Basin Biogeography: A symposium. Bishop
Mus. Press, Honolulu.
208. 1963g. The evolutionary significance of island and host plant transfers in
the genus Nysius and allies (Hemiptera, Lygaeidae) . Proc. Ninth Pac. Sci.
Congr. [Bangkok, Thailand], 9: 26-28.
209. 1964a. The role of Linnaeus in the advancement of entomology. Ann. Rev.
Entomoh, 9: 1-16, portrait.
2 .0. 1964b. The genus Reduviius Fabricius in western North America (Reduvi-
idae, Hemiptera, Insecta). Amer. Mus. Novitates, 2175: 1-15. P. Wygodzin-
sky and R. L. U.
211. 1964c. Description of a new species of Mendanacoris Miller, with notes on
the systematic position of the genus (Reduviidae, Hemiptera, Insecta).
Amer. Mus. Novitates, 2204: 1-13. R. L. U. and P. Wygodzinsky.
2 ,2. l%5a. General Zoology. McGraw-Hill Book Company, Inc., New York, 4th
ed. 741 p. T. 1. Storer and R. L. U.
213. 1965b. New species of bat bugs of the Cimex pilosellus complex (Hemiptera:
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195
Cimicidae) . Pan-Pac. EntomoL, 41(2): 114—117. R. L. U. and N. Ue-
shima.
214. 1965c. Teacher’s Handbook for Zoology. McGraw-Hill Book Company, Inc.,
New York, revised, iv -f- 60 p. T. I. Storer and R. L. U.
215. 1965d. Book review. Tuxen, The Protura. Bull. Entomol. Soc. Amer., 11(2) :
103.
216. 1965e. A new genus of Neotropical Anthocoridae that resembles the bed bug
(Hemiptera). Pan-Pac. Entomol., 41(3) : 200-204. J. Carayon and R. L. U.
217. 1966a. The biosystematics of the Triatominae. Ann. Rev. Entomol., 11: 309-
330. R. L. U., P. Wygodzinsky, and R. E. Ryckman.
218. 1966b. Insects of the Galapagos Islands. Proc. Calif. Acad. Sci., (4)33(7) :
113-196. E. G. Linsley and R. L. U.
219. 1966c. Programming and communication: A message from the president.
Bull. Entomol. Soc. Amer., 12 (1) : 1-2.
220. 1966d. Obituary. Paul Robert Needham, 1902-1964, p. 82-84, In In Memo-
riam. University of California. A. S. Leopold, E. G. Linsley, and R. L. U.
221. 1966e. Copyright bill: Taxonomic works. Science, 152 (3720) : 291-292.
R. L. U. and W. 1. Follett.
222. 1966f. Distributional patterns of selected western North American insects:
Distributional patterns of Cimicidae in western North America. Bull. En-
tomol. Soc. Amer., 12(2) : 112.
223. 1966g. Book review. Gressitt et al., Insects of Campbell Island. Evolution,
20(2) : 246.
224. 1966h. Monograph of Cimicidae. The Thomas Say Foundation [Entomologi-
cal Society of America], Vol. 7. xi -j- 585 p.
225. 1966i. Evolution of orsilline insect faunas on oceanic islands (Hemiptera,
Lygaeidae). Contribution No. 30, p. 233-235, In R. 1. Bowman, ed.. The
Galapagos: Proceedings of the Galapagos International Scientific Project.
Univ. Calif. Press, Berkeley and Los Angeles. R. L. U. and P. D. Ashlock.
226. 1966 j. The discovery of a possibly aboriginal population of the bed bug
{Ciniex lectularius Linnaeus, 1758). Acta Mus. Moraviae, 51: 237-242.
R. L. U. and D. Povolny.
227. 1967a. Life in Rivers and Streams. McGraw-Hill Book Company, Inc.,
New York, 232 p. [Our Living World of Nature Series].
228. 1967b. Message from the parent organization. Proc. North Central Branch,
Entomol. Soc. Amer., 21: 10.
229. 1967c. The introductory course in entomology. In Teaching entomology
panel. Proc. North Central Branch, Entomol. Soc. Amer., 21:35-36.
230. 1967d. Statement regarding public policy on pesticides prepared by special
committee on public policy, ESA. Bio-science, 17 (2) : 76.
231. 1967e. Entomology looks to the future — Men, money and mission. Bull. En-
tomol. Soc. Amer., 13 (1) : 3-5.
232. 1967f. Sur deux Cimex (Insecta, Heteroptera) noveau pour la faune fran-
gaise, parasites des chauvres-souris. Ann. ParasitoL, 42(2): 269-271. R.
L. U. and J. C. Beaucournu.
233. 1967g. Un hemiptere Cimicidae nouveau d’Afrique centrale. Bull, de I’l.
F.A.N., (A) 29(4): 1688-1694. R. L. U. and J. Carayon.
234. 1967h. Book review. Wygodzinsky, A monograph of the Emesinae (Reduvi-
idae, Hemiptera). Pan-Pac. Entomol., 43(3) : 249-250.
196
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 3
235. 1968a. Book review. Wenzel and Tipton, eds., Ectoparasites of Panama. Pan-
Pac. Entomol., 44(1) : 73-75.
236. 1968b. R. L. Usinger Hemiptera collection. Pan-Pac. Entomol., 44(3) : 257.
237. 1969a. Elements of Zoology, 3rd ed. In press. McGraw-Hill Book Company,
Inc., New York. T. I. Storer, R. L. U., and J. W. Nybakken.
238. In preparation. Biology and relationships of Joppeicidae. N. T. Davis and
R. L. U.
LIST OF NAMES PROPOSED
Listed below in their original forms are the 10 subfamilies, 5 tribes, 94
genera, 9 subgenera, 398 species, 18 subspecies, and 4 varieties (a total of
538 names) proposed by Dr. Usinger. The year of description (century
emitted) with a serial letter indicates the paper in which the description
appears; after the colon is cited the page on which the description is to
1 e found followed by a small T’ when one or more figures were included
■V ith the description. An asterisk (*) preceding the name indicates a
f Dssil form.
SCUTELLERIDAE
Goleotichus (Epicoleotichus)
marianensis, 46i:20
V^anduzeeina
aenescens, 30:131
slevini, 30:132
Pentatomidae
Oechalia
acuta, 41g:82
bryani, 41g:81
ferruginea, 41g:85
similis, 41g:88
sinuata, 42e:217
suehiroae, 41g:89
swezeyi, 41g:90
virescens, 41g:77
COREIDAE
-icptocoris
carnivorus, 46i:25, f
lariversi, 52a: 520, f
diptortus
saileri, 52a:521, f
Lygaeidae
Srachynysius, 42d:44
convexus, 42d:44
Clerada
hirsuta, 42g:249
Cligenes
marianensis, 46i:32, f
Gastrodes
arizonensis, 38e:297, f
conicola, 33d: 127
intermedins, 38e:297
remotus, 38e:296
walleyi, 38e:300
Glyptonysius, 42h:44
laevigatus, 42h:45, f
Metrarga
molokaiensis, 59e:102, f
swezeyi, 59e:103
Neseis (Icteronysius) , 42h:82
maculiceps, 42h:84, f
ochriasis
subsp. baldwini, 45a:405
Neseis (Leionysius) , 42h:52
pallidus, 42h:53, f
Neseis (Physonysius) , 42h:50
ampliatus, 42h;51, f
molokaiensis, 42h:50, f
Neseis (Trachynysius) , 42h:55
alternatus, 42h:76, f
chinai, 42h:78, f
cryptus, 42h:58, f
JULY 1969]
ASHLOCK^ — USINGER BIBLIOGRAPHY
197
fasciatus, 42h:80, f
var. hyalinus, 42h, 81, f
subsp. convergens: 42h:81, f
fulgidus, 42h:59, f
hiloensis
subsp. approximatus, 42h:70, f
subsp. intermedins, 42h:71
subsp. interoculatus, 42h:69, f
subsp. jugatus, 42h:68, f
mauiensis
var. pallidipennis, 42h:80, f
nitidus
subsp. impressicollis, 42h:60, f
subsp. consummatus, 42h:62, f
subsp. contubernalis, 42h:61, f
subsp. pipturi, 42h:65, f
oahuensis, 42h:57, f
swezeyi, 42h:73, f
whitei
subsp. brachypterus, 42h:56
Nesocryptias
adamsoni, 59e:115, f
oahuensis, 59e:lll, f
xiopsis, 59e:112, f
Nysius
abnormis, 42h:112, f
chenopodii, 42h:97
communis, 42h:110, f
frigatensis, 42h:94
fucatus, 42h:90, f
fullawayi, 42h:92, f
var. infuscatus, 42h:94, f
subsp. flavus, 42h:94
kinbergi, 59d:92
kirkaldyi, n.n., 37f:443
mixtus, 421i:110, f
neckerensis, 42h:104
nigriscutellatus, 42h:102, f
nihoae, 42h:103
oceanicus, 37e:441, f
picipes, 37e:439, f
salti, 52e:140, f
suffusus, 42h:96
terrestris, 421i;95, f
Oceanides
bimaculatus, 42h:37, f
bryani, 42h:28, f
delicatus, 42h:25, f
dilatipennis, 42h:43, f
fosbergi, 42h:31, f
incognitus, 42li:35, f
membranaceous, 42h:34, f
myopori, 42h:26, f
parvulus, 42b.: 30, f
perkinsi, 42h:42, f
picturatus, 42h:25, f
planicollis, 42h:28, f
rugosiceps, 42h:37, f
sinuatus, 42h:36, f
ventralis, 42h:24, f
Oreonysius, 52e:141
rugosus, 52e:141, f
Pachybrachius
chinai, 46i:30
*Procymus, 40a: 79
*cockerelli, 40a: 79, f
Rhypodes
chinai, 42d:49
myersi, 42d:47
sericatus, 42d:46
stewartensis, 42d:51
Sinorsillus, 38d:140
piliferus, 38d:140
Aradidae
Acanthaptera, 59a: 107
ceylonensis, 59a: 108, f
incrustata, 59a: 110
Acaraptera ( Acaraptera ) , 59a : 148
myersi, 59a: 149, f
Acaraptera (Lissaptera) , 59a:149
completa, 59a:151, f
Acaraptera (Nesiaptera) , 59a: 148
zimmermani, 59a: 153, f
Acoryphocoris, 59a:294
longicorium, 59a: 295, f
Adenocoris, 59a :67
brachypterus, 59a: 68, f
spiniventris, 59a :70
Aglaocoris
rectangulatus, 59a: 141, f
Aneuraptera, 59a: 96
cimiciformis, 59a:96, f
Aneurus
lobatus, 57h:122, f
Aphelocoris, 59a:299
spinosus, 59a: 300, f
Aphyseteres, 59a: 206
borneensis, 59a: 207
198
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
1 Lpteradus
collaris, 59a: 179, f
longicollis, 59a: 176, f
yiradus
brasiliensis, 40f:639, f
furnissi, 36f:500, f
fuscipennis, 36f:504, f
intermedius, 36f:498, f
linsleyi, 361:493, f
mexicanus, 361:503, 1
serratus, 361:496, 1
Ormoneurum, 59a:316
tuberculatum, 59a:318, 1
^ trtabanellus, 57h : 141
infuscatus, 57li:141, 1
Jlanksiessa, 59a: 349
pubescens, 59a: 350, 1
Barcinus
bhoutanensis, 59a: 281, 1
ceramensis, 59a:278, 1
papuensis, 59a: 279, 1
pro ductus, 59 a: 281, 1
]3ergrothiessa, 59a: 326
Biro ana
inf lata, 59 a: 156, 1
Calisius
acutus, 57h:129, 1
'‘‘balticus, 41i:95
dilaticcps, 46i:37, 1
infuscatus, 57h:133, 1
longicornis, 57h:126, 1
micronesicus, 57h:131, 1
saipanensis, 57h:132, 1
tinianensis, 57h:127, 1
trukensis, 57li:135, 1
Oarventaptera, 59a: 161
spinifera, 59a: 162, 1
Carventini, 50b: 176
^ihelonocoris
bloetei, 54b:271, 1
depressus, 54b: 269, 1
ferrugineus, 54b:273, 1
javensis, 54b:265, 1
kormilevi, 54b: 267, 1
mancinii, 54b:276, 1
milleri, 54b:263, 1
' ^helonoderus, 41Z:179
stylatus, 41/ : 180, 1
' Ihinamyersiinae, 59a : 79
Chinamyersia, n.n., 43e:74
Cbinessa, 59a: 269
Cblouocoris, 59a: 209
multispinosus, 59a: 211, 1
Cremastaptera, 59a : 203
tuberculata, 59a: 204, 1
Dasyaptera, 59a: 168
plumosa, 59a: 170, 1
Daulocoris, 59a: 323
Dihybogaster
wygodzinskyi, 59a: 122, 1
Dimorphacaiitba, 59a: 255
distincta, 59a:256, 1
Dolichothyreus, 59a:290
stigmatus, 59a:291, 1
Drakiessa, 59a:230
Emydocoris, 41/: 176
testudinatus, 41/: 177, 1
Eretmocoris
cubeiisis, 59a: 134, 1
gigas, 59a: 139, 1
longicornis, 59a: 138, f
minimus, 59a: 136, f
productus, 59a: 135, f
prominens, 59a: 136, f
Euchelonocoris
elongatus, 59a:213, f
Euricoris
australicus, 59a: 192, f
hollandicus, 59a: 190, f
piliferus, 59a: 187, f
Glochocoris, 59a: 302
tcrminalis, 59a:303, f
Glyptocoris
spinosus, 59a: 125, f
Hoberlandtiessa, 59a:225
convexa, 59a:226, f
Isodermus
crassicornis, 59a :61, f
tenuicornis, 59a: 59
Kema
bloetei: 59a:316, f
Kolpodaptera, 59a: 144
panamensis, 59a: 146, f
prominens, 59a: 145, f
Kormilevia, 59a: 329, f
setifera, 59a: 331, f
Leuraptera, 59a: 158
zealandica, 59a: 160, f
Libicoris
angulatus, 59a: 184, f
JULY 1969]
ASHLOCK USINGER BIBLIOGRAPHY
199
antennatiis, 59a: 181, f
Lissocoris, 57h:137
spiiiipes, 57h:137, f
Lissonotocoris, 59a: 117
membranaceus, 59a:118, f
Lophocoris, 59a:237
lobatus, 59a: 238, f
Mancinia, 59a:218
incrustata, 59a:219, f
Malkinia, 59a:24{)
stigmata, 59a: 241, f
Mastigocoris, 57li:143
angulatus, 57h:144, f
Mezira
cariiiata, 36f:509
pacifica, 36f:506, f
marianensis, 46i:34, f
iiigripennis, 36f:511
‘“succinica, 41i:98, f
vanduzeei, 361:507, f
Mystilocoris, 59a: 341
pubescens, 59a:342, f
Neadenocoris, 59a :71
abdominalis, 59a: 74, f
acutus, 59a: 76, f
glabrus, 59a :78, f
ovatus, 59a: 75, f
reflexus, 59a :79, f
spiniconiis, 59a :72, f
Neasterocoris, 59a: 333
lutulentus, 59a: 334, f
Neocarventus, 59a: 164
angulatus, 59a: 166, 1
Neophloeobia, 59a:232
australica, 59a: 232, f
montrouzieri, 59a: 234, f
Neuroctenus
abdominalis, 57h:147, f
angulatus, 571i:148, f
lobatus, 57h:154, f
obsoletus, 57h:152, f
pacificus, 46i:36, f
palauensis, 57h:151, f
variegatus, 57h:149, f
Notapictinus, 59a: 361
Notoplocaptera, 59a:171
enigma, 59a: 172, f
Notoplocoris, 41Z:170
montei, 41/: 172, f
Ormenocoris, 59a:371
stylatus, 59a:372, f
Oroessa, 59a: 320, f
Overlaetiella
nitida, 59a: 266, f
Parapictinus
alternatus, 59a: 244
Phanocoris, 59a:223
lobatus, 59a: 223, f
Pictinellus, 59a: 288
Pictinus
acuminatus, 57h:160, f
af finis, 57h:162, f
crassicornis, 571i:164, f
dominicus, 361:512
granulatus, 57h:163, 1
longipennis, 57h:161, 1
minimus, 57h:166, 1
Iiigripennis, 57h:158, 1
productus, 57h:157, 1
rectus, 57h:166, 1
Placogenys, 59a: 352
cockerelli, 59a: 353, 1
Prosympiestinae, 59a :62
Prosympiestus
constrictus, 59a :64, 1
dilatatus, 59a: 65, 1
subparallelus, 59a: 66, 1
Proxius (Neoproxius) , 59a:113
Proxius (Nesoproxius) , 59a:114
minutus, 59a:114, 1
Psectrocoris, 59a:304
dilatus, 59a: 306, 1
Rhysocoris, 59a: 129
rugosus, 59a: 130, 1
Rossius, 59a: 258
Saileriessa, 59a: 346
siligmata, 59a:347, 1
Schoutendeniessa, n.n., 59a:287
Stelgidocoris, 59a: 248
minutus, 59a: 249, 1
Strigocoris, 54d:540
pubescens, 59a: 246, 1
schoutedeni, 54d:540, 1
Tretocoris, 59a: 82
grandis, 59a: 83, 1
Trigonaptera, 57h:139
glabra, 57h:139, 1
Woodwardiessa, 59a: 215
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 3
2(!0
quadrata, 59a: 216, f
2 eugocoris, 59a:310
2 ilmmermania, 48(1:261
brachyptera, 48d:263, f
Enicocephalidae
.enictopechinae, 32 : 149
( leratotrachelus, 45b : 338
Clhinella, 45b: 337
r legenicocephalus, 45b : 328
chinai, 45b: 329, f
riesenicocephalus, 391:268
dybasi, 60b: 222, f
hawaiiensis, 391:268, f
philippinensis, 391:269
Oncylocotls
esakii, 60b: 228, f
gracilis, 60b:226, f
swezeyi, 461:39
1 'seudenicocephalus, 45b : 336
Systelloderus
crassatus, 32:151, f
nitidus, 32:152, f
Reduviidae
1 ^demula
distincta, 461:43, f
gressitti, 60c:263, f
reticulatoides, 60d:262, f
/iradellini, 63c :49
Callotriatoma, 39f:36
cubaiia, 39f:37, f
< lavernicolini, 44k: 32
Carayonla
culiciformis, 52d:212
(.hopardlta
villiersi, 49a: 598, f
Dasycnemini, 63c: 49
Dipetalogaster, 391:40
Elasmocorinae, 44a: 612
Elasmocoris, 44a:613
setigerus, 44a: 6 14, f
Emesopsls
amoenus, 60c:252, f
bellulus, 60c:256, f
decoris, 60c:258, f
habros, 60c: 254, f
scitulus, 60c:248, f
pilosus, 461:42, f
Emplcorls
minutus, 461:45
tesselatoides, n.n., 60c: 267
Gardena
catenarium, 60c: 242, f
Hadrocranella
pallidicoxa, 461:41, f
Mendanacorls
milleri, 64c: 2, f
Mestor
humeralis, 391:38
Neolocoptiris, 63e:51
villiersi, 63c: 52, 1
Nesotriatoma, 44k :38
bruneri, 44k: 39, 1
Pessoala
limai, 42k: 292
Physoderes
minor, 461:50, 1
Plolarla
halosydne, 60c: 235, 1
phyllodoce, 60c: 240, 1
thetis, 60c:239, 1
Polytoxus
distinctus, 60c: 270, 1
grandis, 60c:272, 1
marianensis, 461:48, 1
pilosus, 461:46, 1
Reduvllus
sonoraensis, 42c: 198
vanduzeei, 64b: 11, 1
Saicella, 58c:437
smithi, 58c:440, 1
Themonocorini, 58e:257
Themonocoris, 58e:258
bambesanus, 58e:269, 1
kinkalanus, 58e;264, 1
tshikapanus, 58e:262, 1
Trlatoma
barberi, 391:44
capitata, 41c: 52
chilena, 391:45
incrassata, 391:45
lectlcularlus
subsp. floridana, 44k :63, 1
longipennis, 391:48
longlpes
subsp. nigricollis, 44k: 57
mazzotti, 41c: 54
JULY 1969 ]
ASHLOCK — ^USINGER BIBLIOGRAPHY
201
iiitida, 39f:43
peninsularis, 40c :73
phyllosoma
subsp. intermedia, 44k: 61
picturata, 39f:47
protracta
subsp. woodi, 39f:42
sanguisuga
subsp. occidentalis, 44k: 69, f
subsp. texana, 44k :69, f
Tridemula
contumax, 60c: 260, f
Velitra
micronesica, 60c:278, f
Vescia
brachyptera, 42k: 294
Wygodzinskyella, 52b: 56
Xenocaucus, 49g:43
mancinii, 49g:46, f
CiMICIDAE
Afrocimicinae, 66h:451
Afrocimex
constrictus, 57b :8, f
Aphrania
elongata, 66h:412, f
recta, 57b: 11, f
Aphraniola
orientalis, 53d: 138
Bucimex, 63a: 51
chilensis, 63a: 52, f
Cacodmus
aridus, 57b: 10, f
sinuatus, 66h:397, f
sumatrensis, 57b: 11, f
Caminicimex, 66h:468
Cimex
antemiatus, 65b: 115
brevis, 65b: 117
caveriiicola, 66h:338, f
incrassatus, 65b:115
japonicus, 66h:343, £
latipennis, 65b: 114
Crassicimex, 57b: 6
pilosus, 57b :7, f
sexualis, 57b: 7, f
Latrocimicinae, 66h:458
Leptocimex
duplicatus, 60g:86, f
vespertilionis, 60g:8, f
Loxaspis
malayensis, 66h:427, f
setipes, 57b: 10, f
spinosus, 59h:218
Ornithocoris
pallidus, 59h:219
Paracimex
borneensis, 59h:220
caledoniae, 57b: 9, f
capitatus, 66h:382, f
ignotus, 66h:382, f
lamellatus, 57b: 10, f
philippiiiensis, 59h:221
reductus, 66h:385, f
setosus, 57b :9, f
Passicimex, 67g:1690
imprimis, 67g:1693, f
Primicimicinae^
Propicimex, 66h:305
Psitticimex, 66h:471
Stricticimex, 57g:374
antennatus, 57g:376, f
brevispinosus, 59g:63, f
intermedins, 59b :81, f
namru, 60g:85, f
transversus, 57b: 8, f
POLYCTENIDAE
Hesperoctenes
angustatus, 39a: 18, f
eumops, 39a: 19, f
hermsi, 39a :20, f
limai, 45c: 123
parvulus, 45c: 123
Anthocoridae
Astemmocoris, 65e:200
cimicoides, 65e:203, f
Cardiasthethus
minutissimus, 46i:57, f
Lasiochilus
marianensis, 46i:52
swezeyi, 46i:54
Miridae
Aretas
bifasciatus, 46i:84, f
signatus, 46i:82, f
1 In China and Miller, 1955, Ann. Mag.
Natur. Hist., (12)8:263.
02
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
Campylomma
breviceps, 46i:88, f
brunneicollis, 46i:90, f
hawaiiensis, 43g:287, f
marshallensis, 52a: 523, f
pallida, 46i:89, f
Cyrtopeltus (Engytatus)
lysimachiae, 60f:252, f
phyllostegiae, 60f:251, f
Cyrtorhinus ( Cyrtorhinus )
zwaluwenburgi, 44h:148, f
Cyrtorhynus (Reuteriessa) , 51b :3
vitiensis, 51b: 3, f
Dacerla
alata, 57e:7, f
Deraeocoris
guamensis, 46i:67
Felisacus
crassicornis, 46i:70, f
ochraceus, 46i:69, f
Fulvius
angustatus, 46i:68
Halticus
insularis, 46i:85
Hyalopeplus
guamensis, 46i:58, f
Koanoa
williamsi, 37d:437
Lygus
cruzi, 46i:65
fullawayi, 46i:64
var. rubroscutellatus, 46i:65
guamensis, 46i:63
fdacralonidea
hyalinus, 46i:60, f
Nesodaphne
marianensis, 461:66
□rthotylellus
brunnescens, 461:81
pallescens, 461:80
rufescens, 461:79
Paradacerla, 57e:8
azteca, 57e:10, f
Platylygus
vanduzeei, 31:129, f
Psallops, 461:86
oculatus, 461:87, f
Termatophylldea
maculata, 35c: 271, f
Zanchlus
fragilis, 461:76, f
piperi, 461:77, f
virescens, 461:78, f
Cryptostemmatidae
Ceratocombus (Ceratocombus)
yunguensis, 46d:634
Ceratocombus (Xylonannus)
hawaiiensis, 46d:633
Cryptostemma
pratti, 46a: 240
Nesonannus, 461:91
saileri, 461:91
CiMicoiDEA (No Family)
*Electrocoris, 43b :41
'"brunneus, 43b: 45, f
*pubescens, 43b :45
Saldidae
Saldula
marianarum, 461:100
swezyi, 461:101
Leptopodidae
Erlanotes
buenoi, 42f:103
Veliidae
Halovelia
marianarum, 461:98, f
Ocellovelia, 491:345
Mesoveliidae
Mesovella
pacifica, 461:93, 1
Gerridae
Halobates
hawaiiensis, 38c: 79, f
Limnogonus
lundbladi, 461:95, f
Metrobates
trux
subsp. infuscatus, 53e:179
Hydrometridae
Hydrobatodinae, 49c:318
Hydrobatodes, 49c: 314
schoutedeni, 49c:315, f
JULY 1969]
ASHLOCK USINGER BIBLIOGRAPHY
203
Naucoridae
Ambrysinae, 41b: 9
Ambrysus
barberi, 46c: 189
bohartorum, 46c: 195
buenoi, 46c: 199
caliginosus, 46c: 190
convexus, 46c: 196
fossatus, 46c: 191
fuscus, 46c: 198
hintoni, 46c: 206
hungerfordi, 46c: 192
infuscatus, 46c: 188
lunatus, 46c:203
lundbladi, 46c: 205
sonorensis, 46c: 202
vanduzeei, 46c: 207
variegatus, 46c: 200
woodburyi, 46c: 194
Aphelocheirus
australicus, 37a:341, f
philippensis, 37g:307, f
uichancoi, 37g:305, f
Asthenocoris, 37g:301
luzonensis, 37g:303, f
Cataractocoris, 41b ;11
marginiventris, 41b; 13
Cryphocricos
hungerfordi, 47c: 337, f
latus, 47c: 340, f
mexicanus, 47c: 335, f
obscuratus, 47c: 338, f
Heleocoris
mexicanus, 35a: 113, f
Potamocorinae, 41b :8
Homoptera: Fulgoroidea: Achilidae
^Protepiptera, 39g:66
*kaweckii, 39g:66
204
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
Revision of the chordeumid milliped genus Tynomma
Loomis from California
( Chordeumida : Lysiopetalidea : Lysiopetalidae)
John S. Buckett and Michael R. Gardner
California Department of Agriculture, Sacramento and
University of California, Davis
The genus Tynomma Loomis is composed of small to moderate sized
1} siopetalid millipeds which occur in the forests of the California coast
ranges from the vicinity of San Francisco Bay south to Monterey County.
The lysiopetalids may be recognized in the field by their distinct longi-
tudinal crests on all of the body segments, their extremely rapid move-
ment (approaching that of a centipede) and by the milky -white secretion
g iven off by the repugnatorial pores along the lateral margins of the body
v hen they are disturbed. The genus Tynomma can best be distinguished
from other genera by the configuration of the gonopods; the crest and
leg characters are less valuable because of intergeneric similarities.
Work on Tynomma began in 1910 when R. V. Chamberlin described
Lysiopetalum mutans; he based his description on female specimens
a id described no characters which could later be used to assure proper
g meric placement. Loomis (1937) , in his comprehensive work on North
i^merican lysiopetalids, described the genus Tynomma, included two
new entities, T. consanguineum from the Santa Cruz Mountains and T.
sedecimum from Solano County, California, and tentatively included
Z. mutans in the genus also. Chamberlin (1943) later described
Z ynomma messicanum from northeastern Mexico, far removed geo-
graphically from the other species in the genus. Loomis (1966) con-
sidered T. messicanum to a synonym of Abacion texense (Loomis) . We
agree lhat T. messicanum is not a species of Tynomma, at least, and
hmce will not be treated here. Further, it is our opinion that T. sedeci-
n um and T. consanguineum are synonyms of T. mutans, a widespread
s])ecies.
We have been particularly fortunate in this study in being able to
see type material. Through the courtesy of Dr. Ralph E. Crabill, Jr., we
e; ;amined the holotypes of T. sedecimum and T. consanguineum in the
Lnited States National Museum. Also, topotype males and females of
Z mutans were obtained on the Stanford University campus, enabling
us to confirm the identity of that species.
We wish to extend our appreciation to Dr. Richard L. Hoffman
( ILH) for the loan of material from his private collection, and to Dr.
The Pan-Pacific Entomologist 45: 204^216. July 1969
JULY 1969]
BUCKETT & GARDNER TYNOMMA REVISION
205
Paul H. Arnaud, Jr., for the loan of material from the California
Academy of Sciences (CAS) .
Taxonomic Characters
The lysiopetalids possess an abundance of characters useful in distin-
guishing species and genera. Although a brief account of the characters
used in classification of Tynomma will be presented here, a more thor-
ough treatment of the family can be found in Loomis (1937).
The dorsal crests represent the most valuable somatic characters
because of the relatively strict genetic control over them. Within the
family, numbers of crests on the collum vary from 10 to 20, and are
generally constant at the specific level. In specimens we have examined
of Tynomma, 18 crests are apparent on the collum of T. magnum n. sp.,
and 16 in specimens of T. mutans.
The shapes and relative sizes of the primary and secondary crests
have proven especially useful as generic characters. The primary and
secondary crests are situated alternately, with the primary crests bearing
a caudal seta. In the western genera the secondary crests are distinctly
smaller than the primary crests. Only in some of the species of Tynomma
and Diactis Loomis do the secondary crests reach the caudal margin of
the segment.
The segment of the body at which the transition to the full crest
number occurs differs interspecifically and has previously been accepted
as a specific characteristic with virtually no variation. In the case of
Tynomma, however, we feel that infraspecific variation is present, the
transition occurring on segment 11 or 12 in T. mutans.
The legs of lysiopetalids offer other useful specific and generic
characters. The number of leg segments even varies between genera, with
Colactis Loomis and Heptium Loomis possessing 8 segments in middle
body legs, the other genera with 7 throughout. The tarsi of T. magnum
show a suture on the dorsal surface about mid-length, though it is
definitely not indicative of a full joint. An additional leg characteristic
in which the species of Tynomma show relationship is in the eversible
pouch which occurs on the coxae of the pre-genital legs in the male ; most
other genera apparently lack this character. One character which sepa-
rates the two species of Tynomma is the presence of dimorphism in the
mid-body legs of T. magnum. The coxae of the anterior legs of each
segment are truncate mesally, and with flattened sclerotized anterior
processes. The posterior leg-pairs are unmodified as are all middle-
body legs of T. mutans. Other leg modifications which are not of much
taxonomic use are the hair-combs on the first and second legs of males
?06
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
and first through third legs of females, and the tarsal pads on the mesal
f urfaee of the anterior body legs following the first 2 pairs.
Tynomma Loomis
Tynomma Loomis, 1937, Proc. U. S. Nat. Mus., 84: 117.
Type species: Tynomma sedecimum Loomis (= T. mutans (Chamberlin)).
Diagnosis. — Body size from 19-40 mm, segment number from 42-59; eyes in
triangular patch of 19-38 ocelli; collum with 16 or 18 crests; anterior body
segments with primary crests distinctly larger than secondary crests; post-genital
segments with secondary dorsal crests either thin and reaching caudal margin
or wider and not reaching caudal margin; poriferous carinae higher than dorsal
crests, but subequal to lateral crests; all legs 7-segmented; first and second legs
cf male with hair-comb on mesal margin of tarsus; succeeding pairs of legs with
velutinous pad on mesal surface of tarsus; legs 3-6 and several post-genital pairs
I dth bulbous eversible coxal pouches ; legs of central body segments either un-
riodified or with sclerotized processes on coxae of anterior pair. Anterior pair of
tonopods plate-like, with acute distal apex; narrow elongate posterior gonopods
■\dth simple telopodite, thin curved flange produced from telopodite ends in a
small crescent-shaped process.
Relationships
Relationships within the Lysiopetalidae are difficult to ascertain
1: ecause of the large number of characters involved and the difficulty of
c etermining the relative importance of each. The configuration of the
^ onopods is considered to be valuable because of visible similarities
letween the more closely related genera. The segment at which the
transition to full number of dorsal crests takes place is also considered in
general to reflect relationships. The configuration of crests are of some
i se in correlating relationships, but their function is highly limited
because of the great differences even within genera. Similarly, limited
c ata are available on modifications of walking legs.
With the preceding limitations taken into consideration, Tynomma
sppears to be related to Colactis and Heptium. Although distinctly
different for the most part, similarities between the gonopods of the
t iree genera occur in the erect telopodite which possesses a curved,
bifurcate distal process. Similarly, the point of transition to full crest
number forms a series as follows; segments bb, b2 and b4 in Tynomma,
16 and b7 in Colactis, and b8 or b9 in Heptium. Although the number
cf leg segments in Colactis and Heptium is 8 and in Tynomma 7, the dis-
t net suture in the tarsus of T. magnum prevents this character from
S3riously challenging the relationships of the two genera. The dorsal
c rests of T. magnum resemble Colactis and Heptium in that the secondary
crests do not reach to the posterior margin of the segment. Both species
JULY 1969]
BUCKETT & GARDNER TYNOMMA REVISION
207
of Tynomma, however, differ from the species of the other two genera
in possessing definite secondary crests on the anterior 4 segments.
The two species of Tynomma are obviously not closely related, in
fact several characters lend credence to generic separation. The gono-
pods are so similar in form, however, that generic separation does not
seem practical at the present time.
Key to the Species of Tynomma
Transition to full number of crests occurring on segment 14; body large (40
mm), with distinct flattened process on anterior coxae of mid-body
segments in male magnum, new species
Transition to full number of crests occurring on segments 11 or 12; body
smaller (14-22 mm), lacking coxal modifications of mid-body legs in
male mutans (Chamberlin)
Tynomma magnum Bucket! and Gardner, new species
(Figs. 2, 4, 7, 10)
Diagnosis. — Distinguished from T. mutans by larger body size (40 mm) , more
numerous crests on collum (18), transition to full crest number occurring on
segment 14, greater number of segments (59), processes on anterior coxae of
mid-body segments in male and in details of gonopods (Figs. 2, 7, 10).
Description. — Holotype male . — Head dark brown above, lighter brown frontally,
gnathochilarium faintly light brown; antennae with proximal 5 segments dark
brown, distal 2 segments whitish; tergites medium brown, with whitish median
dorsal stripe, and whitish stripe along poriferous carinae; legs and epiproct also
medium hrown. Head pattern minutely tubercled, with numerous microsetae;
coronal suture short, not visibly extending below vertex; frontal area broad,
sunken mesally; clypeus smooth, lacking microsetae on ventral portion; antennae
long, with segment 3 longest, followed in length by segments 2, 5, 4, 6, 1 and 7 ;
all segments covered semi-densely with short setae; 4 terminal sense cones present
on apex of seventh segment; eyes with following numbers of ocelli in longitudinal
rows, starting from corner nearest antennae: left — 8, 7, 8, 5, 4, 3, 2; right — 7,
7, 8, 6, 5, 3, 2, 1.
Body segments 59; collum narrower than head, possessing 18 crests. Transition
to full crest number occurring on segment 14, with 3 primary suprarepugnatorial
crests present on each side versus 2 on preceding segments. Setae of primary
dorsal crests equalling about one-third length of crest in anterior segments and
projecting mesocaudad; setae increasing in length on caudal 6 body segments, sixth
segment from apex of body with setae equalling half the length of crest; ante-
penultimate and penultimate segment with setae exceeding crests in length;
penultimate segment with reduced crest development.
Middle body segments with single secondary crest present between 2 dorsal-
most primary crests; body tapering gradually caudad, secondary crests becoming
reduced caudally, absent on last 5 segments; lateral margins of epiproct evenly
rounded, exceeding anal valves, with one pair of paramedial dorsal setae and
2 pairs of lateral marginal setae; spinnerets prominent, originating just ventrad
of caudal margin of epiproct and produced caudoventrad ; swollen area beneath
?08
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
JULY 1969] BUCKETT & GARDNER — TYNOMMA REVISION
209
caudal margin of epiproct with one pair of setae subtending spinnerets; anal
valves smooth, with one pair of setae removed from anal lips; anal lips narrow,
slightly protruding; hypoproct lenticular, subdivided into 2 lateral regions and 1
median dark region, seta present at caudal apex of each lateral area and one pair
of setae present on caudal margin of median area (cephalad of lateral setae).
Legs 7-segmented, long, with segments from longest to shortest as follows:
7, 4, 3, 6, 5, 1, 2; first and second legs with row of close hairs on mesal margin
of tarsal segment; basal leg segments with several very large semi-spatulate setae
on mesal margin; other legs without such spine-like setae; legs 3 through 13 with
large velutinous pad on mesal face of tarsal segment; second coxae with a pair
of cylindrical distal anterior projections, also a pair of bulbous eversible glands
present on distal portions caudad of cylindrical processes; legs 3-6 and 8-22 with
similar coxal glands; central body segments with anterior coxae modified as
follows: a broad, flat sclerotized process produced cephalad from ventromesal
margin, caudal end of coxa produced so that coxae broadly in contact; posterior
legpair of each of these segments with coxae rounded, not in contact; legs of
last 17 segments unmodified.
Anterior gonopod equal to width of gonopod socket at base, narrowing gradually
distad to % basal width, then abruptly becoming truncate, with an acute mesal
apex (Fig. 7), mesal margins of the anterior gonopods apparently completely
fused; posterior gonopod extremely long and slender, exceeding anterior gonopods
by almost 2 times again their length; posterior gonopod with coxal region broad,
telopodite about % width of anterior gonopod, produced cephalad from posterior
coxa and curving sharply distad from directly behind anterior gonopod; telopodite
gradually widening subapically, then narrowing to mesally curved acute apex;
translucent flange about 3 times as wide as telopodite produced from meso-caudal
margin of telopodite, curving laterad, and abruptly ending except for small meso-
cephalic process terminating in a dorsal spine and meso-caudal spine (Figs. 7, 10).
Holotype male, 7 miles east of Monterey, Monterey County,
California, 19 November 1962 (J. S. Bucket!) . Paratypes: California:
Monterey Co.: 2 males, 5 females. Big Sur, 22 December 1968 (J.S.B.,
M. R. Gardner and R. C. Gardner) ; 1 female, 4 miles northwest of Big
Sur, 21 December 1968 (J.S.B., M.R.G. and R.C.G.) ; 1 male, 3 females,
1 juvenile, 10 miles southwest of Salinas, 21 December 1968 (J.S.B.,
M.R.G. and R.C.G.).
The holotype will be deposited in the Type Collection of the Depart-
ment of Entomology, University of California, Davis. Paratypes will be
deposited in the U. S. National Museum and the authors’ private
collection.
Discussion. — The discovery of this species has required a broad
redefinition of Tynomma, and has filled a significant gap in the character
spectrum of North American lysiopetalids. The point of transition to full
<-
Fig. 1. Coxal region of middle body leg pair, T. mutans. Fig. 2. Coxal region
of middle body leg pair, T. magnum.
^110
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 45, NO. 3
Fig. 3. Middle body segment, dorsal aspect, T. mutans. Fig. 4. Middle body
segment, dorsal aspect, T. magnum. Fig. 5. Anterior gonopod, cephalic aspect,
T. mutans. Fig. 6. Right cyphopod, lateral aspect, T. mutans.
JULY 1969] BUCKETT & GARDNER TYNOMMA REVISION
211
crest number at segment 14 fits exactly between the 12 of T. mutans and
16 of Colactis species. The weak suture in the tarsal segment bridges
the gap between the 7 and 8-segmented lysiopetalids, and the broader,
shorter secondary dorsal crests differ sharply from the narrow, long
secondary crests of T. mutans.
Tynomma magnum occurs in the northern Santa Lucia Mountains. It
has been collected in a coastal Redwood grove at Big Sur and in an oak
woodland 7 miles inland from Monterey. The known range extends
approximately 22 airline miles in the greatest dimension.
Tynomma mutans (Chamberlin)
(Figs. 1, 3, 5, 6, 8, 9)
Lysiopetalum mutans Chamberlin, 1910, Ann. Entomol. Soc. Amer., 3: 233, pi.
30, figs. 1-10, pi. 31, figs. 1, 2.
Tynomma mutans, Loomis, 1937, Proc. U. S. Nat. Mus., 84: 120.
Tynomma sedecimum Loomis, 1937, Ibid., p. 118, figs. 17 j-1, pi. 3, fig. 6. NEW
SYNONYMY.
Tynomma consanguineum Loomis, 1937, Ibid., p. 119, figs. 17 g-i. NEW SYN-
ONYMY.
Diagnosis. — Differs from T. magnum by smaller body size (14-22 mm), with
fewer body segments (43 to 52), transition to full number of crests occurring on
segments 11 or 12, coxal lobes of middle body segments lacking, and in details
of gonopods (Figs. 8, 9).
Description. — Topotype male. — Head dark brown on vertex, lighter brown on
frons and gulae, gnathochilarium whitish; tergites medium brown dorsally, with
longitudinal light stripes on dorsomeson and along repugnatorial carinae; dorso-
meson and repugnatorial carinae with light longitudinal stripe; legs and epiproct
light brown.
Head reticulate at vertex in semicircular area enclosed by arc curving from
behind both left and right eye clusters; coronal suture ending at anterior margin
of semicircular area. Vertex glabrous, frons with dense short hair; gular area
also with dense short setae; antennal segment 2 longest, followed by 3, 5, 4,
6, 7 and 1 ; all segments covered with fine setae ; segment 7 with four terminal
sense cones; eyes 32 in each patch.
Collum % as wide as head, with transverse row of 6 setae cephalad of midtransect,
a paramedian pair slightly caudad, and a pair on lateral margins; dorsum of
collum with 16 longitudinal crests along posterior third of segment, crests subequal
in size, except third and fifth on each side being reduced, sometimes obsolete on
one or both sides; following segments wider than collum, with increased numbers
of crests; segments 2 through 4 with dorsal setae between or on anterior end of
crest, segment 5 with 6 setae on posterior apices of primary crests and one para-
median anterior pair; segment 5 with 10 dorsal setae on caudal apices of primary
crests; median crests of segments parallel; repugnatorial pores beginning on sixth
segment, situated on third primary crest; beginning with eleventh segment, supra-
repugnatorial primary carinae increased to 3; one pair of secondary carinae
present between median primary carinae; segments caudad of 11 with nearly
THE PAN-PACIFIC ENTOMOLOGIST
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BUCKETT & GARDNER TYNOMMA REVISION
213
obsolete secondary dorsal carinae, ventral secondary carinae prominent; penul-
timate segment with normal number of dorsal setae, though crests greatly
reduced; epiproct rounded from above, with one pair of anterior lateral setae;
one pair of setae on lateral corners of caudal margin; one pair of spinnerets
present, located on ventral side of caudal margin; one pair of setae beneath
spinnerets on ventral bulge of epiproct; anal valves smooth, with one pair of
setae near center of valve; anal lips narrow, slightly produced, widening greatly
ventrad; hypoproct lenticular, with one pair of paramedian setae near anterior
margin and one pair of setae on lateral margins.
Legs 1 and 2 with thick hair tufts along mesal margin of tarsus; leg 2 without
anterior coxal process; legs 2 through 7 with large eversible coxal pouches and
legs of middle body segments with coxae unmodified; leg segment 7 longest,
followed by 4, 3, 6, 5, 1, and 2.
Anterior gonopods subtriangular, broad at base, fused along entire mesal margin,
narrowing evenly distad with slight bulge about halfway, with apex sharply
acute and not exceeding posterior gonopod distally.
Posterior gonopods erect, coxae broad, though short longitudinally, joined
together and to anterior gonopod by sclerotized sternal bar; telopodite joined to
coxae near caudal margin of coxa, produced distad and slightly cephalad, with
apex of anterior gonopods fitting between posterior gonopods; telopodite about
half as wide as coxa, dividing apically into mesal curved flange, with narrow
spicule apically ending in a crescent and continuation of telopodite arching slightly
laterad and rounded apically; seminal canal originating proximocephalad of
posterior gonopods, proceeding up anterior face of posterior gonopod, around
lateral margin to posterior surface just beyond coxa, up posterior face to flange,
where it curves sharply mesad and continues to apex of ventral apex of crescent-
shaped process.
Female . — As in male, but with hair combs on mesal face of tarsus of first 3
legs; cyphopods as in figure 6.
Specimens examined. — CALIFORNIA: Alameda Co.: 5 males, 18 females,
Castro Valley, 27 December 1964 (J. S. Buckett and M. R. Gardner). Marin Co.:
1 male. Mill Valley, 27 May 1952 (H. B. Leach), (CAS); 1 male, 7 females,
Samuel P. Taylor State Park, 25 December 1966 (C. W. O’Brien) ; 1 female, San
Antonio Creek, 4 miles south of Petaluma, 25 December 1%4 (J.S.B.). Napa Co.:
1 male, 4 females, 7 miles west of Oakville, 3 January 1958 (F. Raney and R. 0.
Schuster), (RLH). San Benito Co.: 3 females, between Salinas and San Juan
Bautista, 21 December 1968 (J.S.B. & M.R.G.). San Mateo Co.: 1 female. Lake
Pilarcitos, 17 February 1966 (C.W.O.) ; 2 males, 4 females, Stanford University,
29 December 1964 (J.S.B. & M.R.G.). Santa Clara Co.: 1 female, 6 miles south-
east of Holy City, 29 December 1966 (M.R.G., R.C.G., S. E. Harrison) ; 1 female,
7.5 miles south of Los Gatos, 23 February 1963 (M. Irwin, V. Vesterby, R. West-
cott) ; 1 male, 6 females, Stevens Creek Reservoir, near Cupertino, 28 December
1966 (M.R.G., R.C.G. & S.E.H.). Santa Cruz Co.: 1 male, 1 immature, Ben
Fig. 7. Gonopods, cephalic aspect, holotype of T. magnum (drawn approxi-
mately one-half size in relation to T. mutans) . Fig. 8. Gonopods, lateral aspect,
T. mutans. Fig. 9. Gonopods, caudal aspect, T. mutans. Fig. 10. Gonopods,
lateral aspect, holotype of T. magnum.
214
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 45, NO. 3
Fig. 11. Distribution map of Tynomma.
JULY 1969] BUCKETT & GARDNER TYNOMMA REVISION
215
Lomond, 28 December 1964 (l.S.B. & M.R.G.). Solano Co.'. 2 males, 6 females,
2 miles west of Cordelia, 27 November 1964 (J.S.B. & F. W. Chapman) ; 3 females,
Green Valley, 1 January 1958 (F. Raney & R.O.S.), (RLH). Sonoma Co.'. 4
males, 8 females, 3 miles west of Forestville, 26 November 1964 (J.S.B.) ; 1
female, west of Mark West Reservoir, 22 January 1958 (F.R. & R.O.S.), (RLH) ;
2 females, 4 miles northeast of Penngrove, 26 November 1965 (J.S.B.) ; 1 female,
6 miles south-soutbwest of Santa Rosa, 26 November 1964 (J.S.B.) ; 1 female,
7 miles northeast of Santa Rosa, 26 November 1964 (J.S.B.) ; 15 males, 10
females, 19 miles northeast of Santa Rosa, 27 November 1965 (J.S.B.).
Discussion. — The collection of T. mutans topotypes from Stanford
University as well as examination of the holotypes of T. sedecimum and
T. consanguineum plus much new material have made necessary the
synonymy of the latter two names. T. mutans was described from a
female, the cyphopods of which were drawn by Chamberlin and accom-
pany the description.
Loomis (1937) distinguished T. consanguineum from T. sedecimum
by its having “larger eyes, two more crests on segment 1, and the transi-
tion to the full number of dorsal crests occurring one segment farther
back.” Accepting eye number as a criterion of eye size, as Loomis
apparently did, there is an average difference with more ocelli in
southern populations. However, the topotypes of T. sedecimum north-
east of the San Francisco Bay have as many as 34 ocelli, and specimens
from the Santa Cruz Mountains as few as 28. The difference in crest
numbers on the collum is probably due to variation, since this character
could not be discerned in the specimens we have examined. The
gonopods of the two forms are virtually identical; the 3-pronged process
shown by Loomis projecting from the center of the crescent-shaped
process of T. consanguineum was not observed by us in the type or other
material. The only remaining character which distinguishes the forms
is the location of the segment of transition to full crest number. Speci-
mens from north and east of the Bay and from Stanford have the tran-
sition occurring on segment 11, those from the Santa Cruz Mountains
have it occurring on segment 12. However, in our opinion, this single
feature is not adequate evidence to indicate specific integrity of these
populations, so we interpret this difference as infraspecific geographic
variation.
Tynomma mutans occurs widely in forested areas of the central coast
ranges, both in oak and redwood. Its southern extremity reaches to
within about 15 airline miles of the northern record of T. magnum.
Literature Cited
Chamberlin, R. V. 1910. Diplopoda from the western states. Ann. Entomol. Soc.
Amer., 3(4) : 233-262, pis. 30-43.
:'16
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
1943. On Mexican millipeds. Bull. Univ. Utah, biol. ser. 34(7) : 3-103,
illust.
I.OOMIS, H. F. 1937. Crested millipeds of the family Lysiopetalidae in North
America, with description of new genera and species. Proc. U. S. Nat.
Mus., No. 3006 84 97-135, figs. 16—18, pis. 3, 4.
1966. Millipeds from the region of Monterrey, Mexico. J. Kans. Entomol.
Soc., 39(3): 513-524, figs. 1-10.
New Species of Psenini
(Hymenoptera : Sphecidae)
R. M. Bohart and E. E. Grissell
University of California, Davis
In connection with a generic revision of the Sphecidae certain un-
described species have turned up which either significantly extend the
I nown geographical range or alter generally accepted generic limits.
5 he two new species of the pemphredonine genus Psenulus fall into the
lirst category above. The two new Pseneo fall in the second category.
Holotypes of Psenulus mayorum and P. aztecus, together with Pseneo
irwini are deposited in the University of California at Davis Entomology
1/Iuseum (UCD). The holotype of Pseneo leytensis is in the California
Academy of Sciences (CAS).
Psenulus mayorum Bohart and Grissell, new species
Female holotype. — ^Length 7 mm. Black, marked with pale yellow as follows:
iiandible mostly, palpi, scape in front, pronotum dorsally including lobes except
lor dark humeri, midventral stripe on meso thorax, foretibia externally, midtibia
tiid hindtibia on basal third, basitarsi of fore and midleg mostly; reddish are:
laandible apically, flagellum beneath, scape partly; tarsi partly, and tegula; wings
faintly stained. Pubescence well distributed, most conspicuous on lower face,
mostly silvery but yellowed on vertex and scutum, dense toward midventral line
cf mesothorax; petiole pubescence nearly absent dorsally, sparse but longer than
jetiole diameter laterally; a short hair fringe apically on sternites IV and V.
Punctation mostly fine and sparse, entire body rather polished except propodeum
laterally and posteriorly where it is moderately rough to subreticulate (Fig. 6).
Ulypeus narrowly projecting at apex which is set off by a weak transverse groove
( Fig. 1); mandible dentate near middle of upper inner margin and bidentate
apically; head broader than long, least interocular distance less than eye breadth
ir front view; frontal carina keel-like, very slightly broadened between antennae,
connecting with a transverse carina which angles obtusely and fades out near eyes
(Fig. 1); occipital carina nearly straight dorsally, ending ventrally far before
reaching midventral line; humeral angle sharp but obtuse (Fig. 2) ; notaulices
The Pan-Pacific Entomologist 45: 216-221. July 1969
JULY 1969]
BOHART & GRISSELL NEW PSENINI
217
2
Psenulus mayorum ?
Fig. 1. Head, anterior. Fig. 2. Pronotal ridge, dorsal. Fig. 3. Petiole and
tergite I, profile. Fig. 4. Tergite VI. Fig. 5. Forewing near stigma. Fig. 6.
Propodeum, posterior. Fig. 7. Head, anterior. Fig. 8. Propodeum, posterior.
Fig. 9. Forewing near stigma. Fig. 10. Pronotal ridge, dorsal. Fig. 11. Petiole
and tergite I, profile. Fig. 12. Propodeum, hindcoxa and extreme base of petiole,
lateral.
218
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO.
o
o
Fig. 13. Ocellar triangle, dorsal. Fig. 14. Hindfemur, inner. Fig. 15. Head,
aaterior. Fig. 16. Pronotal ridge, dorsal. Fig. 17. Forewing near stigma. Fig. 18.
I etiole, tergite I and hindleg, profile. Fig. 19. Tergite VI. Fig. 20. Propodeum,
posterior. Fig. 21. Head, anterior. Fig. 22. Hindfemur, inner. Fig. 23. Pronotal
r dge, dorsal. Fig. 24. Forewing near stigma. Fig. 25. Ocellar triangle, dorsal.
IiG. 26. Petiole, tergite I and hindleg, profile. Fig. 27. Propodeum, posterior.
I IG. 28. Tergite VI with enlarged integument surface at right.
JULY 1969]
BOHART & GRISSELL NEW PSENINI
219
nearly complete; propodeal enclosure longitudinally carinate, posterior propodeal
groove linear but broadened above (Fig. 6) ; forewing with first recurrent inter-
stitial and second recurrent received well over on third submarginal cell (Fig. 6).
Petiole nearly cylindrical, strongly decurved near thorax, dorsum as seen in profile
about 1.3 times as long as hindfemur, no lateral or dorsal carinae or sulci except
a tiny depression just before tergite I (Fig. 3) ; pygidium reduced to a median
Carina (Fig. 4).
Male unknown.
Holotype female (UCD), Jalapa, Tabasco, Mexico, November.
Systematics. — This is the first species to be described from the
Neotropical Region. It differs from the four previously described
American species in many respects; the pronotum and legs are ex-
tensively pale, the clypeus is snout-like, flagellomere II is considerably
longer than broad, the humeral angles of the pronotum are rather sharp,
the propodeum is extensively polished dorsally behind the enclosure,
and the pygidium is linear.
Psenulus aztecus Bohart and Grissell, new species
Male holotype. — Length 9 mm. Black, marked with off-white as follows;
mandible mostly, palpi, scape in front, pronotal lobes, forefemur apically, fore-
tibia and midtibia entirely, hindtibia on basal third, tarsi mostly except article V ;
yellow are: flagellum beneath, tegula; reddish-brown are: mandible apically,
flagellum above, abdomen mostly; wings unstained. Pubescence well distributed,
most conspicuous on lower face, silvery throughout; petiole pubescence nearly
absent dorsally, sparse and shorter than petiole diameter laterally, moderately
dense ventrally near base. Punctation mostly fine and sparse, body mostly polished
except propodeum laterally and posteriorly where it is roughly sculptured (Figs.
8, 12). Clypeus emarginate, bidentate; mandible bidentate apically, not dentate on
inner margin; head a little broader than long (Fig. 7), least interocular distance
equal to eye breadth in front view; frontal carina keel-like, not broadened between
antennae, connecting with a transverse carina which angles downward only slightly
and nearly reaches eye margin; flagellum cylindrical, articles and scape subequal
in length (Fig. 7) ; occipital carina gently curved dorsally, ending ventrally far
before reaching midventral line; humeral angle rounded (Fig. 10) ; notaulices
complete but faint posteriorly; propodeal enclosure longitudinally carinate, pos-
terior propodeal groove linear but slightly broadened above (Fig. 8) ; forewing
with first recurrent received near proximal edge of second submarginal cell and
second recurrent received at proximal fourth of third submarginal cell (Fig. 9).
Petiole nearly cylindrical, decurved near thorax (Fig. 11), dorsum as seen in profile
about 1.2 times as long as hindfemur, short carinae present dorsolaterally toward
base, broad shallow depression present just before tergite I.
Female unknown.
Holotype male (UCD), 3 Ml. s. E. Plan de Barrancas, Jalisco,
Mexico, 8 July 1963 (F. D. Parker, L. A. Stange).
Systematics.^ — ^The second species of Psenulus to be described from
the Neotropical Region, it differs from P. mayorum by the dark prono-
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
:\2o
t im, except for the lobes, by the much more extensively pale tibiae and
tarsi, the rounded humeri, the broader third submarginal cell, the more
curved occipital carina dorsally, and by the entirely different clypeal
shape (the sexes of the types are different, however, and this may
account for some of the difference). Psenulus aztecus may be dis-
t inguished from the remaining American Psenulus by the pale markings
and the large polished area on the propodeum behind the enclosure.
Pseneo leytensis Bohart and Grissell, new species
Female holotype. — Length 11 mm. Black, shading to reddish on antenna,
tegula, legs and abdomen laterally and ventrally; forewing moderately stained.
I ubescence light golden, appressed and abundant over most of body, obscuring
sculpture on face as well as pleuron and propodeum (Fig. 20) except part of
enclosure; moderate on notum; hindfemur densely pubescent over entire inner
sirface (Fig. 14) ; petiole hair sparse dorsally, becoming longer and more con-
spicuous posteriorly, sparse and erect laterally; pubescence dense and short on
tergites and second sternite but not obscuring punctation; pygidium with about
s x long bristles on each side (Fig. 19). Punctation fine and close on clypeus,
lecoming less dense toward vertex where it is moderate; notum with medium,
veil spaced punctures, pleuron with well spaced micropunctures; propodeum
posteriorly granulate, enclosure longitudinally striate (Fig. 20) ; petiole with
scattered medium punctures laterodorsally and laterally; tergites I to V with
snail punctures rather evenly spaced about one or two puncture diameters apart;
pygidium smooth except for setigerous pits (Fig. 19). Clypeus beveled and truncate
at apex (Fig. 15) with two longitudinal carinae; frons weakly rounded below
ccelli; a prominent shiny swelling adjacent to compound eye at level of midocellus;
a weak furrow between hindocelli, postocellar line curved into ocellar triangle (Fig.
13) ; inner eye margins rather evenly convex, not converging below; humeri
rounded (Fig. 16) ; notaulices disappearing beyond middle of scutum; petiole
dorsum as seen in profile about 1.8 times as long as hindfemur, petiole with
nearly complete dorsolateral and lateral carinae (Fig. 18) ; pygidium angled at
about 30 degrees, not carinate medially (Fig. 19).
Male unknown.
Holotype female (CAS), Tacloban, Leyte Islands, Philippines,
riovember, 1944 (E. S. Ross).
Systematics. — Pseneo leytensis differs in many respects from the
cnly other Oriental species {P. townesi van Lith) from the island of
l.azon in the Philippines. The clypeal apex is truncate rather than
ernarginate, the punctation of the notum is more sparse and less coarse,
the pubescence is dense and golden rather than silvery to light brownish,
t ie propodeum is posteriorly granulate (beneath the pile) rather than
coarsely reticulate, the second recurrent has a more basad termination
(Fig. 17) and the inner surface of the hindfemur has no special sub-
apical hair tuft (Fig. 14). From American species P. leytensis differs
ty having two longitudinal carinae of the clypeal bevel (as in P.
JULY 1969]
BOHART & GRISSELL NEW PSENINI
221
townesi) , rather than three, by the granulate propodeal cheeks beneath
dense appressed pile, and by the generally pubescent inner surface of
the hindfemur.
Pseneo irwini Bohart and Grissell, new species
Female holotype. — Length 9 mm. Black, tinted with reddish brown on tegula,
tarsi and abdominal segments, especially apically, faint gold and green reflections
on propodeum; forewing lightly stained. Pubescence silvery and dense on face,
pale and sparse but conspicuous on mesopleuron, thicker on notum and propodeum,
fulvous on pronotum and scutum, hindfemur with narrow strip of pubescence on
inner surface for nearly entire length (Fig. 22) , abdomen with off-silvery to
yellowish hair becoming more dense on posterior segments, pygidium with abundant
yellow bristles of moderate length (Fig. 28) . Punctation of clypeus fine and close,
fine and somewhat spaced on frons, sparse on vertex, practically absent on gena;
punctures of scutum mixed fine to coarse, moderately dense, coarse ones somewhat
striatiform ; scutellum with moderate punctation ; mesopleuron to base of propodeum
laterally nearly impunctate; propodeum irregularly reticulate, enclosure as in
fig. 27 ; petiole smooth and polished ; tergite I polished, II to V becoming micro-
punctate laterally and posteriorly; pygidium with dense microreticulation and
scattered large punctures (Fig. 28). Clypeal apex thin, weakly bidentate (Fig. 21) ;
inner orbits converging slightly below; frons evenly rounded below level of ocelli;
a longitudinal furrow between hindocelli, postocellar line curved into ocellar tri-
angle medially (Fig. 25) ; gena with a prominent smooth swelling opposite lower
outer angle of eye; humeral angle slender and projecting strongly outward (Fig. 23) ;
notaulices disappearing beyond middle of scutum; petiole dorsum as seen in
profile about as long as hindfemur (Fig. 26), petiole carinae absent except dorso-
laterally toward base; pygidium angled at about 42 degrees, not carinate medially
(Fig. 28).
Male unknown.
Holotype female (UCD), Quezaltepeque, El Salvador, 6 August
1963 (D. Q. Cavagnaro, M. E. Irwin).
Systematics. — We have seen examples of most of the described
species of Pseneo, and several unique features are displayed by P. irwini.
It is the only species with the clypeal apex thin and bilobed (Fig. 21).
The pronotal humeri project outward but are lobular rather than sharp
as in most species (Fig. 23). The pygidium is nearly flat, extensively
bristly and minutely pebbled (Fig. 28). The second recurrent vein is
only weakly curved anteriorly and is interstitial (Fig. 24) . Finally,
the hindfemur has the inner distal patch of hair prolonged basad (Fig.
22). The species is named for M. E. Irwin who has collected wasps
extensively in El Salvador.
222
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
A New Dictya in California, with Biological Notes
(Diptera : Sciomyzidae)
T. W. Fisher and R. E. Orth^
University of California, Riverside
For reasons of consistency, Dictya fontinalis, new species, is described
ir close accordance with the format used by G. C. Steyskal (1954). In
h:s key, D. fontinalis runs to D. montana Steyskal in the “Typical
Group.” In this sense, D. montana (in California) has been split so that
now we recognize three species of Dictya in California. The third
species, D. texensis Curran, occurs with D. montana in the southern
part of the state.
Dictya fontinalis Fisher and Orth, new species
Holotype Male. — Wing length 5.3 mm. Prosternum bare. Background color
of body rosy-pink, especially apparent in fresh specimens, and usually persisting
or postscutellum of pinned material. Postabdomen as in fig. 1. Surstylus with
dtrsal tip well projecting, not sharply angulate, but blunted or somewhat rounded,
la;eral line not strongly S-shaped. Ventral process of hypandrium without
pieterminal lobe, apex with anteriorly directed tip, subapical portion flattened
w th anterior margin turned outward as viewed ventrally. Ventral processes of
hypandrium parallel in anterior view (fig. 2) , quite narrow in lateral view, appear-
ing much broader when flattened or viewed at an angle so that the maximum
w dth can be perceived (fig. 3A). Ventral process of epandrium with small,
si ort posterior lobe.
For comparison, fig. 3B illustrates the flattened ventral process of the hy-
pe ndrium of D. montana.
Allotype Female. — Wing length 6.3 mm. Prosternum bare. Color as in male.
Postabdominal sternites as in fig. 4A. Sternite VI rectangular, nearly twice as
bioad as long, anterior corners almost right-angled, posterior corners slightly
rcunded. Sternite VII deeply bimarginate anteriorly, anterior margin of median
lobe nearly straight. Posterior margin of sternite VIII shallowly concave without
distinct lobes; apodemes (fig. 4B) subrectangular, apparent thin indistinct con-
necting structure an illusion created in part by infolding and adnation of the
Specialist and Laboratory Technician, respectively.
Fig. 1. Dictya fontinalis Fisher and Orth, Holotype: Postabdomen, dextral
view, inverted; ep, epandrium; hy, hypandrium; ss, surstylus. Fig. 2. Dictya
fontinalis, Holotype: Anterior view of hypandrium inverted. Fig. 3. Terminal
p( rtion of ventral process of hypandrium, flattened: A. Dictya fontinalis, Para-
topotype, 3 August 1965 (T. W. Fisher and R. E. Orth) ; B. Dictya montana
Steyskal, U. S. A., California, Los Angeles Co., Gorman, 3,800 ft., AS-596 (R. E.
0 th). Fig. 4. Dictya fontinalis, Allotype: A. Sternites VI-X; B. Internal view
of sternites VH and VIH to show apodemes.
Tie Pan-Pacific Entomologist 45: 222-228. July 1969
JULY 1969]
FISHER & ORTH A NEW DICTYA
223
4B
4A
224
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
n argins of sternites VII and VIII. Lateral lobes on anterior margin of sternite IX
each with small sublateral tooth directed posterad but slightly elevated from
sl ernite, and from postero ventral view connected by weakly developed narrow shelf.
Holotype Male, Boca Spring, Nevada County, California, 5,900 ft.
elevation, approximately 39° 26' North latitude, 120° 04' West longitude
aid shown as Boca Spring on U.S.G.S. Boca Quadrangle 7.5 minute
series map, 1955, the actual site being near SW corner of NW 14 NE 14
Section 10, T 18 N, R 17 E, Mt. Diablo BM, 7 June 1966, Field Accession
0 . AS-443 (T. W. Fisher and R. E. Orth) .
Allotype. — Shasta Co., Cassel, 5 July 1955 (J. W. MacSwain) ;
approximately 40°54' North, 121 °33' West. Paratopotypes. — 81 S
(average wing length, 5.2 mm), 18 $ (average wing length, 6.2 mm) ;
3 August 1965 (AS-350) , 7 June 1966 (AS-443), 22 September 1966
(A.S-539) (T. W. Fisher and R. E. Orth). See Fig. 5. Other Para-
topes. — Mendocino Co., 1 d , 14 mi. W. Willits, 30 June 1951 (W. C.
Eentinck) , approximately 39°25' North, 123°21' West. 2 2 , Nevada Co.,
I' evada City, Goldflat Road, 2,500 ft. elevation, 13 June 1965 (AS-320)
(P. W. Fisher), approximately 39'°16' North, 121°00' West. 1 d, % mi.
SW. Boca Spring, 5,750 ft., 22 September 1966 (AS-540) (T. W.
Fisher and R. E. Orth).
Other Material Seen. — 1 d , 1 2 (in alcohol) , Trinity Co., Wal-
dorff Ranch (near Big Bar), 3 July 1964, 2,400 ft. (J. C. Borden),
approximately 40.44 North, 122.57 West.
Deposition of Type Material. — Holotype, allotype, to California
T^ cademy of Sciences, CAS Type No. 10207; plus twelve paratopotypes.
Fourteen paratopotypes to U. S. National Museum. Twelve paratopo-
types to Cornell University. Sixty-two paratopotypes and 4 paratypes in
Museum of Department of Entomology, University of California, River-
side.
Distribution. — Although we have collected extensively throughout
California for six years, D. fontinalis has been found only in latitudes
39° and 40° North in mountainous terrain.
The Trinity County specimens and all but one of the specimens that
V e collected were taken at flowing cold springs. Thus, the specific name
fontinalis (Latin — of or from a spring) .
Discussion
We have collected approximately 3,000 specimens of Dictya through-
out California which we now place in D. montana, but a number of
localized forms can be recognized on the basis of gross coloration, size,
a id by differences in certain structures of the male postabdomen. How-
JULY 1969]
FISHER & ORTH— A NEW DICTYA
225
Fig. 5. Dictya fontinalis Fisher and Orth, Paratopotype, male.
ever, at present we consider D. montana to be a polytypic species. This
concept was reinforced by reciprocal crossmating tests, which showed
various degrees of crossing-compatability, between individuals and/ or
small groups of individuals from four populations of montana from
widely separated localities in the state.
Attempts at crossmating D. fontinalis with two forms of D. montana
were unsuccessful. In fact copulation was never observed during these
tests, and no eggs were deposited by females of either species. Interest-
ingly, montana-texensis crossmating tests did result in attempted copu-
lation both ways, and numerous non-fertile eggs were laid by D. montana
females mated by D. texensis males. No eggs resulted from the recip-
rocal cross. No attempts were made to cross D. texensis and D. fonti-
nalis.
Field evidence that D. fontinalis is reproductively isolated from D.
montana is afforded in the general Boca Spring area where D. fonti-
nalis is the most abundant sciomyzid fly at the type locality, a marshy
meadow adjoining the spring proper. A half mile away at a flowing
stream-marshy meadow habitat at a slightly lower elevation, D. montana
is the dominant species of Dictya. From a total of four collections at
226
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
Table 1. Comparison of total numbers of sciomyzid flies collected
a two nearly contiguous sites in Nevada County, California.
Boca Spring
(4 collections)
Stream site
(3 collections)
Antichaeta testacea Melander
0
4
Atrichomelina pubera (Loew)
1
0
Lictya fontinalis F & 0, n. sp.
100
1
Dictya montana Steyskal
3
15
L '.mnia severa Cresson
24
5
Pherbellia humilis (Loew)
1
0
Pherbellia nana (Fallen)
2
3
Pherbellia obscura Ringdahl
0
1
Pherbellia sclioenherri (Fallen)
21
29
Sepedon borealis Steyskal
18
1
S ^pedon capellei Fisher & Orth
1
4
T etanocera nanciae Brimley
21
7
T stanocera soror Melander
2
0
tbe spring and three collections at the stream totals of D. fontinalis and
L . montana were 100:3 at the spring and 1:15 at the stream.
The type locality, Boca Spring, consists of a broad gently sloping
marshy meadow of low grasses and Eleocharis, and is bordered by pines.
^ /ater comes to the surface at the easterly edge of the meadow and flows
in a number of cold, narrow, shallow, clear streamlets in a westerly
direction toward the Little Truckee River, a half-mile distant. In the
past, some of the water was diverted into a small catchment built of
concrete and rocks, now partly in ruins, which early settlers named
“Boca Spring” after the railroad station “Boca” (Spanish — mouth)
h eated near the mouth of Little Truckee River where it enters the
Truckee River. The damming of Little Truckee River created Boca
heservoir. The catchment at Boca Spring is choked with water cress
a id this plant and water celery grow profusely along the water courses
hi the adjacent meadow. By far the most numerous mollusk is a tiny
aquatic prosobranch snail, Lithoglyphus turhiniformis (Tryon) (Fam-
ily: Hydrobiidae) which clings to the submerged leaves and stems of
c ess and celery as well as rocks both in the catchment and in the
n eadow. A fingernail clam is a common benthic mollusk particularly
in the catchment. Intensive searching in the area revealed a few slugs,
ai individual succineid snail, and a few small lynmaeid snails.
The lower stream habitat where Dictya is predominantly represented
hy D. montana is a somewhat drier meadow in a wide gulley choked
V ith grasses and sedges 24 or more inches in length. Scattered pines
JULY 1969]
FISHER & ORTH — A NEW DICTYA
227
provide sparse shade. The stream flowing through it originates at a
nearby spring area in a water course independent from Boca Spring and
forms small (3'' X 3') basins as it flows westerly toward Boca Reservoir,
and it appears to be subject to silting from seasonal runoff in the
gulley that continues for perhaps IV 2 miles above the collecting site.
The course of the stream in the meadow is shaded by tall grasses which
nearly conceal it. In August, water temperature at the stream site (as
well as Boca Spring) was 54° F at 11 A.M. The dominant snail in this
waterway is Physa gyrina Say. We have never found hydrobiid snails
at this site. Competition for Physa with other sciomyzid flies, Pherbellia
schoenherri (Fallen) in particular, probably is reflected in lower total
numbers of D. montana taken. Total numbers of all species of Scio-
myzidae taken at both sites are shown in Table 1. Collecting was
performed with the D-Vac suction collector and conventional aerial-
sweep nets. The collecting dates over a three-year period fell between
7 June and 22 September. The lower (stream) site affords a more
typical habitat, as indicated by its dominant mollusk, the ubiquitous
Physa, with which many sciomyzid flies are associated in California.
Operculate snails are unsuited as host species for most Sciomyzidae
because feeding larvae become pinched as the operculum is pulled into
the aperture of the snail. However, Neff and Berg (1962) concluded
that Littorina littorea (L.), a shoreline species having a coriaceous
operculum, is the normal prey of H oplodictya setosa (Coquillett ) . Also,
Knutson and Berg (1967) state that third-instar larvae of Knutsonia
tritaria (Loew), a European species, eat portions of the soft parts of
certain operculate snails (Hydrobia sp. and Melanopsis algerica (Pils-
bry), a spring-pool inhabitant in southern Spain).
In the laboratory, larvae of Dictya fontinalis rarely could be reared
through the second instar solely on a diet of very small juvenile Physa
virgata Gould (the species available at Riverside, Calif.) and tbe indi-
vidual which was reared through all larval stadia on P. virgata died as
a pupa. Development was completed, however, on a diet solely of
Fontelicella calif or niensis Gregg and Taylor (Hydrobiidae) obtained
from a spring area along San Felipe Creek in northeastern San Diego
County, California. During earlier tests, two other species of Hydro-
biidae, Lithoglyphus turbiniformis (Try on, 1865) from Boca Spring
and Tryonia sp. from a spring along the Westgard Pass road east of
Bigpine, Inyo County, California, were eaten by these larvae. Newly
hatched first-instar larvae of D. fontinalis were able to attack and feed
on the mature hydrobiid snails mentioned [Maximum-sized Fonteli-
cella = 1.2 mm (aperture) X 2.9 mm (length), Lithoglyphus = 1.4 X
228
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
2 5, Tryonia = 1.1 X 3.3]. Although we were unable to provide a com-
plete range in sizes of those snails, it seems reasonable to assume that
newly emerged hydrobiid juveniles, not only being smaller, but having
a less rigid operculum, would be more vulnerable to attack by newly
hatched first-instar larvae of D. fontinalis. Following first-instar de-
velopment on hydrobiid snails, late second- and third-instar larvae
readily attacked small to half-grown Physa, Lymnaea, and Planorbella,
a.i well as hydrobiids.
Acknowledgments
Figures 1-4 were drawn by R. E. Orth and figure 5 was photographed
by^ E. B. White, Department of Entomology, Division of Biological Con-
ti ol, University of California, Riverside, California. Determinations of
snails were by D. W. Taylor, Arizona State University, Tempe, Arizona.
4 he manuscript was reviewed by L. V. Knutson, Systematic Entomology
Laboratory, U. S. Department of Agriculture, U. S. National Museum.
Literature Cited
K NUTSON, L. V., AND C. 0. Berg. 1967. Biology and immature stages of mala-
cophagous Diptera of the genus Knutsonia Verbeke (Sciomyzidae) .
Bull. Inst. Roy. Sci. Natur. Belg., 43(7) ; 1-60.
INeff, S. E., and C. 0. Berg. 1962. Biology and immature stages of Hoplodictya
spinicornis and H. setosa ( Diptera: Sciomyzidae ) . Trans. Amer. Ento-
mol. Soc., 88(2) : 77-93.
Steyskal, G. C. 1954. The American species of the genus Dictya Meigen. Ann.
Entomol. Soc. Amer., 47(3): 511-39.
JULY 1969] HYNES — IMMATURE STAGES OF RHABDOMASTlX
229
The Immature Stages of the Genus Rhabdomastix
(Diptera : Tipulidae)
C. Dennis Hynes
California State Polytechnic College, San Luis Obispo
Since Alexander’s (1920) review of the knowledge of immature stages
of the genus Rhabdomastix, no futher information about the American
species has been published. Descriptions of the immature stages of
Rhabdomastix fascigera Alex., R. trichophora Alex,, R. flaviventris
Alex., and R. setigera Alex., as well as a discussion of the relationships
of R. leonardi Alex., R. hynesi Alex., R. calif or niensis Alex., and R. lipo-
phelps Alex., are contained in this paper.
Rhabdomastix fascigera is widespread in its distribution, ranging
from northern Washington to southern California and in altitude from
sea level to eight thousand feet. The larvae are found in a wide variety
of microhabitats: in streams to depths of four feet, in sand at the sides
of streams, and on the rhizomes of mosses growing at the sides of
streams. The larvae have a characteristic twisting-coiling behavior
when disturbed, usually coiling around small twigs or strands of algae.
Moreover, R. fascigera exudes a sticky substance from the anal opening
which aids in holding it in position in the stream. This phenomenon has
not been observed in any other member of the tribe Eriopterini, but has
been noted in several members of the genus Hexatoma in the Limnophilini.
In the sandy area of their restricted geographical ranges, R. tricho-
phora, R. hynesi, and R. leonardi are also found with R. fascigera. R.
setigera and R. flaviventris occupy habitats in which the sand is finer
and drier than that characteristic of the habitat of R. fascigera.
Descriptions or discussions of the last instar larvae and the pupae
are based on the following numbers of specimens: Rhabdomastix
fascigera (larvae — 20, pupae — 10) , R. subfascigera (larvae — 4, pupae —
2), R. lipophelps (larvae — 2, pupae — 1), R. calif or niensis (larvae — 4,
pupae — 1), R. trichophora (larvae — 20, pupae — 5), R. leonardi (lar-
vae — 10, pupae — 5), R. hynesi (larvae — 6, pupae — 2), R. setigera
(larvae — 20, pupae — 5) , R. flaviventris (larvae — 20, pupae — 10) .
Rhabdomastix (Sacandaga) fascigera Alexander
Last Instar Larva . — Length 16.6-22.9 mm; dextrosinistral and dorsoventral
diameters both 0.6-0.9 mm. Body cylindrical with numerous microscopic yellow
setae, becoming brownish-gold at posterior of eighth segment and anterior of
ninth segment. Pencils of elongate setae reduced. Abdominal segments without
^ This investigation is supported by National Science Foundation Grant GB-4532.
The Pan-Pacific Entomologist 45: 229-237. July 1969
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 45, NO. 3
5
JULY 1969] HYNES IMMATURE STAGES OF RHABDOMASTIX
231
creeping welts. Spiracular disk (fig. 1) obliquely truncate with five fleshy lobes;
markings on face of lobes slightly colored; spiracles dark brown, ringed with
lighter brown on margins. Four slender, white anal gills, subequal in length,
tapering to rounded points at distal ends. Head capsule (fig. 6) typically eriop-
terine, sclerotization generally weak, length 0.784r-0.860 mm (from posterior
margin of maxillary segment to anterior apices of labruni) , width 0.136-0.170 mm
(at posterior mandibular articulation), depth 0.102-0.145 mm, dorsal bars longer
than lateral or ventral bars, curving basally at tips, triangular plates lightly
sclerotized except at margins of premaxillary suture, heavy sclerotization at
caudal margins of suture, lateral bar half thickness of dorsal or ventral bar, labial
portion of ventral bar slightly thiekened, continuing forward as slender rod until
joining submental and premental apparatus, anterior end of bar not expanded or
toothed, maxillae elongate, extending beyond tip of labrum, pointed and curved
slightly laterad distally, mandibles toothed with tips inclined ventral and mesad
from lateral and dorsal articulation.
Pupa . — Length 7.7-9.0 mm; dextrosinistral diameter 1.12-1.20 mm; dorsoventral
diameter 1.10-1.15 mm. Body (fig. 9) pale yellow, wing pads dark brown to black.
Antennal scape with three spines spaced equally along ventral edge medially,
frontally, and laterally, with spine located immediately below lateral spine and
in line with spine located at cephaloventral edge of eye sheath, another series of
spines located in facial margins in groups of four on both sides of body. Pronotum
with medial tubercle appressed closely over flagellar portion of antennal sheaths,
lateral tubercle with three or four elongated setae; mesonotum with lateral carina
ending in small slit indicating tuberculate mesonotal breathing horns; medial
carina raised, sharp, ending at pseudosuture. From pseudosuture a thin brownish
line divided by white line extends over dorsal crest, dorsal crest brown except in
areas next to patch of heavy spines. Spiracular openings simple, not projecting
laterally. Leg sheaths end midlength of fourth abdominal segment, prothoracic
and mesothoracic leg sheaths subequal in length, metathoracic sheaths longer.
Male cauda with spiracular area depressed, rising abruptly to four large spines
forming transverse row. Sheaths of inner dististyles bulbous at bases, narrowing
abruptly, curving dorsally, and becoming pointed at tips, sheaths of outer disti-
styles flat, joined along entire length, curving mesally at tips. Female cauda with
only remains of spiracular openings as disk markings. Cauda with four large
spines, each with single elongate seta. Anteapical spines of tergal valve directed
sharply dorsad; sternal valves with distal tips remaining in contact with bulbous
tergal valves along their entire length.
Rhabdomastix (Sacandaga) trichophora Alexander
Last Instar Larva. — Length 15.0-16.8 mm; dextrosinistral and dorsoventral
diameters both 0.55-0.60 mm. Body description similar to Rhabdomastix fascigera
Fig. 1. Three-quarter view of spiracular disk of R. fascigera. Fig. 2. Three-
quarter view of spiracular disk of R. trichophora. Fig. 3. Three-quarter view
of spiracular disk of R. calif or niensis. Fig. 4. Three-quarter view of spiracular
disk of R. flaviventris. Fig. 5. Three-quarter view of spiracular disk of R.
setigera.
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 3
32
JULY 1969] HYNES IMMATURE STAGES OF RHABDOMASTIX
233
with following exceptions. Fine yellow, closely appressed, microscopic setae not
as numerous as in R. fascigera. Pencils of setae evident. Head capsule similar
to that of R. fascigera; length 0.582-0.607 mm, dextrosinistral diameter 0.101 mm,
dorsoventral diameter 0.250-0.304 mm. Posterior spatulate plates bent slightly
laterad, triangular plate merely touching inner edge of spatulate area. Spiracular
disk (fig. 2) with five fleshy lobes; dorsal lobe ending in thick, horny tubercle
projected caudally, narrowing to sharp, ventrally directed point; dorsolateral
lobes with large horny spine near tip of lobe directed mesad, lower edge with
thin, dark brown stripe, upper edge with barely discernible marking, triangular
area above spiracles dark brown, center of spiracles yellow with rims of spiracles
same color; anal lobes fleshy and white, larger and pointed at lateral margins.
Papa . — Length 6.5-8.5 mm; dorsoventral and dextrosinistral diameters both
0.8-0.9 mm. General description similar to R. fascigera except for following
details. Heavily sclerotized, transverse mesonotal ridge followed by a series of
smaller ridges less heavily sclerotized behind spines of dorsal crest (fig. 10).
Few or no spines forming posterior transverse line on dorsum of abdominal
segments four, five, and six. Male and female cauda similar to that of R. fascigera
except for small spines in area homologous to spiracular disk of larva.
Rhabdomastix (Sacandaga) californiensis Alexander
Last Instar Larva. — ’Length 12.0-16.1 mm; dextrosinistral and dorsoventral
diameters both 0.45-0.55 mm. Larval and pupal characteristics essentially similar
to those of Rhabdomastix trichophora except for following details. Dorsolateral
lobe of spiracular disk with ventral and dorsal stripe of dark brown, center of
spiracle yellow, ventral circumference dark brown, remainder of circumference
yellow or same color as center of spiracle (fig. 3).
Papa. — Length 5.48-5.70 mm; dextrosinistral diameter 0.66-0.75 mm; dorso-
lateral diameter 0.70-0.73 mm. Series of smaller ridges behind mesonotal ridge
lacking.
Rhabdomastix (Sacandaga) leonardi Alexander
Last Instar Larva. — Length 12.5-16.6 mm; dextrosinistral and dorsoventral
diameters both 0.45-0.55 mm.
Papa. — Length 8.0 mm; dextrosinistral and dorsolateral diameters both 1.0 mm.
Remainder of larval and pupal characteristics similar to those of R. californiensis.
Rhabdomastix (Sacandaga) hynesi Alexander
Last Instar Larva . — Length 14.6-18.5 mm; dextrosinistral and dorsoventral
diameters both 0.55-0.60 mm.
<-
Fig. 6. Head capsule of R. fascigera; D dorsal view, V = ventral view. Fig. 7.
Ventral (maxillary) bar and maxillary plate of R. setigera. Fig. 8. Ventral (maxil-
lary) bar and maxillary plate of R. flaviventris. Fig. 9. Lateral views of anterior and
caudal ends of pupa of R. fascigera. Fig. 10. Lateral views of anterior and
caudal ends of pupa of R. trichophora. Fig. 11. Lateral views of anterior and
caudal ends of pupa of R. setigera.
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
i:34
Pupa . — Length 8. 1-9.6 mm; dextrosinistral and dorsoventral diameters both
C. 9-1.0 mm. Remainder of larval and pupal characteristics similar to those of
11. californiensis.
Rhabdomastix (Sacandaga) setigera Alexander
Last Instar Larva . — Length 11.6-14.0 mm; dextrosinistral and dorsolateral diam-
eters both 0.6-0.7 mm. Body whitish yellow, cylindrical, tapered at anterior end;
tioracic segments slightly expanded at eighth abdominal segment. Spiracular
cisk (fig. 5) flat, tapering gently from face obliquely to tips of ventral spines;
( orsal and dorsolateral lobes not discernible. Markings on spiracular disk
forming inverted Y with stem between spiracles, marking in upper center of disk
ct base of Y nearly diamond shaped, continuing basad, indenting, medially, then
expanding and continuing in separate arms to base of spines, spiracles small,
1 eddish-brown in centers with outer rims lighter, ventral lobes end in thick
spines curving basally; four bulbous, white anal lobes, posterior lobes approx-
imately twice length of anterior pair. Head capsule different from that of
Rhabdomastix fascigera-, length 0.633 mm.; dextrosinistral diameter 0.136-0.145
rim; dorsoventral diameter 0.171-0.195 mm; antennal buttress continuing for-
1 rard and medially into heavily sclerotized anteclypeal fold, antennae four to
live times as long as wide, lightly sclerotized; papillae elongate, ending approxi-
mately one-quarter the distance from tip of maxillae; maxillae expand to tip
cf labrum, blunt and glove-shaped; mandible heavily sclerotized and angled
I asally and mesally ; hypopharynx sclerotized, anchored to inner margins of
maxillary plate; maxillary plate expanded at anterior tip of maxillary bar, ending
ia three or four small blunt teeth anteriorly (fig. 7), separated centrally by small
tbin membranous area.
Pupa . — Length 6.3-7.6 mm; dextrosinistral diameter and dorsoventral diam-
eters both 1. 0-1.1 mm. Frontoclypeal area, eye and antennal sheaths with spines,
iintennal sheath crenate on basal margin. Anterior margin of pronotum with small
lobe on either side of median carina; posterior to these small lobes are two
more bulbous lobes each with two setae directed dorsally; beneath these is a
large, ridged lobe at lateral margin of, and overhanging, antennal sheath;
1 reathing horns small, bulbous; median carina with small transverse ridges on
either side extending from anterior margin of mesonotum to pseudosuture, median
line forming slight V-shaped dip up to and beyond dorsal crest at pseudosuture.
Patch of spines (fig. 11) covering dorsal crest, medial spines larger, becoming
smaller laterally; area posterior to dorsal crest slightly ridged or variegated to
I osterior margin of mesonotum. Base of haltere sheath visible as slight ex-
1 ansion near dorsal margin of wing pad. Leg sheaths end approximately midlength
cf abdominal segment four. Male cauda with gonosheath thick, bulbous at base
V ith spine laterally directed at tip of sheath. Inner dististyle with base thick,
then abruptly thinner, ending in spine at tip; sheath bulbous at base, narrowing
to spines at tip. Small tubercle directed caudally between bases of tergal arms,
transverse row of small spines laterally at base of each tergal arm. Female cauda
with tergal arms similar to those of male, tergal valve sheath with ridges on dorsal
surface to dorsally directed spine, three or four setae found caudally at base of
spine. Ventral side of sternal sheath ridged, curving dorsally to unarmed tip.
1 abdominal segments two through seven with two to three folds except for
cbdominal segment eight with spines on lateral edges of sternum but not on
r redial sternal area or on pleural area.
JULY 1969] HYNES IMMATURE STAGES OF RHABDOMASTIX
235
Rhabdomastix (Sacandaga) flaviventris Alexander
Last Instar Larva . — ^Details same as given for Rhabdomastix setigera with
exceptions as follow. Head capsule with maxillary plate (fig. 8) having anterior
teeth much more pronounced than in Rhabdomastix setigera. Spiracular disk
(fig. 4) with inverted Y much lighter brown than in R. setigera, stem of Y
divided down medial line by whitish-yellow stripe, base of Y much thicker with
lateral margin nearly straight line to base of ventral lobe spine, inner margins of
arms of Y slanting from center of disk to base of spines.
Pupa . — The pupa of R. flaviventris cannot be separated from that of R. setigera
on basis of specimens in my possession.
Discussion
A basic problem arises in trying to explain the relationships of these
species which show two very different types of larval forms, but which
differ only slightly in the adult stages. The species under discussion
may be divided into three groups based upon the morphological char-
acters of the immature stages given above. The first of the three groups
will be called the fascigera group. I have reared specimens of R. sub-
fascigera and R. lipophleps, and found them to be similar in external
morphology to R. fascigera. Many characteristics were investigated,
but the materials in my possession do not allow separation of larval or
pupal specimens of these species with any degree of confidence. How-
ever, separation by their known geographical ranges is possible with
the exception of those found in the northern Washington area. In this
area, I have reared R. fascigera at 2000 feet, and from the same stream
have also reared R. suhfascigera at 5000 feet.
The second group will be referred to as the thichophora group, con-
sisting of R. trichophora, R. calif or niensis, R. leonardi, and R. hynesi.
These specimens are similar to each other and to the fascigera group in
body form and morphology of the head capsule. They differ from the
fascigera group in details of the spiracular disks. There is no trouble
in separating larvae of Rhabdomastix trichophora from the other species ;
however, I cannot separate larval specimens of R. calif orniensis, R.
leonardi, and R. hynesi. Again, separation of these species can be made
on the basis of geographical location. I have not been able to separate
pupal stages of any species in this group on the basis of morphology,
but can easily separate them as a group from the fascigera and lurida
groups.
The third group has been referred to as the lurida group (Alexan-
der, 1943) of which R. setigera and R. flaviventris have now been
reared. The larvae are different from those of the other groups in body
236
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
form, especially in the ratio of length to width, in morphology of the
spiracular disk, and in many details of the head capsule, especially the
toothed maxillary plate at the anterior end of the ventral bar.
Investigations to determine whether or not members of each of the
groups are conspecific on the basis of adult characteristics were con-
sidered beyond the scope of the present project. Based on morphologi-
cil characters found in the head capsule and in the spiracular disk, the
j'lscigera and trichophora groups are more closely related to each other
t ran to the lurida group. Pupal and, to a greater extent, adult characters
place the lurida group in the genus Rhahdomastix. These pupal char-
acteristics of the lurida group are the shape and structure of the frontal
and cephalic crest areas. The characteristic adult features involve wing
venation and the genital structures.
There are two possible explanations of the differentiation seen in
t lese groups. The first explanation is that the larval stages of these
groups have progressed in their evolution to such an extent that they
Slow a greater magnitude of divergence than do the adult stages. This
ejcplanation would, moreover, indicate that the species belonging to the
lurida group are correctly considered to be a part of the genus Rhahdo-
mastix. The basis for the generic definition of Rhahdomastix is the
norphology of the adult stage. If we use characteristics of this stage
alone as our criteria for the genus, there are two morphologically dis-
t act types of larvae in one genus — a rare situation in the family
Tipulidae and inconsistent with general concepts of the generic category
i 1 cranefly families.
The seeond possible explanation is that the larval stages of
t le lurida group have retained their original lines of differentiation
and have continued their own development along divergent lines.
T he adult stages have converged with those of the fascigera and
t ichophora groups on the basis of comparisons of wing venation and
genital structure. Alexander (1920) considered the genus Rhahdo-
mastix to be closely allied to the genus Gonomyia. The lurida group,
\'ith characteristies of the larval head capsule such as the toothed
riaxillary plate, details of the frontoclypeal area, shape of the head
c apsule, as well as the absence or near absence of lobes on the spiracular
cisk, may be more closely related to the genus Teucholahis in which
t iese characteristics are also present.
The evidence also suggests that members of the lurida group are not
correctly classified to the generic level and that, with further study, a
rew genus may have to be proposed to indicate correctly the relation-
strip between the genus Rhahdomastix and the genus Teucholahis,
JULY 1969]
SCIENTIFIC NOTES
237
Literature Cited
Alexander, C. P. 1920. The crane flies of New York. Pt. 11. Biology and
phylogeny. N. Y. Agr. Exp. Sta. Ithaca Mem., 38: 699-1133, 87 pis.
1943. Records and descriptions of North American craneflies (Diptera).
Pt. III. Tipuloidea of the Upper Gunnison Valley, Colorado. Amer.
Midland Natur., 29: 147-179, 38 fig.
SCIENTIFIC NOTES
The location of the holotypes of Bruchus cubiculus Casey and Mylabris
wheelocki Blatchley (Coleoptera: Bruchidae). — In an earlier paper I dis-
cussed the problems involved in locating type-specimens of Bruchidae deposited
in museums in the United States, listed where types were located and designated
some lectotypes (Johnson, 1968, Ann. Entomol. Soc. Amer., 61: 1266-1272). The
present note lists pertinent information concerning the holotypes of two species not
included in the above paper, Mylabris ivheelocki Blatchley (Blatchley, 1930, Can.
Entomol., 62: 35) and Bruchus cubiculus Casey (Casey, 1884, Contributions to the
Descriptive and Systematic Coleopterology of North America. Part II. p. 183).
The holotype of M. wheelocki is deposited in the Entomology Research Collec-
tion, Purdue University, Lafayette, Indiana. Through the kindness of Ross H.
Arnett, Jr. and Eric H. Smith I was able to examine the specimen and found it
bears the following labels: “3837,” “TYPE,” “Royal Palm Park, Fla. WSB,
3-21-29,” ‘‘'Mylabris wheelocki sp. nov.” According to Dr. Smith (and Blatch-
leyana, 1930, p. 59) this is Blatchley Type •#374.
The holotype of B. cubiculus is deposited in the United States National Museum.
Dr. John M. Kingsolver relayed to me that the specimen bears an “Ari.” label
and bears type number 49229.
L. J. Bottimer (1968, Can. Entomol., 100: 1034, 1035) included both species
in the genus Megacerus and discussed the taxonomic status of Megacerus
cubiculus. — Clarence D. Johnson, Northern Arizona University, Flagstaff.
238
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
Occurrence of Exoteleia burkei in the Sierra Nevada (Lepidoptera :
G elechiidae). — During June 1965, dying shoots infested with small lepidopterous
la fvae were observed on knobcone pines, Pinus attenuata Lemm., growing at the
Ir stitute of Forest Genetics, Placerville, El Dorado County, California.
Adults reared from the shoots were identified by J. A. Powell, University of
C ilifornia, Berkeley, as Exoteleia burkei Keifer, which had been reported only
from the vicinity of the San Francisco Bay area, and from Monterey pine, P. radiata
D Don, and Digger pine, P. sabiniana Dough (Keifer, 1931, Pan-Pac. Entomoh, 8:
61-73; Burke, 1937, Proc. West. Shade Tree Conf., (4) : 21-31; Burdick and
Powell, 1960. Can. Entomoh, 42(4): 310-320). The previous easternmost record
for E. burkei was from the Coast Range in eastern Alameda County (Burdick and
Powell, 1960). PowelU has since reared E. burkei from P. coulteri D. Don from
tie La Panza Range, (part of the Coast Range) San Luis Obispo County, and
Brown and Eads (1967, Calif. Agr. Exp. Sta. Bull. 843, 72 pp.) report that it
commonly infests P. radiata in Southern California. The Sierra Nevada observation
represents a range extension of about 100 miles eastward, but whether the species is
ir fact new there or merely previously unnoticed is not known. Neither is it known
if E. burkei is more widely distributed east of the Central Valley.
Knobcone pine is native to the Placerville area; however the infestation that I
oljserved was in a small plantation used for tree breeding work. The infested trees
were from 20-30 feet tall. Ponderosa pine, P. ponderosa Laws., is the predominant
ni tive pine species at Placerville; sugar pine, P. lambertiana Dough and Digger
pine also occur naturally. Indeed most species of Pinus are represented in the In-
stitute’s plantations, but only P. attenuata was observed to be infested.
Previous authors have reported a number of larval habits for E. burkei, including
needle mining, feeding in staminate cones, and boring in the developing shoots. At
Placerville only the last habit was noted, but the early instar larvae were not seen.
It appeared that a single shoot was mined by each late-instar larva. Generally, only
or e larva occupied a shoot, but there were rare instances of multiple infestation.
Tie shoot was thoroughly mined out for a distance of 3.5-5.0 cm back from the
tip. Needle bases were also mined out from within the shoot, and the needle bundles
were thus cut off within the fascicle. Needle elongation was less than 2.0 cm when
gi owth was halted as a result of larval activity.
Contrary to the assumption of Burdick and Powell (1960) that larvae leave the
feeding site to pupate, pupation in this instance took place within the mined shoots.
Tie pupa was most often found head outward in a mined-out base of a needle
bundle but occasionally one was found within the shoot itself. Before pupating, the
la va cut a hole in the needle sheath from within to provide for egress of the adult,
ar d also built a silk ramp inside to direct the adult out the hole. — Robert E.
Stevens, Rocky Mountain Forest and Range Experiment Station, Fort Collins.^
Personal communication, October 17, 1968.
"Central headquarters maintained at Fort, Collins, in cooperation with Colorado State University. At
the time these observations were made, the author was on the staff of the Pacific Southwest Forest and
Range Experiment Station, Berkeley, California.
JULY 1969 ]
SCIENTIFIC NOTES
239
First recorded occurrence of the genus C allant hidium in Idaho with
notes on three nests of C. formosum (Cresson) (Hymenoptera ; Mega-
chilidae). — The genus Callanthidium contains only the two species, C. formosum
(Cresson) and C. illustre (Cresson), which are limited in distribution to the
temperate regions of western North America. Grigarick and Stange (1968, Bull.
Calif. Ins. Surv. 9: 1-113) present a key to these species, give their distributions,
and review the biology of C. illustre. Until now, nothing has been reported on
the nest construction of C. formosum.
An examination of trap nests placed in Idaho in 1967 revealed three nests of
C. formosum which were collected 3 miles northwest of Carey, Blaine County,
Idaho. Members of this species have also been collected at Craters of the Moon
National Monument, Butte County, Idaho, on flowers of Lupinus caudatus Kellogg
and Phacelia hastata var. leucophylla (Torr.) Cronq. and in a Malaise trap.
These constitute the first published records of Callanthidium in Idaho.
The nests were established in a nest block which had been placed in a bush of
Artemesia tridentata Nutt. The block had a series of approximately 80 regularly
spaced holes, 5-9 mm in diameter and 10-15 cm long; three holes had been used,
all by C. formosum. Nest I was approximately 65 mm long and 8 mm in diameter.
There were two cells, 20 and 22 mm long, separated by a 4^5 mm long tomentum
plug. A similar plug was also at the bottom of the nest. Each cell contained a
mammalate cocoon 13 mm long. Tomentum tightly enclosed each cocoon. The
top of the upper cell was sealed with a resin cap 0.8 mm in thickness followed by
a 20 mm long tomentum plug. There were no pebbles or debris within this nest.
Nest II was 9 mm in diameter and 107 mm long. There were two cells, 22 and 23
mm long, with the tomentum similar to nest I. The top of the upper cell was
sealed with a plug of pebbles 8 mm in length intermixed with some resin, followed
by a 8 mm long tomentum plug. Nest III was 5 mm in diameter and 83 mm long.
There were two cells, each 20 mm long and again with tomentum between the
cells and at the bottom of the nest. The upper cell was without a resin cap or
tomentum plug, but had 8 mm of pebbles with no resin, suggesting that this nest
was either incomplete or had been disturbed.
The nest size, materials used, and number of cells are much the same as was
described by Hicks (1929, Can. EntomoL, 61(1): 1-8) for C. illustre. Grigarick
and Stange (1968) state in their generic key that members of Callanthidium use
resin in the construction of the nest cap. This note verifies the usefulness of
this part of the Callanthidium couplet.
I wish to thank W. F. Barr, University of Idaho, Moscow, for setting out and
collecting the nest blocks used in this study. — Donald S. Horning, Jr., University
of California, Davis.
240
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
ZOOLOGICAL NOMENCLATURE: Announcement A. (n.s.) 83
Required six-month’s notice is given on the possible use of plenary powers by the
International Commission on Zoological Nomenclature in connection with the fol-
io ving names listed by case number:
(see, Bull. Zool. Nomencl. 25, pt. 6, 28 February 1969)
1859. Validation of emendation to patchae of patchiae (Schizoneura) Bdrner &
Blunck, 1916; suppression of Schizoneura patchi Meunier, 1917 (Insecta,
Hemiptera) .
1854. Suppression of Dicyphus tamaricis Puton, 1886 (Insecta, Hemiptera).
1858. Type specimen for Anthocoris pini Barensprung, 1858 (Insecta, Hemiptera).
Comments should be sent in duplicate, citing case number, to the Secretary, In-
te national Commission on Zoological Nomenclature, c/o British Museum (Natural
H story), Cromwell Road, London S. W. 7, England. Those received early enough
will be published in the Bulletin of Zoological Nomenclature. — W. E. China, Act-
ing Secretary to the International Commissioner on Zoological Nomenclature.
ZOOLOGICAL NOMENCLATURE; Announcement A. (n.s.) 84
Required six-month’s notice is given on the possible use of plenary powers by the
Ir ternational Commission on Zoological Nomenclature in connection with the
following names listed by case number:
(see. Bull. Zool. Nomencl. 26, pt. 1, 12 May 1969)
1848. Suppression of Papilio sehrus Huebner, 1824/26 (Insecta, Lepidoptera) .
1869. Type-species for Physothrips Karny, 1912 (Insecta, Thysanoptera) .
1871. Type-species for Xyletinus Latreille, 1809 (Insecta, Coleoptera) .
Comments should be sent in duplicate, citing case number, to the Secretary,
Ir ternational Commission on Zoological Nomenclature, c/o British Museum
(Natural History), Cromwell Road, London, S.W.7, England. Those received early
er ough will be published in the Bulletin of Zoological Nomenclature. — ^W. E.
C IINA, Acting Secretary to the International Commission on Zoological Nomen-
clature.
JULY 1969]
BOOK REVIEWS
241
BOOK REVIEWS
Ecology and Biogeography of High Altitude Insects. By M. S. Mani. 528
pages, 80 figs., 41 tables, cloth. 1967 (W. Junk N. V., The Hague). ($27.80,
U.S.)
In 1962 Professor Mani published his “Introduction to High Altitude Entomology”,
which was primarily concerned with the insect fauna of the high Himalaya. During
the intervening years Dr. Mani has maintained his interest in alpine environments
and has continued to expand the scope of his research. In this newer and much
larger volume he includes hundreds of personal observations made during field trips
into the Caucasus, the Alai-Pamirs, the Ala-Tau, and the Tien Shan mountains. He
has also incorporated pertinent information resulting from the field research of
hundreds of other entomologists and ecologists throughout the world, as evidenced
by text references to each of the 1141 books and articles cited in the bibliography.
Dr. Mani devotes several pages to a general description of typical mountain
terrain and the difficulties associated with existence above timberline. Nine
chapters (260 pages) are devoted to specific mountainous regions throughout the
world (“The Mountains of Equatorial East Africa”; “The Himalaya”; “The Moun-
tains of the North Temperate Regions of the New World”; “The Subarctic and
Arctic Mountains” and so forth) . These chapters include not only the descriptions
of the ranges and peaks but also analyze the significant ecological factors involved.
Precipitation, atmospheric movements, temperature means and extremes, and other
climatic aspects are documented. The altitudinal biotic zonation is discussed in
some detail, with an enumeration of typical genera and speeies of plants that
characterize each zone. Separate sections are concerned with the insect life of each
zone, and previously-published data concerning the fauna of each mountainous area
are repeated here, including the significant ecological and biological observations
of numerous authors. These zoogeographic discussions alone will make this book a
very valuable addition to the libraries of all scientific institutions, and the remain-
ing chapters include such an abundance of biological information that every serious
animal and plant ecologist will want a copy in his own personal library. Chapter
five is a somewhat taxonomic discussion of the high altitude arthropod fauna, how-
ever no attempt is made to describe the insects or to aid the reader in identification.
Each Order of insects that contributes to the high altitude fauna is covered briefly,
and within the larger orders several families are more specifically discussed. In
most cases details of altitudinal and zoogeographic distribution are presented and
the biological and ecological aspects of the major autochthonous hypsobiont ar-
thropods are discussed.
Probably the most fascinating and stimulating portion of the book for most
readers is the detailed analysis of the physical factors that are so significant at
high altitudes — reduced atmospheric pressure, temperature, and humidity, and the
great increase in wind velocity, solar insolation, and ultraviolet radiation. Emphasis
is placed on the importance of snow cover, rock mantle, soil moisture, and food
availability. The behavioral and morphological adaptations that enable certain
insects to survive in the alpine environment are considered in great detail. Im-
portant morphological adaptations include melanism, reduced body size, and
atrophy of wings, while such traits as terricoly, hygrophily and cryophily are of
major significance. Ingenious diagrams visually indicate many of the complex inter-
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 45, NO. 3
2^:.2
relationships between various environniental, behavioral, and morphological factors
that are associated with insect survival at high altitudes. Dozens of charts and
graphs help to clarify the correlation between altitude, latitude, temperature, humid-
ity, edaphic factors, and so forth. For example, one table compares air temperatures
with readings at and below the soil surface at ten different high altitudes. Others
correlate atmospheric pressure with oxygen tension at various elevations, indicate
temperature gradients in the air at various distances from sunny rock surfaces, and
compare the hourly temperature fluctuations at various depths in the soil during a
ty aical summer day. Comparisons such as these provide a valuable insight into the
nature of the contrasting microhabitats and ecological niches available to high
al itude insects, and help to explain how animal survival is possible in that essen-
tia lly hostile environment.
This remarkable volume is replete with facts, observations, and theories never
before brought together in a single volume. Biologists and ecologists will find it a
be ok that can be read with much interest and enjoyment, discussed in great depth
with one’s colleagues, then kept within easy reach for frequent utilization as a
va.uable reference source. J. Gordon Edwards, San Jose State College, California.
Sii-LEGGED Science. By Brian Hocking, Schenkman Publishing Co., Inc.,
Cambridge, Mass., vii -|- 199 pp., illustrated, 1968. $4.50.
The contents of this book can be divided into three parts. The first part, com-
prising chapters 1 through 7, is a general discussion of the attributes of insects.
Included in this discussion is an anecdotal commentary on the value and procedure
of scientific research (ch. 1), a brief statement of what insects are and how they
are related to other arthropods (ch. 2), and a discussion of sex-determination
(ch. 3), flight (ch. 4), audition (ch. 5), chemoreception (ch. 6), and vision (ch.
7). Within the second part, chapter 8 provides the reader with the names of
eighteen of the commoner orders together with the habits of one or two common
insects per order, while chapters 9 through 14 contain more extensive information
about the habits of the Coleoptera, Lepidoptera, Diptera, and Hymenoptera. The
th rd part, consisting of chapters 15 through 18, contains a discussion of some
in er actions between men and insects.
The author states in the preface that the book is for embryonic entomologists,
bi t any appeal the book may have to a later instar, larva, pupa, or imago is all
fo the better. Curiously enough, the development of the book, from the stand-
point of style and information content, parallels the development of insects, or,
fo: that matter, of human beings. The first part of the book is very infantile in
nature, has comparatively little information, and stylistically seems extremely
fo ced and contrived. For example, the author, in explaining about the chelicerae
of arthropods (p. 22) , points out that some creatures have grown knives and forks
in the form of cutters and jabbers and, as such, are spared from the toils of
di ihwashing. Presumably the reader is supposed to laugh. While this style of
wiiting may appeal to young children, it would hardly appeal to sophisticated
ad ults.
The second part of the book has a more mature, sophisticated presentation.
He :re the author seems less concerned with turning an amusing phrase at every
corner and more concerned with discussing matters in a lively, factual manner.
Chapter 8 has a novel and interesting manner of presenting the insect orders —
JULY 1969]
RECENT LITERATURE
243
each order is introduced by a few pertinent lines of verse. Unfortunately the
verses are, with a few notable exceptions, disappointingly uninspired and mediocre.
On the whole, this part will probably appeal to sophisticated adults, but its style
is a little too complex and intricate to have much appeal for children.
The third part of the book also has a mature presentation, but in places has
elements which resemble those of the first part. Just as humans lapse into a
“second childhood,” so also this part begins to lapse into a “senescence.” In
perusing this part, one can periodically come across puns and diagrams that, from
a biological standpoint, are quite amusing. For example, the author has a diagram
(p. 150) that depicts two stick figures of humans, one male with a white face,
the other female with a black face. Between the figures is an X of the type that
geneticists use to signify crossing, and there is a progeny diagram that depicts,
as progeny of this pair of stick figures, one white termite, one blank ant, and two
khaki ants. The uninitiated reader, perusing this book at random by flipping
pages, might come to the conclusion that either the author is given to using
euphemisms or the author is not familiar with the literature of genetics.
The book as an integral whole can best be described as an anomaly. Parts of
it seem designed for children, while other parts seem designed for adults. Although
it unquestionably has much good factual information that would be useful to the
embryonic entomologist, stylistically it may not settle well with the sophisticated
reader who wants to learn something about insects. — C. N. Slobodciiikoff, Uni-
versity of California, Berkeley.
RECENT LITERATURE
Occasional Papers, Bureau of Entomology, California Department of Agriculture.
Since the appearance of the notice (in Pan-Pac. Entomol. 41 (2) : 89, 1965), con-
cerning the first six issues of the Occasional Papers, Bureau of Entomology, Cali-
fornia Department of Agriculture, ten numbers of this publication have been
published in the past 3 1/2 years. These are:
No. 7, New California grasshoppers of the genus Melanoplus (Orthoptera, Acrid-
idae), by Ashley B. Gurney and George M. Buxton, 7 pp., 4 pis. 1965.
No. 8, A key to the genera of the subfamily Nitidulinae (Nitidulidae, Coleoptera)
and description of a new genus and new species, by Lorin R. Gillogly, 24 pp.,
2 figs., 1965.
No. 9, A new species of Ceratophyus found in California (Coleoptera : Scarabae-
idae), by 0. L. Cartwright, 7 pp., 2 pis., 1966.
No. 10, Keys for identifying larvae of Scarabaeoidea to the family and subfamily
(Coleoptera), by P. 0. Ritcher, 8 pp., 39 figs., 1967.
No. 11, Revision of the milliped genus Harpaphe Cook from Western North
America (Polydesmida : Xystodesmidae) , by John S. Buckett and Michael R.
Gardner, 51 pp., 12 pis., colored frontis, 1968.
No. 12, Revision of the Nearctic moth genus Abagrotis Smith with descriptions of
new species (Lepidoptera rNoctuidae) , Part 1, by John S. Buckett, 21 pp.,
1968.
No. 13, Grasshoppers of the genus Karokia and of the saltator, immunis, and
harperi species groups of the genus Melanoplus in California and Oregon
2i4
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 3
(Orthoptera : Acrididae) , by Ashley B. Gurney and George M. Buxton, 56
pp., 120 figs., 1968.
No. 14, Revision of the Nearctic moth genus Abagrotis Smith with descriptions of
new species (Lepidoptera : Noctuidae) , Part 2, by John S. Buckett, 16 pp.,
1968.
No. 15, Revision of the Nearctic moth genus Abagrotis Smith with descriptions of
new species (Lepidoptera : Noctuidae) , Part 3, by John S. Buckett, 29 pp.,
1968.
No. 16, Revision of the Nearctic moth genus Abagrotis Smith with descriptions of
of new species (Lepidoptera : Noctuidae), Part 4, by John S. Buckett, 27 pp.,
1968.
The masthead states that individual copies will be sent free of charge upon re-
q test and a regular mailing list will be maintained, including libraries and special-
ists, by writing to the editor — John S. Buckett, Bureau of Entomology, California
Department of Agriculture, 1220 N Street, Sacramento, California 95814. — P. H.
Arnaud, Jr., California Academy of Sciences, San Francisco.
SOCIETY NOTICES
A DVERTiSEMENT OF EXCHANGES. — A special advertising page is now available to
n embers and non-members who wish to advertise entomological exchanges, sales,
a inouncements, etc. We can offer an ad in four issues for $1.50 per line. Ads
V ill be arranged in order of their receipt. Contact the Advertising Manager, W. H.
lange. Department of Entomology, University of California, Davis, California
95616.
5 ALE OF Past Issues. — We are overstocked in early issues of the Pan-Pacific
Intomologist, and need to reduce our inventory to provide more storage space.
Therefore, we can offer Society members complete sets of volumes 1 through 16
(1924 through 1940) at the extremely reduced rate of $1.50 per volume or $24.00
for the entire first 16 volumes. These are available to Society members only,
c n a first-come, first-serve basis as long as the surplus lasts. Contact the Treasurer,
I. H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San
Francisco, California 94118.
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Insect Pins vi
D-Vac Company vi
Entomological Research Institute vii
Chemago Corporation viii
Entomological Exchanges and Sales ix
Publications of the Society x
Chevron Chemical Corporation Back cover x
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IX
ENTOMOLOGICAL EXCHANGES AND SALES
This page is available to members and non-members who wish to advertise
e itomological exchanges, sales, announcements, etc. We can offer an ad in four
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Vol. 45 OCTOBER 1969 No. 4
THE
Pan-Pacefic Entomologist
In Memoriam
HOWARD LESTER McKENZIE, JR.
1910-1968
MILLER, BOHART AND WILKEY- — Howard Lester McKenzie, Jr., 1910-
1968 245
OPLER — Two new species of Caloptilia associated with Rhus in California
(Lepidoptera: Gracilariidae) 259
BUTLER AND JOHNSON — The development of Cactobrosis fernaldialis and
Albareda parabates in relation to temperature (Lepidoptera: Pyralidae) 266
AKRE AND TORGERSON — Behavior of Vatesus beetles associated with army
ants (Coleoptera: Staphylinidae) 269
WICKMAN — A crossbreeding study of the cedar tree borer, Semanotus ligneus
amplus, and the fir tree borer, S. litigiosus (Coleoptera; Cerambycidae) 282
WILLIAMS — A new species of Syntropis from Baja California Sur, Mexico
with notes on its biology (Scorpionida: Vejovidae) 285
ALEXANDER — A new species of crane fly associated with the plant genus
Lopezia in Mexico (Diptera: Tipulidae) 291
BLACKWELDER — The nature of taxonomic data 293
CHEMSAK — Records and descriptions of Mexican and Central American
Tiilomorphini (Coleoptera: Cerambycidae) 303
VAN VOLKINBURG — Thinoseius spinosus found in new and unusual habitat
(Acari; Eviphididae) 318
NUTTING — Distribution and flights of rare North American desert termites of
the genus Amitermes (Isoptera: Termitidae) 320
BOOK REVIEW 325
NEW PAGE CHARGES 326
INDEX TO VOLUME 45 327
SAN FRANCISCO, CALIFORNIA • 1969
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
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Howard Lester McKenzie, Jr., 1910-1968.
The Pan-Pacific Entomologist
Vol. 45 October 1969 No. 4
Xtstcr HilrEenzie, Jr.
) 9 ) 0-1968
D. R. Miller, R. M. Bohart, and R. F. Wilkey
Systematic Entomology Laboratory, ARS, USD A, Washington, D.C.;
University of California, Davis; and
California State Department of Agriculture, Sacramento
Howard Lester McKenzie, Jr., age 58, died of cancer in Sacramento,
California on 17 October 1968 at the peak of his scientific career.
He was born in Lennox, Iowa on 25 May 1910 and was one of seven
children. While in Iowa, Howard McKenzie, Sr. supported his family
on the income derived from a combination furniture and hardware store.
He also made several real estate and stock investments which were to
support his family long after his death.
In 1923 the McKenzie family loaded their belongings into a rented
boxcar and moved to Pomona, California. Within a few years after
their arrival, Howard senior died and although this left Etta McKenzie
with a large family to support, her careful management of previous
investments left her with no financial problems.
Throughout grammar and high school, Howard was not overly con-
cerned with his education. He was on the high school basketball team
and spent most of his spare time doing odd jobs. When he finished
his secondary education he had no intention of going on to college, but
through the strong urging of his mother he entered Chaff ey Junior
College in 1928. While at Chaffey he was impressed by the teaching
of Professor George P. Weldon, Economic Entomologist and Pomologist,
who inspired Howard to continue his education. When Howard was
asked by Dean Booth what his intentions for the future were, he could
not answer except to say that he had enjoyed the subject matter of the
courses taught by Professor Weldon. Since, at the time, there was a
large demand for entomologists. Dean Booth suggested that he continue
his education with entomology as his field of emphasis.
The Pan-Pacific Entomologist 45; 245-259. October 1969
;!48 THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
So with this encouragement and inspiration, Howard entered the Uni-
ersity of California at Berkeley where he soon came under the influence
of Professor E. 0. Essig. In later years, Howard often spoke kindly of
Professor Essig and frequently reminisced about the summer field
course and the many evening discussions led by Essig. In a letter
vvritten to Essig, Howard expressed his appreciation in these words,
“I just can’t thank you enough for all that you have done for me, Professor
Essig, and remember if ever I can help you in any way, just let me
know, and I will surely do so.” Other instructors at Berkeley who helped
kindle Howard’s entomological enthusiasm were Professors W. B. Herms
and E. C. Van Dyke.
One of the requirements of those days was a Bachelors thesis. Under
the direction of Professor Essig, Howard did a rather extensive paper
on the biology of the ladybird beetle Hyperospis lateralis. This was
later to be the basis for his first publication (I) .
After graduating from Berkeley in 1932, Howard accepted a job at
the Riverside Experiment Station as Laboratory Assistant under the
auspices of Professor H. J. Quayle. Although his salary was meager,
he enjoyed his stay at the experiment station. He continued to work on
biological problems and as a result published major works on avocado
insects (6) and the gladiolus thrips (5) . In his own words he describes
his duties at Riverside: “Recorded information on life histories and
habits of California avocado insects and mites, the gladiolus thrips and
other insects attacking certain ornamental and subtropical plants, con-
ducted systematic work in identifying insects submitted to the University,
conducted extensive fumigation experiments with HCN to control scale
insects on avocado.” The diversity of tasks performed in this position
greatly broadened his entomological background.
While at Riverside he began courting Virginia Burdette Rountree
whom he married in 1933, These were wonderful days for Howard,
but he was bothered by extended stays at Encinitas Research Station
which was too far from Riverside to allow for daily commuting.
After Howard had worked in Riverside for over a year. Professor
Quayle recognized his potential as an entomologist, and with the hope
of encouraging him to continue his formal studies, offered to pay him
75 dollars a month to return to Berkeley and get a Masters degree.
Realizing the need for more education, he and Virginia accepted the
offer and spent the year of 1933-34 in Berkeley. His Masters thesis
was on a portion of the work which he had undertaken at Riverside and
was entitled, “Biology and control of the omnivorous looper Sahulodes
OCTOBER 1969] MILLER, BOHART, WILKEY HOWARD L. MCKENZIE 249
caberata Guenee, on avocado.” As before his major professor was E. 0.
Essig.
In 1934 he returned to Riverside and worked on diverse problems.
During this period some of his attention was turned toward strictly
taxonomic problems which were ultimately his greatest love. In later
years he stated that his discovery of a method to distinguish between
red and yellow scales was one of his most important achievements (12) .
The known taxonomic differences up to that time were extremely subtle,
requiring a detailed knowledge of the group. Since only red scale was
a serious economic pest, the need for a simple, accurate means to dis-
tinguish it from yellow scale was a pressing problem especially for
people involved with quarantine and biological control programs.
While Howard was working on the red scale paper in 1936, he became
associated with Professor G. F. Ferris of Stanford University. Ferris
reviewed the paper and made several helpful suggestions. From that
time until the death of Professor Ferris in 1958, they remained close
friends. Ferris urged Howard to continue his taxonomic studies on
the scale insects and taught him a great deal about the Coccoidea.
Howard often described his first attempt to draw a scale under the
direction of Ferris. Much to Howard’s chagrin his first illustration was
rejected with instructions to redo the entire illustration. Greatly dis-
appointed, “^‘Mac,” as Ferris often called him, did so but it was again
rejected. Finally bis third attempt was accepted. This lesson greatly
impressed Howard and even his last illustrations reflect the care de-
manded by Ferris many years ago.
In 1937 Howard accepted a position as Assistant Entomologist with
the U. S. Department of Agriculture to work on scale insects attacking
pines. Although his winter headquarters were in Berkeley, he spent his
first two summers in Arizona working on a margarodid in the genus
Matsucoccus. During this period he did some excellent biological work
and proved that this scale was causing serious damage to the pines.
In so doing he was also able to work out the biology of several species.
In his spare time he continued taxonomic studies and published an
extensive revision of the diaspidid genera Aonidiella and Chrys omphalus
(18).
These were pleasant days for Howard; when he was not working,
he and Virginia spent their time on picnies in many of the scenic areas
near Prescott, Arizona.
During the first winter in Berkeley, Howard spent all of his spare
time with Professor Ferris; they frequently worked together long after
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
2 SO
n lidnight. Howard was becoming an important entomologist in his own
r ght under the influence of two of the foremost workers of that time —
E. 0. Essig and G. F. Ferris. Also, his interest in taxonomy was be-
coming much stronger even though he was employed primarily to do
biological research.
From 1939 to 1942, Howard continued his work with the margarodids
believing, but never substantiating, the fact that Matsucoccus on an
i)idividual tree often preceded attack by bark beetles. During this
period he continued his field work, but it was apparent that his greatest
interest was taxonomy. He disliked having to stay away from his family
for extended periods of time and therefore began to search for new
e mployment.
With a broad entomological background he qualified for Assistant
Systematic Entomologist with the California State Department of
rgiculture, and he began working in this position 1 July 1942. He
V rites in a letter to Professor Quayle, “It certainly is a real pleasure
to be working with Entomologists who appreciate what scale insects
can do, and also who appreciate the value taxonomy has in connection
\ ith many of the economic insect problems.” For the next 16 years
Howard was able to concentrate on taxonomic problems and since this
\ as his greatest love, his publications were both plentiful and valuable.
He continued to confer with Professor Ferris and as a result their
friendship became stronger; rather than teacher and student, they were
r ow colleagues. The diversity of Howard’s interests within the Coc-
coidea made him one of the leading coccidologists in the world; he
1 ad published on the taxonomy of the Diaspididae, Asterolecaniidae,
£nd Margarodidae, with studies on zoogeography, life histories, plant
c amage, and possible mechanisms of control. During his employment
T/ith the California State Department of Agriculture, his major con-
tribution was his first book entitled, “The Armored Scale Insects of
(California” (49).
Although his publications during this period were restricted to the
scale insects, his identification responsibilities required an extensive
1 nowledge of the systematics of the insects as a whole. He was therefore
£ ble to maintain a broad understanding of the current taxonomic work
i 11 many groups other than the Coccoidea, such as Orthoptera, Hemiptera,
the other groups of the Homoptera and miscellaneous orders.
Much time was spent on collecting trips with Hartford Keifer,
Howard’s “boss” while with the State. They were mostly in search of
s oft scales of the genus Lecanium. As a result, there are many hundreds
OCTOBER 1969] MILLER, BOHART, WILKEY — HOWARD L. MCKENZIE 251
of slide mounts of these scales in the CDA collection which will hopefully
help to unravel this confused genus in the future.
In September 1954 Ferris was nearly through with volume 7 of his
“Atlas of the Seale Insects of North America” when suddenly he lost
part of the vision in his right eye. In addition he was having a great deal
of difficulty organizing one of the more complicated keys in this volume.
He writes in a letter to Mac, “My mind has simply given out for things
like that. When I try to work with anything at all difficult my mind
simply quits work and will not function. I have tried twice to complete
this key, with no results. The doctor suggests that the years of high
blood pressure have caused some capillaries in the brain to deteriorate,
with an incipient cerebral haemorrhage . . . Do you suppose you could
come down for a day and help me out?” This Howard did and the
seventh volume of the atlas was completed. The fact that Ferris asked
Howard to help him, demonstrates the great amount of respect that
Ferris had for his Sacramento colleague.
Although Ferris recovered from his blood pressure problems, he was
never again able to work to full capacity. He passed away on 21 April
1958 leaving Howard with the responsibility of carrying on and im-
proving upon the work to which he had dedicated his life.
In that same year Howard was given the opportunity to assume this
responsibility as a full-time endeavor. Mainly through the efforts of
Dr. Alfred Boyce of the University of California at Riverside, Howard
was offered a position as Associate Entomologist in the Experiment
Station at the University of California either at Davis or Riverside.
He gladly accepted the offer and exclaimed to his wife that it did not
seem right for someone to actually pay him to do the one thing that he
loved most — scale taxonomy.
He decided to do a companion volume to his first book as a revision
of the mealybugs of California. He was so enthusiastic about his work
that by the time he had been at Davis for three years, he had established
himself as a world authority on the mealybugs. Since he had never
published on this extremely large and difficult group of scale insects,
this was truly a remarkable feat.
In 1960 it was apparent that Howard was quite capable of filling the
shoes of Professor Ferris. Realizing this, officials at Stanford Uni-
versity felt that it was a waste not to have the Ferris collection in the
hands of an active coccidologist and, therefore, presented the entire
Stanford Coccoidea collection as a gift to the University of California,
Department of Entomology, Davis.
With this collection as a basis, Howard worked diligently on his final
252
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
masterpiece. In 1967 the most important publication of his life was in
print. It was published by the University of California Press as a
technical book and is entitled “Mealybugs of California, with Taxonomy,
Biology, and Control of North American Species” (67). This book
treated coccidology broadly with sections on economic importance,
control, ecology, biology, cytology, field and laboratory methods,
morphology, and most important, taxonomy. The book is truly spec-
tacular with many water color paintings, color photographs, and minutely
detailed line drawings. This classical book shall always remain as a
memorial to Howard L. McKenzie.
Howard was an extremely meticulous man and his work reflects this
characteristic. People who worked with him often found it difficult
since he demanded so much attention to detail, but it was this detail
that made his work of such excellent quality. He was also a person
wiio would not accept “no” for an answer. If there was any possibility
of achieving his goal he would keep at it until it was fulfilled. He had
the ability to get people to do just about anything. Only the combination
of these talents could have brought together the book described above.
Howard was well known by his friends as a jovial, happy-go-lucky
individual. He could always cheer a person up no matter what the
cenditions. He nearly always had new jokes to tell and although they
teided to be somewhat “corny,” it was not possible to keep from
la aghing with him. Along with his sense of humor he had the admirable
quality of being extremely optimistic. In addition, he intensely disliked
hurting anyone and would go to extremes to avoid any sort of unfavor-
able confrontation. His friends and colleagues are grateful to have
known him and sad to have lost him.
Within the scientific community Howard maintained a strong interest
in entomological affairs. He was a member of the Entomological Society
of America and the Pacific Coast Entomological Society. At the time
of his death he was President-Elect of the latter society.
In Sacramento he was a well known and well respected member of
the community. He was very active in his church and by this means
was an inspiration and friend to many people.
Throughout his life he developed several hobbies. His early interest
was in breeding show pigeons and as most people who knew him will
recall, he never did anything halfheartedly. Virginia McKenzie de-
sc ribed his pigeon facilities as very extensive and smiles as if to empha-
size the word “very.” He was apparently quite successful in this endeavor
fer he amassed an extensive collection of ribbons.
After arriving in Berkeley he was encouraged to make several class
OCTOBER 1969] MILLER, BOHART, WILKEY — HOWARD L. MCKENZIE 253
insect collections and with these as a nucleus he continued until he had
built a collection which contained approximately 14,000 specimens. As
was typical of “Mac” the collection was well curated, with each specimen
carefully placed and identified. Through the courtesy of Virginia Mc-
Kenzie, this collection was recently donated to Davis where it is
presently being incorporated.
A third hobby and perhaps one of Howard’s greatest loves was
gardening. His love of plants was probably influenced or perhaps even
started by Professor Essig who was a world renowned horticulturist.
In a large percentage of the extensive correspondence between Essig
and McKenzie the topic of gardening endeavors nearly always came up.
In later years, Howard spent a day or two of vacation each month
working in his garden. For those who had the privilege to visit his
home, his backyard always reflected the “green thumb” of its owner.
Howard is survived by his wife, Virginia, a daughter, Mrs. Linda
Briggs, a son, David, 2 grandchildren, 4 sisters, and a brother.
The following is a bibliographical list of scientific writings by Howard
L. McKenzie:
1 .
2 .
3.
4.
5.
6 .
7.
8 .
9.
10 .
11 .
12 .
13 .
1932. The biology and feeding habits of Hyperaspis lateralis Mulsant.
Univ. Calif. Publ. EntomoL, 6 (2) : 9-20, illus.
1933. Observations on the genista caterpillar Tholeria reversalis Guenee.
Bull. Calif. Dep. Agr., 22 (7) : 410-12, illus.
1933. Note on the tarantula hawk wasps. Pan-Pac. Entomol., 9 (4) : 159.
1934. The green fruit or peach beetle, Cotinis texana Casey, in California.
J. Econ. Entomol., 27: 1110.
1935. Life history and control of the gladiolus thrips in California. Univ.
Calif. Agr. Exp. Sta. Circ. 337, 16 pp., illus.
1935. Biology and control of avocado insects and mites. Univ. Calif. Agr.
Exp. Sta. Bull. 592, 48 pp., illus.
1935. Life history and control of latania scale on avocado. Calif. Avocado
Yearb., 1935: 80-82.
1936. Say’s plant bug, Chlorochroa sayi Stab, on beets in California. Pan-Pac.
Entomol., 12 (1) : 18.
1936. A bermuda grass diaspine scale new to California. Pan-Pac. Entomol.,
12 (2) : 96.
1936. An anatomical and systematic study of the genus Anatis of America.
Univ. Calif. Publ. Entomol., 6 (10) : 263-72, illus.
1936. Rhigopsis effracta Lee. attacking grape buds in California. Bull. Calif.
Dep. Agr., 25 (2) : 272.
1937. Morphological differences distinguishing California red scale, yellow
scale, and related species. Univ. Calif. Publ. Entomol., 6 (13) : 323-
35, illus.
1937. Atmospheric HCN fumigation for latania scale on avocado fruits. Bull.
Calif. Dep. Agr., 25 (3) : 369-76, illus. (by Lindgren, D. L. and H. L.
McKenzie) .
254
14
1937.
15
1937.
16
1937.
17
1938.
18
1939.
19
1941.
20
1941.
21
1941.
22,
1942.
23
1942.
24.
1942.
2£.
1943.
2f.
1943.
2U
1943.
21.
1943.
2(,
1944.
3(.
1945.
3 .
1946.
32.
1946.
33.
1946.
1946.
3 ).
1947.
3 5.
1947.
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45 , NO. 4
Contributions to the knowledge of the Coccoidea V. Microentomology,
2 (2) : 47-101, illus. (hy Ferris, G. F., D. D. Jorgensen, and H. L.
McKenzie) .
Avocado fumigation investigation. Bull. Calif. Dep. Agr., 26 (3) :
311-19, illus. (By McKenzie, H. L. and D. L. Lindgren) .
Generic characteristics of Aonidiella Berlese and Leonardi, and a
description of a new species from Australia. Pan-Pac. EntomoL, 13
(4) : 176-80, illus.
The genus Aonidiella. Microentomology, 3 (1) : 1-36, illus.
A revision of the genus Chrys omphalus and supplementary notes on
the genus Aonidiella. Microentomology, 4 (2) : 51-77, illus.
A new species of the genus Matsucoccus attacking pinon pine in Cali-
fornia. Microentomology, 6 (1) : 2-5, illus.
Injury hy sugar pine matsucoccus scale resembles that of blister rust.
J. Forest., 39 (5) : 488-89.
Matsucoccus bisetosus Morrison, a potential enemy of California pines.
J. Econ. Entomol., 34 (6) : 783-85, illus.
New species of pine-infesting Margarodidae from California and south-
western United States. Microentomology, 7 (1) : 1-18, illus.
Seasonal history of the margarodid scale, Matsucoccus bisetosus Morri-
son, occuring on ponderosa and Jeffrey pines in California. Micro-
entomology, 7 (1) : 19-24, illus.
Two new species related to red scale. Bull. Calif. Dep. Agr., 31 (3) :
141-47, illus.
Miscellaneous diaspid studies including notes on Chrysomphalus. Bull.
Calif. Dep. Agr., 32 (2) : 148-62, illus.
Errata. Change of deposition of types of Aonidiella paucitatis Mc-
Kenzie. Bull. Calif. Dep. Agr., 32 (3) : 187.
The seasonal history of Matsucoccus vexillorum Morrison. Micro-
entomology, 8 (2) : 42-52, illus.
Notes on Matsucoccus vexillorum Morrison. Microentomology, 8 (2) :
53-57, illus.
Miscellaneous diaspid scale studies. Bull. Calif. Dep. Agr., 33 (1) :
53-59, illus.
A revision of Parlatoria and closely allied genera. Microentomology,
10 (2) : 47-121, illus.
Supplementary notes on the genera Aonidiella and Parlatoria. Micro-
entomology, 11 (1) : 29-36, illus.
A new species of Lepidosaphes attacking Dendrobium orchids in Hawaii
and California. Proc. Hawaiian Entomol. Soc., 12 (3) : 611-13, illus.
General distribution of red scale, Aonidiella aurantii (Maskell) in
California. Bull. Calif. Dep. Agr., 35 (2) : 95-99, illus.
Distribution of red scale in California. Calif. Citrogr., 31 (11) : 428-29,
illus.
Diaspid scale studies, with notes on California species. Bull. Calif.
Dep. Agr., 36 (1) : 31-36, illus.
Miscellaneous diaspid scale studies. Part V. Bull. Calif. Dep. Agr.,
36 (3) : 107-14, illus.
OCTOBER 1969] MILLER, BOHART, WILKEY HOWARD L. MCKENZIE 255
37. 1947. Correction: Lepidosaphes noxia McKenzie. Proc. Hawaiian Entomol.
Soc., 13 (1) : 31.
38. 1948. The Prescott scale {Matsucoccus vexillorum) and associated organisms
that cause flagging injury to ponderosa pine in the southwest. J. Agr.
Res., 76 (2) : 33-51, illus. (by McKenzie, H. L., L. S. Gill, and D. E.
Ellis).
39. 1949. A new introduced diaspidid scale of possible economic concern in
California. Scale studies- — part VI. Bull. Calif. Dep. Agr., 38 (3) :
123-26, illus.
40. 1950. The genera Lindingaspis MacGillivray and Marginaspis Hall. Micro-
entomology, 15 (3) : 98-124, illus.
41. 1951. Miscellaneous diaspidid scale studies. Scale studies — part VH. Bull.
Calif. Dep. Agr., 40 (2) : 1-3, illus.
42. 1951. Present status of the kuno scale, Lecanium kunoensis Kuwana, in
California. Bull. Calif. Dep. Agr., 40 (3) : 105-09, illus.
43. 1952. New parlatoriine scales from India and Egypt, and supplementary notes
on other related species. Scale studies — part IX. Bull. Calif. Dep. Agr.,
41 (1) : 9-18, illus.
44. 1952. Present status of the olive pollinia scale, Pollinia pollini (Costa), in
California. Bull. Calif. Dep. Agr., 41 (2) : 115-21, illus. (by Armitage,
H. M. and H. L. McKenzie) .
45. 1952. Distribution and biological notes on the olive parlatoria scale, Parla-
toria oleae (Colvee), in California. Scale studies — part X. Bull. Calif.
Dep. Agr., 41 (3) : 127-38, illus.
46. 1953. A new scale insect from the Ryukyu Islands related to red scale. Scale
studies — part XI. Bull. Calif. Dep. Agr., 42 (1) : 1-4, illus.
47. 1953. Two new Selenaspidus scales infesting Euphorbia in California. Scale
studies — part XII. Bull. Calif. Dep. Agr., 42 (2) : 53-58, illus.
48. 1955. A new species of Lepidosaphes scale infesting umbrella pine in Cali-
fornia. Pan-Pac. Entomol., 31 (4) : 187-90, illus.
49. 1956. The Armored Scale Insects of California. Bull. Calif. Insect Surv.,
vol. 5. Univ. Calif. Press, 209 pp.
50. 1957. A new armored scale insect on Elaeagnus from Texas. Scale studies —
part XHI. Bull. Calif. Dep. Agr., 46 (3) : 218-20, illus.
51. 1958. A new asterolecaniid scale on succulents from Mexico. Pan-Pac.
Entomol., 34 (3) : 169-72, illus.
52. 1959. Gordon Floyd Ferris as a student of the scale insects. Pan-Pac. En-
tomol., 35 (1) : 25-28.
53. 1959. Soft scales infesting walnut. Calif. Agr., 13 (5) : 6, 13, illus. (by
Michelbacher, A. E., H. L. McKenzie, and C. Q. Gonzales).
54. 1960. A new subterranean Rhizoecus mealybug from Arizona. Pan-Pac.
Entomol., 36 ( 3) : 139-41, illus.
55. 1%0. Taxonomic study of California mealybugs, with descriptions of new
species. Hilgardia, 29 (15) : 681-770, illus.
56. 1960. Taxonomic position of Parlatoria virescens Maskell, and descriptions
of related species. Scale studies — part XIV. Bull. Calif. Dep. Agr.,
49 ( 3) : 204^11, illus.
57. 1961. Second taxonomic study of California mealybugs, with descriptions
of new species. Hilgardia, 31 (2) : 15-52, illus.
25 (.
58. 1961.
59. 1962.
60. 1962.
61. 1962.
62. 1963.
63. 1964.
64. 1964.
65. 1965.
66. 1967.
67 1967.
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45 , NO. 4
Systematic status of the Pseudococcus maritimus — malacearum complex
of mealybugs. Bull. Calif. Dep. Agr., 50 (4) : 245^9, illus. (by Wilkey,
R. F. and H. L. McKenzie) .
Third taxonomic study of California mealybugs, including additional
species from North and South America. Hilgardia, 32 (14) : 637-88,
illus.
Two new species and additional collection records for the genus
Protodiaspis. Hilgardia, 33 (4) : 133-39, illus. (by McKenzie, H. L.
and W. A. Nelson-Rees) .
Evolutionary patterns in tbe armored scale insects and their allies.
Hilgardia, 33 (4) : 141-70a, illus. (by Brown, S. W. and H. L. Mc-
Kenzie) .
Miscellaneous diaspidid scale studies, including a new asterolecaniid
from Florida. Scale studies — part XV. Bull. Calif. Dep. Agr., 52 (1) :
29-39, illus.
Fourth taxonomic study of California mealybugs, with additional
species from North America, South America, and Japan. Hilgardia,
35 (10) : 211-72, illus.
Two new eriococcid scales from California. Scale studies — ^part XVI.
Bull. Calif. Dep. Agr., 53 (1) : 21-25, illus.
Fifth taxonomic study of North American mealybugs, with revisional
notes on seven species. Hilgardia, 37 (1) : 1-15, illus. (by McKenzie,
H. L. and D. J. Williams).
A systematic study of Ovaticoccus Kloet and its relatives, with a key to
North American genera of Eriococcidae. Hilgardia, 38 (13) : 471-539,
illus. (by Miller, D. R. and H. L. McKenzie).
Mealybugs of California; with taxonomy, biology and control of North
American species. Univ. Calif. Press, Berkeley, 525 pp.
List of Names Proposed by Howard L. McKenzie
TFIBE
Margarodidae
Pi yococcini (22)
GENERA
Diaspidid AE
A1 riconidia ( 36 )
Helaspis (62)
Ntopinnaspis (39)
Pirlagena (30)
Ptrlaspis (30)
Asterolecaniidae
Sclerosococcus (51)
Margarodidae
Desmococcus (22)
Pityococcus (22)
PSEUDOCOCCIDAE
A] ithelococcus (63)
Chorizococcus (55)
Circaputo (59)
Paradoxococcus (59)
Pygmaeococcus (55)
Scaptococcus (63)
SPECIES
DiASPmmAE
Acutaspis
subnigra (35)
tingi (35)
Af riconidia
halli (36)
Annutaspis
singularis (62)
Aonidiella
comperei (12)
ensifera (24)
eremocitri (16)
OCTOBER 1969] MILLER, BOHART, WILKEY HOWARD L. MCKENZIE 257
inornata (17)
longicorna (18)
marginipora ( 31 )
messengeri (46)
paucitatis (24)
Chrysomphalus
nulliporus (18)
variabilis (25)
Diaspidiotiis
mccombi (62)
Diaspis
conocarpi (36)
ferrisi(36)
gilloglyi (62)
parasiti (36)
Helaspis
mexicana (62)
Hemiberlesia
fraxini (29)
meiidax (25)
pseudorapax (41)
Lepidosaphes
mackieana (25)
noxia (32)
sciadopitysi (48)
Lindingaspis
ferrisi (40)
fusca (25)
magiiifica (25)
neorossi (40)
similis (40)
tingi (4-0)
Melanaspis
elaeagni (50)
tenax (29)
Neopiiinaspis
harperi (39)
Parlagena
inops (30)
Parlatoria
acalcarata (56)
boycei (43)
citri (25)
crypta ( 25 )
desolator (56)
marginalis (30)
morrisoni (25)
multipora (30)
perplexus (43)
vandae (56)
Protodiaspis
chichi McKenzie and Nelson-Rees
(60)
cinchonae ( 29 )
didymus McKenzie and Nelson-Rees
(60)
P seudoparlatoria
browni (62)
Selenaspidus
albus (47)
rubidus (47)
Asterolecaniidae
Sclerosococcus
bromeliae (62)
ferrisi (51)
Margarodidae
Desmococcus
captivus (22)
sedeiitarius (22)
Matsucoccus
moiiophyllae ( 19 )
Pityococcus
deleoni (22)
ferrisi (22)
rugulosus (22)
Eriococcidae
Ovaticoccus
calif ornicus (64)
senarius (64)
PSEUDOCOCCIDAE
Anisococcus
abnormalis (63)
imperialis (67)
Anthelococcus
simondsi (63)
Balanococcus
takahashii (63)
Chorizococcus
abortivus (67)
abroniae (55)
brevicruris (55)
calif ornicus (63)
coxindex (67)
fistulosus (67)
interruptus (63)
microporus (55)
253
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
penultimatus (67)
polyporus (57)
psoraleae (55)
senarius (67)
snellingi (67)
variabilis (63)
'ivilkeyi (55)
'kvilsoni (57)
yuccae (57)
Cii caputo
Ihiirsutus (59)
Cryptoripersia
tubulata (63)
cococcus
spectabilis (57)
Du tchlicoccus
arundinis (67)
megacirculus (67)
Dy smicoccus
desertorum (59)
burdi (59)
pinicolus (63)
racemus (67)
vacuatus (67)
He liococcus
adenostomae (55)
atriplicis (63)
Hi mococcus
caritus (55)
ceraricus (63)
inornatus (55)
Ps radoxococcus
mcdanieli (59)
Phenacoccus
advena (63)
alleni (63)
cajonensis (67)
destitutus (67)
dicoriae (57)
echeveriae (55)
eschscholtziae (57)
giganteus (63)
graminosus (55)
hurdi (63)
incomptus (63)
infernalis (59)
lotearum (55)
megaulus (67)
milleri (63)
multisetosus ( 67 )
nonarius (63)
tibiaegracilis (55)
Pseudococcus
aberrans (59)
dispar (59)
diversus (63)
importatus (55)
macswaini (59)
megasetosus (67)
microcirculus (55)
pertusus (67)
prunicolus (63)
pseudobscurus ( 63 )
sparsus (59)
Puto
acirculus (55)
albicans (67)
atriplicis (57)
calif omicus (67)
decorosus (67)
echinatus (57)
laticribellum (57)
mimicus (67)
nulliporus (55)
pacificus (67)
pricei (55)
profusus (55)
simmondsiae (57)
usingeri (59)
Pygmaeococcus
morrisoni (55)
Rastrococcus
chilensis (63)
Rhizoecus
bicirculus (67 )
bituberculatus (55)
bobarti (54)
browni (57)
disjunctus (67)
eluminatus (55)
gracilis (57)
menkei (59)
neomexicanus (59)
pritchardi (55)
sonomae (55)
spinosus (55)
stangei (59)
OCTOBER 1969]
OPLER TWO NEW CALOPTILIA
259
Scaptococcus
californicus (63)
milleri (67)
Spilococcus
cactearum ( 55 )
ceanothi (67)
corticosus (67)
haigi (59)
keiferi (55)
parkeri (59)
parvicirculus (55)
quercinus (59)
ventralis (67)
T rionymus
frontalis (67)
furvus (67)
haancheni (55)
myersi (57)
quadricirculus (67)
winnemucae (67)
Some of the insects and related groups named in honor of Howard
L. McKenzie are as follows:
ACARINA
Aculodes mckenziei Keifer (Eriophyidae)
Amblyseius mckenziei Schuster and Pritchard (Phytoseiidae)
HOMOPTERA
Dysmicoccus mackenziei Beardsley (Pseudococcidae)
Humococcus mackenziei Ezzat (Pseudococcidae)
Lindingaspis mackenziei Williams (Diaspididae)
Ovaticoccus mackenziei Miller (Eriococcidae)
Parlagena mckenziei Balachowsky (Diaspididae)
Selenediella mckenziei Takahashi (Diaspididae)
DIPTERA
Tujunga mackenziei Steyskal (Otitidae)
Two New Species of Caloptilia Associated with Rhus
in California
(Lepidoptera: Gracilariidae)
Paul A. Opler
University of California, Berkeley
Sweet (1969) carried out an extensive study of the life histories and
host preferences of two species of Caloptilia which occur in California.
While attempting to identify these species it was discovered that they
are both undescribed. In this paper I will describe both of these species
and will compare them with Caloptilia rhoifoliella (Chambers) : New
Combination, the other species in the complex. All three species feed
upon members of the genus Rhus (Anacardiaceae) .
The generic name Caloptilia is used for these species in preference to
Gracilaria in accordance with the revision of the family by Vari (1961) .
The Pan-Pacific Entomologist 45: 259-266. October 1969
260
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Caloptilia ovatiella Opler, new species
(Figs. 1, 6, 7, 10)
Male. — Alar expanse, 11-12 nun. Head', palpi upturned, scaled with white and
dark brown intermixed; Irons covered with white scales, few dark brown scales in-
termixed; vertex dark brown, paler posteriorly; antennae dark brown. Thorax:
light brown on dorsum, dark brown on pleural areas and sternum; pro- and meso-
tloracic legs dark brown basally; tibia and tarsi white-scaled; metathoracic legs
w th coxae and based third of trochanters white-scaled; apical two-thirds of tro-
chanters dark brown ; femora, tibiae, and tarsi with mixture of white and pale brown
scales. Forewing: brown, with lavender iridescence when alive or recently so, fad-
irg to pale sheen in museum specimens; patch of dark brown scales along basal
fi;th of costal margin followed by long white patch extending nearly to apex en-
closing about six small patches of dark brown; few light brown scales intermixed
tl roughout white patch as well ; narrow strip of dark brown scales along middle
h df of inner margin; remainder of wing brown with small scattered patches of
dirk brown scaling, becoming heavier apically; ventral surface dark brown, nar-
rc wly edged with light brown on margins; fringe on outer margin of broadened
dirk brown scales; fringe scales along inner margin long and tan. Hindwing: tan
with pale fringe. Abdomen: clothed with tan scales. Genitalia: vesica with linear
row of 24 approximately parallel straight spines; aedeagus strongly curved; valvae
with dorsally directed flap-like process one-third width of valva projecting from
ventral margin of inner face at basal one-third; transverse bar-like process on inner
fi.ce of valva from base at dorsal edge to ventral edge at two-thirds; process with
slightly bulbous area at juncture with ventral margin covered with long inwardly
d rected setae.
Female. — Alar expanse, 12-13 mm. In male except: female genitalia with two
e lually sized curved signa.
Holotype male, Eaton Canyon, near Altadena, Los Angeles County,
California, Harold Sweet laboratory colony, reared from Rhus ovata,
emerged 26 August 1968, P. Opler genitalia prep. 7, deposited in Cali-
f Drnia Academy of Sciences.
Paratypes: 22, same data except emerged July 1968 (5), 19 August-3 Septem-
ber 1968 (17) ; 10 same locality data reared from Rhus laurina, emerged 2-24
/ ugust 1968; and 29 from South Laguna Beach, Orange County, California, Harold
Sweet laboratory colony, reared from Rhus integrifolia, emerged 2-20 August
1968; one paratype San Gabriel Wash, 5 mi. S. Azuza, Los Angeles County, Cali-
fornia, 29 March 1968, JAP 68C72, reared from Rhus laurina, emerged 21 April
1968, P. A. Opler. Paratypes deposited in collections of California Academy of
Figs. 1-3: Fig. 1. Caloptilia ovatiella, $, Eaton Cyn., Los Angeles Co., Calif., H.
fweet lab. colony, reared from Rhus ovata, emerged July 1968. Fig. 2. C.
civersilohiella, $, Strawberry Cyn., Alameda Co., Calif., H. Sweet lab. colony,
reared from Rhus diversiloba, emerged 17 August 1968. Fig. 3. C. rhoifoliella,
o , New Brighton, Pa., 18 August 1907, ex Merrick Museum, U.S.N.M.
OCTOBER 1969]
OPLER TWO NEW CALOPTILIA
261
232
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Sciences, California Insect Survey, Canadian National Collection, and U.S. National
]V useum.
Discussion. — Sweet (1969) reared this species from Rhus ovata, R.
k urina, and R. integrifolia. The distribution of this species coincides
\^ith that of its hosts in southern California (Sweet, 1969) and presum-
ably in northern Baja California.
Caloptilia diversilobiella Opler, new species
(Figs. 2, 4, 8, 11)
Male. — Alar expanse, 11-13 mm. Head: palpi upturned, tip of distal segment
white-scaled, remainder dark brown; frons white-scaled; vertex covered with an-
teriorly directed dark brown scales; antennae blackish brown. Thorax: dark brown
O’ orangish brown on dorsum and pleural areas; sternum white; pro- and meso-
tlioracic legs with tibiae and tarsi white; femora and trochanters dark brown;
coxae white; metathoracic legs with coxae and basal third of trochanters white-
scaled with few scattered dark brown scales; apical two-thirds of trochanters dark
brown; femora, tibiae, and tarsi with light tan scales. Forewing: with purplish irides-
cmce when alive, gradually fading in museum specimens; ground dark brown or
reddish-brown with scattering of dark brown scales, variable, if ground reddish-
brown then with patch of dark brown at base of costal margin; irregular white
patch along costal margin beginning at basal one-fifth, terminating just before
apex, widest at base; reddish-brown scales occasionally invading patch along inner
edge, sometimes white completely replaced by reddish-brown; narrow black marks
along costa within patch; ventral surface dark brown, narrowly edged with light
brown on margins; fringe on outer margin of broadened dark brown scales; fringe
clong inner margin of long pale tan scales. Hindwing: pale tan with fringe of long
pale tan scales. Abdomen: clothed with gray scales. Genitalia: vesica with posteri-
crly directed row of 15-17 hook-like spines; aedeagus relatively straight; valvae with
long thin setae on apical half of inner face extending to just basad of transverse bar
£ long ventral margin ; patch of relatively short stout setae near ventral edge of valva
1 asal to transverse har.
Female. — Alar expanse, 10-12 mm. External features as described for male;
genitalia not distinguishable from that of C. ovatiella.
Holotype male, Strawberry Canyon, Alameda County, California,
Harold Sweet laboratory colony, reared from Rhus diversiloha, emerged
17 August 1968, P. Opler genitalia prep. 6, deposited in California Acad-
emy of Sciences.
Paratypes: 55, same data except emerged May to June 1968 (20), 9-22 August
1968 (30), 23 November 1965 (3), 8 December 1965 (2); 64, Arroyo Seco, Los
/.ngeles County, California, Harold Sweet laboratory colony, reared from Rhus
civersiloha, emerged 2-19 August 1968; 22, Jarboe Gap, Butte County, California,
Harold Sweet laboratory colony, reared from Rhus diversiloha, emerged 9 June-6
July 1968; 13 additional paratypes collected as adults in field as follows: California,
(.'ontra Costa Co.: El Cerrito, 5 February 1961, 2 June 1960, 27 September 1960
(C. D. MacNeill, CAS) ; Richmond, 22 May 1959, 9 July 1959 (C. D. MacNeill,
OCTOBER 1969]
OPLER TWO NEW CALOPTILIA
263
Figs. 4-9. Male genitalia of Caloptilia species, Fig. 4. C. diversilobiella, right
valva removed. Fig. 5. C. rhoifoliella, left valva. Fig. 6. C. ovatiella, right valva
removed. Fig. 7. C. ovatiella, aedeagus. Fig. 8. C. diversilobiella, aedeagus. Fig.
9. C. rhoifoliella, aedeagus.
254
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Figs. 10-11. Female genitalia of Caloptilia species, Fig. 10. C. ovatiella. Fig. 11. C.
iHversilobiella.
CAS), 8 October 1966 (A. J. Slater, CIS) ; Russell Property, 4% mi. NE Orinda, 2
November 1967 (P. A. Opler, CIS) ; Humboldt Co.: near Myer’s Flat, 31 August
1960 (C. D. MacNeill, CAS) ; San Francisco Co.: San Francisco, ex Quercus agri-
,olia, 30 August 1926 (H. H. Keifer, CAS) ; San Mateo Co.: San Mateo, 2 mi. W.,
7 September 1926 (H. H. Keifer, CAS) ; Santa Clara Co.: New ALmaden, 5 July
964, 14-16 September 1963 (P. A. Opler, CIS). Paratypes deposited in collections
OCTOBER 1969]
OPLER — TWO NEW CALOPTILIA
265
of California Academy of Sciences, California Insect Survey, Canadian National
Collection, Los Angeles County Museum, and U.S. National Museum.
Discussion. — Caloptilia rhoifoliella, a species which occurs in eastern
North America, can be distinguished from C. diver silobiella only by ex-
amination of the male genitalia. The aedeagus of C. rhoifoliella (fig. 5)
has the vesica with a row of 24 triangularly shaped unhooked spines in
contrast with the lesser number of hooked spines on the vesica of C. div-
ersilobiella (fig. 8) . The valva of C. rhoifoliella has a more extensive
group of short, stout, setae on the inner face and, concomitantly, fewer
long narrow setae (fig. 5) than does C. diver silobiella (fig. 4). It is not
surprising that such a close relationship should exist between these two
species since the hosts, Rhus diversiloba for C. diver silobiella and Rhus
toxicodendron for C. rhoifoliella are closest to each other within the
genus Rhus in the taxonomic sense.
Sweet (1969) has reared Caloptilia diver silobiella from Rhus diversi-
loba', the moth occurs throughout the range of its host in California,
commonly in coastal localities and more rarely in the Sierra Nevada.
Caloptilia rhoifoliella has been reared from Rhus toxicodendron (Cham-
bers, 1876; Criddle, 1927) and Rhus copallina (Chambers, 1876; Raiz-
enne, 1952) . C. rhoifoliella occurs from Manitoba east to Ontario thence
south to Kentucky and North Carolina.
The specific distinctness of the two new species as well as the limits of
their host selection was clearly documented in the laboratory by Sweet’s
studies. Individuals from different populations within species would
freely breed in the laboratory, whereas in no case did copulation occur
between individuals of different species. Females of either species would
not oviposit on the host(s) of the other.
Five females, collected at Devil’s Punchbowl, Los Angeles County,
California at light on 1 May 1968 by P. Opler and J. A. Powell, appear
to be closely related to C. ovatiella, but they do not appear to fall within
the range of variation for the species as presented by the type series.
None of the hosts reported by Sweet occurs in that area according to
Munz and Keck (1959), although another species, Rhus trilobata, does
occur there. Thus, there is a distinct possibility that a third species of
the complex in California is involved. No attempt to deal with these
specimens will be made in the absence of associated males and a sub-
stantiated host association.
Acknowledgments
I wish to thank Harold Sweet, San Mateo College, whose provision of
specimens and enthusiastic encouragement formed the raison d’etre of
266
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
t\ is paper. Donald R. Davis, U.S. National Museum, loaned two speci-
nens of Caloptilia rhoijoliella to the author. Jerry A. Powell kindly re-
V ewed the manuscript of this paper.
Literature Cited
C iAMBERS, V. T. 1876. Tineina. Can. Entomol., 8(2) : 30-35.
C UDDLE, N. 1927. Lepidoptera reared in Manitoba from poison ivy. Can. Entomol.,
59(5) : 99-101.
]V UNZ, P. A. AND D. D. Keck. 1959. A California flora. Univ. Calif. Press, Berke-
ley and Los Angeles, 1681 pp.
RilzENNE, H. 1952. Forest Lepidoptera of Southern Ontario and their parasites.
Dep. Agr., Ottawa, Canada, 277 pp.
S VEET, H. 1969. Investigation of distribution, biology, ecology, natural enemies and
host preferences of the poison oak leafroller, Caloptilia diversilobiella.
Univ. Calif. Berkeley, Ph.D. thesis, unpubl.
V ^Ri, L. 1961. South African Lepidoptera. Volume 1. Lithocolletidae. Transvaal
Mus. Mem., 12, 238 pp., 112 pis.
The Development of Cactobrosis fernaldialis and Albareda
par abates in Relation to Temperature^
(Lepidoptera: Pyralidae)
George D. Butler, Jr.^ and Paul H. Johnson
Department of Entomology, University of Arizona, Tucson
Cactobrosis fernaldialis (Hulst) is the most important known vector of
tlie bacterial disease, Erwinia carnegieana Standring, of the giant cactus
or saguaro, Cereus giganteus Engelm. (= Carnegiea gigantea Britt, and
F ose). Butler and Werner (1965) gave light-trap records for this
s jecies as well as Albareda par abates (Dyar) and Cahela ponder osella
(B. and McD.), which are also cactus-eating moths, Boyle (1949) ob-
served C. fernaldialis in the field and laboratory and concluded that there
V as but one generation a year. Eggs required 30 days to hatch, the
larval stage in the giant cactus lasted from November to May, and the
pupal stage was from 28 to 33 days. The present study indicates that the
IJoumal Paper No. 1452 of the Arizona Agricultural Experiment Station.
2 Present address: USDA Cotton Insects Research Laboratory, 2000 East Allen Road, Tucson, Arizona
81719.
The Pan-Pacific Entomologist 45: 266-269. October 1969
OCTOBER 1969] BUTLER & JOHNSON CACTOBROSIS DEVELOPMENT 267
life cycle of this insect is considerably different when feeding on semi-
synthetic media in the laboratory. These results also provide informa-
tion to predict the rate of development for rearing programs, for estimat-
ing low and high temperature thresholds of development, and to use as a
major input for computer simulation of population fluctuations.
Methods and Materials. — Larvae of C. fernaldialis were collected
from saguaro buds in early June, 1966. They were placed in individual
1-oz. paper cups containing a lima bean-agar medium developed by
Shorey (1963), which is used as a standard laboratory diet for boll-
worms, cabbage loopers, salt-marsh caterpillars, and other larvae. The
cups were placed in a plastic bag in a 1-ft^ wire cage. The larvae chewed
holes through the cups when full-grown and crawled out. Some spun
cocoons between the cups while others chewed their way out of th^
plastic bag and pupated in the corners of the cage. Moths were held
in 1-qt cartons with a 1-dr vial of sugar-water in the lid. A strip of 32
X 32-mesh saran screening was placed in each carton on which most of
the eggs were laid. The eggs were removed daily, held in constant tem-
perature cabinets, and examined daily for hatching. First-instar larvae
were placed in 8-oz cups of media until about half-grown and then trans-
ferred to individual 6-dr cotton-stoppered shell vials. They were held
at different constant temperatures and examined daily to determine when
they pupated and when adults emerged. Several generations were reared
in the laboratory.
Constant temperature cabinets were maintained at 68°, 77°, 81°, 86°,
and 95°F and thermoswitches held the temperatures to ± 2.5°F. Humid-
ity was held near 50%. Lights were on for a 15-hr period each day
starting at 6 A.M. A regression equation, y = a + bX, where y is the re-
ciprocal of the duration of a stage and X is the temperature, was cal-
culated for the different periods of development, as discussed by Butler
(1966). Fahrenheit temperatures were used to permit calculation of
developmental time using U. S. Weather Bureau data. Transformation
of the regression equations to Celsius can be made according to Zar
(1969).
Results. — The duration of the egg stage of C. fernaldialis was 9 days
for one group of eggs at 68°F, 4 and 5 days for 2 groups of eggs at 77°F,
and 3 days for 6 groups of eggs at 86 °F. The duration of the larval and
the larval-plus-pupal stages at different temperatures is given in Table 1.
The larval stage varied from 34.2 days at 68° to 22.2 days at 81 °F and
increased in duration as the temperature rose above this point. The re-
gression equation for the larval stage between 68° and 81°F is y =
0.00124X - 0.0547 (r^ = 0.99) , where y is the reciprocal of the duration
268
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Table 1. The mean duration of the larval and the larval-plus-pupal
stages of Cactohrosis fernaldialis in relation to temperature.
Tempera-
ture
°F
Larval stage
Larval-plus-pupal stages
No. in-
Mean no.
No. in-
Mean no.
dividuals
days ±
SD
dividuals
days
± SD
68
23
34.2 ±
10.6
19
57.4
Hh
11.2
77
52
23.9 ±
7.0
46
38.1
6.2
81
14
22.2 ±
1.5
11
35.9
2.3
86
70
22.6 ±
6.7
52
35.4
6.5
95
7
28.0 ±
2.1
3
40.7
2.2
ol the larval stage and X is the temperature. The duration of the larval-
plas-pupal stages decreased from 57.4 days at 68° to 35.4 days at 86°F.
T]ie regression equation for this period between 68° and 86°F is y ==
0. 30083X — 0.0387 (r^ = 0.97) , where y is the reciprocal of the duration
oi the larval-plus-pupal stage in days and X is the temperature.
The life cycle of A. parabates was studied in a manner similar to that
ol C. fernaldialis except that moths were collected from a light trap. A
si igle group of eggs required 6 days to hatch at 68°F. The duration of
tb e larval and the larval-plus -pupal stages at 3 temperatures are given in
T ible 2. The regression equation for the larval stage is y = 0.00095X -
0.0385 (r^ = 0.93), where y is the reciprocal of the duration of the
laival stage and X is the temperature. The regression equation for the
larval-plus-pupal stages is y = 0.00080X - 0.0384 (r^ = 0.93), where y is
ti e reciprocal of the duration of the larval-plus-pupal stages and X is
tl e temperature. The equation for the larval-plus-pupal stages is almost
ic entical to that for C. fernaldialis, so the two species develop at a sim-
ilar rate and in a similar time.
Table 2. The mean duration of the larval and the larval-plus-pupal
stages of Alhareda parabates in relation to temperature.
Larval stage Larval-plus-pupal stages
1 empera-
ture
ojp
No. in-
dividuals
Mean no.
days
SD
No. in-
dividuals
Mean no.
days
SD
61
11
40.9
6.7
6
66.8
7.0
Ti
10
27.0
4.0
10
39.1
3.7
8(1
10
24.1
5.0
10
34.0
■±_
3.6
OCTOBER 1969] AKRE & TORGERSON VATESUS BEHAVIOR
269
Literature Cited
Boyle, A. M. 1949. Further studies of the bacterial necrosis of the giant cactus.
Phytopathology, 39(12) : 1029-1052.
Butler, G. D., Jr. 1966. Development of beet armyworm and its parasite Chelonus
texanus in relation to temperature. J. Econ. EntomoL, 59(6) : 1324^-1327.
Butler, G. D., Jr. and F. G. Werner. 1965. Light-trap records of three cactus-
eating moths in Arizona. Pan-Pac. Entomol., 41(1) : 10-12.
Shorey, H. H. 1963. A simple artificial rearing medium for the cabbage looper. J.
Econ. Entomol., 56(4) : 536-7.
Zar, j. H. 1969. The effect of the choice of temperature scale on simple linear re-
gression equations. Ecology 49(6) : 1161.
Behavior of Vatesus Beetles Associated with Army Ants
(Coleoptera: Staphylinidae) 1
Roger D. Akre and Richard L. Torgerson^
Washington State University, Pullman
In previous papers on staphylinid guests associated with army ants
(Akre and Rettenmeyer, 1966), guests were separated into categories on
the basis of their trail-following behavior and degree of host association.
Vatesus were placed into the group having the closest possible relations
with their host. Since then, more data has been obtained on the behavior
of Vatesus beetles and is reported.
The tribe Vatesini contains only one genus and more than 25 species
(Seevers 1965). All members are limulodid (horseshoe crab-shaped) in
form, but differ from other Trachyporines in that they are associated
with army ants. Discussion in this paper will be limited to the behavior
of 3 species of Vatesus on which both field and laboratory data were col-
lected.
Akre and Rettenmeyer (1966) , in the first behavioral paper on staphy-
linids associated with army ants, discussed the behavior of the Aleo-
charinae, Paederinae, and Staphylininae, but omitted the Vatesini. The
report also contained methods used in studying army ants and their
guests.
Rettenmeyer (1961) includes an excellent discussion and much in-
formation on 6 species of Vatesus he collected with army ants in the
^ Scientific Paper No. 3238, College of Agriculture, Washington State University. Work done under
Project 1802.
2 The critical reading of the manuscript by Dr. R. F. Harwood is appreciated. This study supported
in part by National Science Foundation Grant GB-5220.
The Pan-Pacific Entomologist 45 : 269-281. October 1969
270
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Canal Zone. Other than this report, all other papers on Vatesus are pri-
mirily taxonomic and give at most only host data.
This report was part of a study made in Panama during 1963, 1967,
ai d 1968 at the Smithsonian Tropical Research Institute, Canal Zone.
Vatesus Clypeatus (Wasmann) Species Complex
Although Borgmeier (1961) included V. clypeatus and V. gigas as
separate species in his key to Vatesus, Seevers (1965) considered V.
ch^peatus, V. gigas, and V. panamensis as a species complex since he
cculd find no means of differentiating them. The antennal characters
given hy Borgmeier do not hold true for large samples.
Rettenmeyer (1961) stated there was some doubt as to the taxonomic
status of V. panamensis and V. clypeatus, but included his data on the
V %tesus clypeatus complex by using V. panamensis as the species name
si (ice Barro Colorado Island was the species locality for V. panamensis.
V bile all our specimens were collected on Barro Colorado, we prefer to
uj e the species designation V. clypeatus to refer to this complex until
taxonomic difficulties are resolved.
Rettenmeyer (1961) collected 59 V. clypeatus {= panamensis) with
Eciton burchelli (Westwood), E. hamatum (Fabricius), E. vagans
(Oliver), E. dulcius crassinode Borgmeier, and possibly E. mexicanum
Rager. We collected 189 adult V. clypeatus from the following hosts: 16
colonies of E. burchelli (39), 24 colonies of E. hamatum (136), one col-
ony of E. dulcius crassinode (5), and 3 colonies of E. vagans (9). In
addition, 7 colonies of E. burchelli, 4 colonies of E. hamatum, 3 colonies
o E. dulcius crassinode, 3 colonies of E. vagans, and 4 colonies of E.
miexicanum had none. If V. clypeatus is found with E. mexicanum, it
n ust be quite rare. All but 2 of the beetles were collected from emigra-
tian columns. One of these 2 beetles was collected in a refuse deposit
column and the other was collected while digging out a subterranean
bivouac of E, dulcius crassinode.
We found Vatesus clypeatus in emigration columns only when brood
was being carried (Fig. 6) . They usually appeared after brood had been
carried for about 20 minutes and ant traffic was heavy. A few Vatesus
were collected after the queen had passed and the emigration was nearly
over. Only one Vatesus was collected at the end of the emigration after
all the ants had passed, and it appeared slightly injured. The inquilines
a I ways ran in the center of the column. They were never seen to deviate
f ] om the trail like other guests such as the staphylinid T etradonia mar-
ginalis Reichensperger (Akre and Rettenmeyer, 1968) . None was ob-
OCTOBER 1969] AKRE & TORGERSON VATESUS BEHAVIOR
271
served riding on the ants or brood. Whenever an emigration column
was disrupted by collecting, the ants would pile the booty and brood in
clusters under leaves or in concealed crevices. Vatesus were observed
buried in these piles of brood at least 8 times. During these periods the
beetles were ignored by the ants.
Vatesus spp. have functional wings and have been collected in quan-
tity at light (Borgmeier 1961) . Although we ran ultraviolet and incandes-
cent lights and a malaise trap for a total of 8 months, no Vatesus cly-
peatus was collected. While this beetle has functional wings, all 180
specimens examined had these wings broken off at lines of weakness
anterior to tbe distal margins of the elytra.
Experiments revealed the antennae of V. clypeatus are the sites of trail
chemoreception. Chemical trails were made using the methods given by
Akre (1964), and such trails were tested with unmodified beetles as well
as ones that had suspect chemosensory appendages removed. Ether had
no effect on trail following ability since control specimens readily
detected a trail as soon as they recovered from the anesthesia, so this
anesthetic was used to permit removal of appendages.
The antennae of V. clypeatus are 11 segmented; a long scape, short
pedicle, and a 9 segmented flagellum. Removal of both flagella resulted in
cessation of all movement. With one flagellum removed specimens were
very active and readily followed a trail. Lateral movements from one
side of the trail to the other, compensating for the loss of one appendage,
were exaggerated and suggestive of klinotactic orientation. The succes-
sive removal of flagellomeres revealed that speciments were able to sense
a trail with 2 flagellomeres remaining on each antenna. None of 4 speci-
mens with only the basal flagellomere were able to sense a trail, but 3
specimens with the 2 basal flagellomeres readily followed a trail 12-24
hours after amputation.
When following a trail Vatesus exhibits rapid lateral antennal move-
ments, but the antennae do not appear to touch the substrate. Distinct
trichoid sensillae are located on the flagellomeres; several on the ter-
minal segment and 3 on the distilateral margin of each remaining flagel-
lomere. These hairs are probably the site of chemoreception.
The possession of chemosensory ability on nearly all flagel-
lomeres would seem to be of distinct advantage if specimens frequently
lose parts of their antennae from being attacked by the ants. However,
examination of many specimens did not reveal missing flagellomeres as is
frequently found in the myrmeeophilous thysanuran, T richatelura manni
(Caudell) .
272
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Three Vatesus adults were kept alive in laboratory nests with their
host ants for 22 days in 1963 and 12 Vatesus were kept alive for periods
oi 10-15 days in 1967 and 1968, other V. clypeatus were kept alive for
slorter periods. During these times the Vatesus were observed through
a broad field binocular microscope at least one hour daily.
The ants under observation were fed booty and brood from other army
aiit colonies. The beetles were frequently observed feeding on this food.
hile they were able to chew holes in dead army ant brood with great
difficulty, they were usually unable to cut holes in booty without an
assist from the ants. Four Vatesus had been in a nest without food for
one day when some wasp (Polybiinae) larvae were placed in the nest.
T ie staphylinids immediately crawled on top of the larvae and the
mouth parts of the beetles moved every rapidly over the surface of the
larvae (strigulation) . Several tried to chew the larvae, but no beetle
was seen to cut a hole in the tough and pliable integument. The Vatesus
W3re constantly chased by the ants also feeding upon the larvae. Other
V %tesus ate dead E. hamatum, E. burchelli, E. vagans, E. dulcius cras-
si’iode, and E, inexicaiium brood. The adult Vatesus were never seen
k lling live brood as has been observed with other myrmecophiles ( Akre
and Rettenmeyer, 1966; Akre, 1968). More frequently than eating the
b ood themselves, Vatesus fed on juices oozing from holes cut in the lar-
vae by the ants or by other guests such as histerids. Frequently the
beetles attempted to feed on the brood while the ants were feeding but
were always chased away. The ants were extremely intolerant of Vatesus
and frequently attacked them. When the workers first chased Vatesus,
tl e beetles took a very zigzag course and usually lost the attackers. If the
workers chased them farther than a few centimeters, the inquilines dug
down in the soil of the nest until only the tops of their pronota and elytra
w ere exposed. Although the ants bit at the beetles, they were unable to
g asp their smooth top surfaces and quit shortly.
In nests, the Vatesus spent most of the time near the brood, or near
bDoty that was supplied as food. The ants always made a compact pile of
ti e brood and then clustered on top of it. The staphylinids frequently
b aried themselves in the pile of brood or dug down in the soil under the
p le, remaining there for hours. It was assumed they were feeding, but
this was not observed; alternatively they might associate with brood to
receive booty or to ensure a colony would not leave without them. The
1 atesus were frequently attacked by ants standing on the brood as they
ran between the ants’ legs. However, only one Vatesus was seen killed
bj^ the ants. This staphylinid appeared moribund and an ant grasped
OCTOBER 1969] AKRE & TORGERSON VATESUS BEHAVIOR
273
one of its legs and turned it over. The workers soon cut off all the inqui-
line’s legs, rendering it helpless. The next day only bits of the pronotum
and elytra could be located.
The Vatesus spent many hours in the laboratory nests buried under
the soil with only the tops of their pronota and elytra exposed. In this
position they were impervious to attack by the ants. However, when a
worker passed a partially buried Vatesus, it sometimes sensed the beetle
and began to dig frantically in the soil. Soon nearby workers began dig-
ging, probably in response to alarm pheromone. If the digging got too
close, the Vatesus scurried away and dug down in another location.
Life Cycle
Rettenmeyer (1961) reported the first collections and identification of
Vatesus clypeatus {pauamensis) larvae (Fig. 1) from 2 colonies of E.
hamatum and one colony of E. dulcius crassinode. The latter may not
have been V. clypeatus as Vatesus goianus Borgmeier is also associated
with crassinode.
Table 1 lists data for 200 larvae of Vatesus clypeatus collected from 2
colonies of E. burchelli (9), 8 colonies of E. hamatum (167), and one
colony of E. vagans (24). Rettenmeyer (1961) collected nearly all his
larvae (over 70) at or near the end of the emigrations. We collected 48
larvae from various colonies after all the ants had passed or at least when
ant traffic was sparse at the end of emigrations. All the remaining larvae
(except 8 collected in refuse deposits) were collected throughout the
emigration after brood was being carried, with no larvae passing after
all the ants were gone. The entire emigration was observed for all col-
onies listed in Table 1 except Ec-151, E. vagans, where we observed only
the last 20 minutes.
Table 1 also shows the sex of most of the adults collected in these
colonies. Although some adults may have left the colony or died when
the colony was in statary, this total represents all adult Vatesus in the
colony when we watched it. However, one colony was not observed dur-
ing the entire emigration. Colony Ec-115 had 54 larvae but only one
adult female Vatesus. Other collections listed in Table 1 also indicate
that only a few females must have laid all the eggs.
Larvae collected in emigrations ran either in the center of the column
with ants running over and around them, in the center of the column
with frequent deviations to the edge, or on the edge of the column out of
the ants’ way. Many stopped and hid under leaves or in crevices if there
was any disturbance of the column. In marked contrast to their behavior
Cfn
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
FIG.4
Fig. 1. Larval Vatesus, mature and ready to pupate. Fig. 2. Prepupa. Fig. 3. An
0 der prepupa starting to pupate with the abdomen already emerging from the pre-
pLipal integument. Fig. 4. Pupa that was unable to shed the larval integument.
IiG. 5. Pupa. Fig. 6. Vatesus adult in emigration column of Ection hamatum.
i nder laboratory conditions, the ants paid no attention to the larvae re-
ardless of their position in the column. It was evident from observing
OCTOBER 1969] AKRE & TORGERSON VATESUS BEHAVIOR
275
Table 1. Vatesus clypeatus collected with Colonies.
Adults
Nomadic-Statary
Colony #
Species Male
Female Total
Larvae
Collected
Day
E -321
E. hamatum
9
4
6
12
end of emig.
1st. nomadic
Ec- 13
E. burchelli
2
1
8
7
end of emig.
4th. nomadic
Ec- 85
E. hamatum
?
3
refuse deposit
Statary
Ec-111
E. hamatum
5
3
8
29
5 refuse de-
posit, rest
scattered
1st. nomadic
Ec-115
E. hamatum
5
1
6
54
Scattered
1st. nomadic
Ec-122
E. hamatum
4
3
7
22
Scattered
throughout
1st. nomadic
Ec-134
E. burchelli
9
?
2
2
end of emig.
1st. nomadic
Ec-141
E. hamatum
1
?
2
8
Scattered 5,
3 at end of
1st. nomadic
emig.
Ec-146
E. hamatum
6
7
13
36
Scattered
1st. nomadic
Ec-151
E. vagans
4
1
5
24
end of emig.
Prob. 2 or 3rd
nomadic
Ec-166
E. hamatum
3
5
8
3
Scattered
Prob. 3 or 4th.
nomadic
larvae found at the end of emigrations that they sense and use the chemi-
cal trails of the ants as do the adult Vatesus (Rettenmeyer, 1961; Akre
and Rettenmeyer, 1968).
Adult Vatesus are extremely good trail followers, making them ideal
for bioassaying the length of duration of trails in the field. Specimens
were released on host trails 12-24 hours following their collection. Dur-
ing the intervening time they were without host contact.
Fourteen specimens were released a total of 8 times on host trails, but
only once did a specimen appear to sense the trail or follow it for any dis-
tance. The morning subsequent to the emigration of colony Ec-2, E.
hamatum, a specimen was released near the old bivouac site and con-
tinuously observed for 2.25 hours. During this time it moved 17 m along
the emigration trail. Little difficulty was encountered when the speci-
men moved on roots or lianas, but in leaf litter or at points where the
ants had formed an emigration bridge the specimen encountered con-
siderable difficulty sensing the trail.
A Vatesus released on the emigration trail of Ec-11, E. humatum, less
than 48 hours after the ants moved, chose to follow a trail freshly laid by
50-100 ants released a few hours earlier, rather than the emigration trail
which at times was 5-7 ants wide.
S76
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Whether specimens were released in darkness or during the day
ssemed to have no effect. A Vatesus released on an E. burchelli trail,
s bout 16 hours after the ants had moved, was observed for one hour with
red light. During this period it seemed to be following the trail about
10% of the time. In one hour it moved 178 cm from the release point. It
explored all roots and lianas lying in the path of the emigration route,
following those routes which most readily facilitated its passage even
t lough these features were not part of the emigration route.
Twenty minutes after the last ant of colony Ec-141 {E. hamatum) emi-
grated, a Vatesus was released 2.5 m from the old bivouac. Within 10
iiinutes the specimen covered 20 m of trail over large lianas and leaf
litter. We could detect no difference in the specimen’s ability to sense
tlis quite fresh trail on either surface. This same specimen was released
£t the identical location 10.5 hours later, but gave no indication of
f bility to follow the trail. A similar procedure was repeated after the
emigration of Ec-159, E. hamatum. Upon release the specimen initially
1 noved toward the old bivouac 4 times. Each time it was re-aspirated for
fear of losing it. The fourth time it moved 4.74 m in 3 minutes, in the
( lirection of the emigration appearing to have little difficulty sensing the
trail.
From these observations we conclude that only very fresh trails can
lie used by Vatesus in host finding. That trail following on old trails is
]iot a method of host finding is further substantiated by Schneirla’s ob-
servations (1933) that ant trails lack polarity.
Our collections indicate Vatesus lays eggs and the larvae develop dur-
ing the host’s statary phase. As mentioned above, we collected 8 larvae
from the refuse deposits of E. hamatum', three from colony Ec-85, 14
days after we had located the colony which was in statary at that time,
and the other 5 larvae from refuse deposits of Ec-111 the day before it
lecame nomadic. In all cases the larvae were collected 10-50 cm from
the base of the bivouac, running among the discarded cocoons and booty
efuse. However, the larvae probably do not pupate in this location. We
excavated 4 bivouac sites of statary colonies immediately after the ants
lecame nomadic. All soil and debris around the bivouac site and to a
depth of 10 cm was removed and placed in plastic bags. This was brought
lack to the laboratory and searched very carefully without finding any
Vatesus larvae or pupae, indicating the larvae leave with the ants on the
irst nomadic day.
Larvae of various sizes and developmental stages were found in all
colonies investigated indicating adults do not lay their eggs at the same
OCTOBER 1969 ] AKRE & TORGERSON VATESUS BEHAVIOR
277
40
35
NUMBER Of INDIVIDUALS
■ ■
_L_
JJ
^PUPA^
■ ■
.65
.95 1.05
HEAD CAPSULE WIDTH
115
nm
U5 U5 145
Fig. 7. Head capsule widths of 154 Vatesus larvae.
time or else one adult lays her eggs over an extended period of time.
Alternate explanations could include differential embryonic development
or larval growth. Figure 7 shows the head capsule widths of 154 larvae
fall into 3 major groups so that at least 3 instars are present. However,
the head capsule growth may slow down in later instars (Ludwig and
Abercrombie, 1940) and head measurement is not recommended for
determining instars because some instars may be left out (Gaines and
Campbell, 1935). Since at least some staphylinids have only 3 instars,
this indicates we may have collected all larval stages (Paulian, 1941).
An attempt was made to rear the larvae to maturity by placing them
in round plastic containers, 15 cm diameter, with saran screen lids and
plaster of Paris/charcoal bottoms. Sterile soil (heated for 30 minutes)
was placed over the plaster to a depth of ca 1 cm. No ants were placed
in these boxes because the ants always killed the larvae. Larvae were
fed army ant brood, live phorid maggots, and another myrmecophile,
the thysanuran, T lichatelura manni (Caudell). Most of the larvae did
not feed, and either changed into prepupae or died. However, a few
were voracious. Four larvae, placed in a box with E. hamatum brood,
began to feed immediately. Although the brood was still alive, one beetle
larva devoured 3 small ant larvae in rapid succession. In several days
nearly all the brood (about 50 small larvae) had disappeared and was
assumed eaten. Several other larvae fed on dead brood of various army
ants, but the larvae seemed to prefer to kill and eat T. manni. This silver-
fish was their most readily accepted food. The integument of the silver-
fish is quite soft and the larvae readily killed and ate even the largest
individuals. However, the larvae had difficulty catching T. manni unless
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
i:78
]»oth were placed in small containers. Therefore, the T. manni were in-
jured before being placed in the 15 cm containers to facilitate feeding.
^Anywhere from 3—10 larvae were placed in each container. A total of
79 were retained in this manner. Eleven of the larvae changed to pre-
jmpae within 4^11 days after their collection and 3 pupated within 5-8
days. Most of the remaining larvae died within 6-7 days but 4 were still
£ live and active 16 days after their capture.
Before changing into prepupae, the larvae dug into the soil and made
£ n earthen cell where they became quiet. The prepupae are described by
llettenmeyer (1961) as curved with the head and legs drawn close to-
gether (Figs. 2, 3). He also stated that no movement could be detected
in the prepupae when they were handled with forceps. However, all pre-
pupae we handled exhibited characteristic pupal movements and any that
failed to move were usually dead or dying. Although the larvae changed
into prepupae beneath the soil, the soil in our laboratory nests was not
deep enough or was of incorrect texture because the movements of the
])repupae uncovered most of them before they pupated and may have
c ontributed to their death.
Only 3 of our larvae pupated. One of these, under the soil in an
earthen cell, managed to cast off the prepupal skin completely. The other
Iwo prepupae were on the top of the soil and died without completely
ehedding the larval skin (Fig. 4) . The first pupa was subsequently
lulled by a predaceous mite and none of the pupae completed develop-
inent. Of these 3 pupae, 2 lived for 10 days in the pupal stage. Whether
i he pupal stage normally lasts this long is not known, but even if it lasted
lor only a few days the ants would leave the Vatesus behind.
We have only one record of a colony from which we collected larvae
ctill following the emigration on the fourth nomadic day. In this in-
stance only 7 larvae were collected. It appears that the Vatesus larvae
urobably follow the ants with a few larvae dropping out of the column
£ind pupating each day.
The pupa that was able to completely free itself from the larval skin
;s shown in Fig. 5. Careful examination of the 3 pupae revealed fully
ormed secondary wings. Since all adult beetles examined had the
vings broken off, it seems likely that the newly hatched Vatesus fly to
their host ants and then lose the tips of their wings.
Vatesus goianus Borgmeier
Although V. goianus was reported previously only from Brazil with
T. dulcius, we collected 3 specimens from an emigration colunm of E.
OCTOBER 1969] AKRE & TORGERSON VATESUS BEHAVIOR
279
dulcius crassinode on Barro Colorado Island that fit the description of V.
goianus well, and Seevers (Chicago) indicated they probably were that
species. In addition to the 3 adult V. goianus, 5 larval Vatesus were col-
lected at the end of the emigration after all the ants had passed. Since
both Vatesus clypeatus and V. goianus occur with E. dulcius crassinode,
the larvae may have been of either species. However, since the entire
emigration was watched and no adult V. clypeatus were collected; it is
probable the larvae are V. goianus. The larvae have not been examined
to reveal differences.
The behavior of adult V. goianus was similar to V. clypeatus in the
field and laboratory. The adults were picked out of the center of the
emigration column when brood was being carried. In laboratory nests
these staphylinids fed on the brood and booty provided the ants as food,
but spent most of their time in the nests partially buried in the soil.
All these V. goianus had lost the tips of their wings along lines of
weakness just like V. clypeatus.
Vatesus praedatorius Seevers
Seevers (1958) description and illustrations of V. praedatorius are in
error and identification cannot be made from his paper. Type specimens
were borrowed from the Snow Entomological Museum and identification
was made by using these specimens.
Twenty V. praedatorius were collected in a raid column of Labidus
praedator (F. Smith) on 26 June 1968. The raid column was found at
8:40 p.m. and at that time was 10-12 ants wide. All the ants were ap-
parently returning to the bivouac with booty. The column emerged from
a hole in the ground, moved over the top of the soil for about 3 meters,
and then disappeared into another hole in the ground. The column was
observed from 8:40-11:30 p.m. when it ceased abruptly. One moment
it was 10 ants wide and suddenly there were none. The Vatesus were
collected throughout this time running in the center of the column and 2
were collected at the very end after the ants had passed. One Vatesus was
collected riding on the posterior end of a piece of booty carried by an
intermediate worker.
Upon returning to the laboratory, the Vatesus were placed in a labora-
tory nest with their host ants. Eighteen were kept alive for 6 days, and
the remaining 2 for 11 days before preservation. During the period
they were kept in the laboratory, observations were made on their be-
havior for at least 2 hours daily.
The behavior of V. praedatorius was also similar to that of V. cly-
il80 THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
yeatus. However, these small Vatesus seemed to be constantly active in
tile nests and ran from one cluster of ants to another. If any did re-
main still they were on the top of the soil or in the tunnels dug by the
c nts. They did not bury themselves as did V. clypeatus.
The Vatesus were very active, feeding on booty collected along with
the ants and on dead E. hamatum brood furnished the ants as food.
^Although the beetles were observed feeding many times, we could not
(ietermine if they actually chewed holes in the brood and booty. The
mouth parts of the beetles strigulated over the surface of the food, and
several times the beetles were seen chewing with their mandibles on
booty and brood. However, the mandibles appeared to be quite ineffi-
cient and the beetles seemed to feed more on the juices and materials
c ozing out of the holes cut by the ants.
The host ants constantly attacked and chased V. praedatorius but were
never seen to be successful in capturing or killing a specimen.
All 20 specimens of V. praedatorius had functional secondary wings
end could probably fly, although none attempted to fly even when
dropped.
Discussion
These three species of Vatesus are closely integrated with their re-
spective host army ants. Even though they are attacked constantly in
ihe unnatural situation present in a laboratory nest, the beetles always
lemained close to the ant cluster and brood. All have the ability to use
] resh chemical trails laid by the army ants.
All 3 species are predaceous on army ant brood but in nature they prob-
c bly feed entirely upon the booty brought in by the ants. In either
situation they are heavily dependent upon the ants both to furnish the
food and to tear it open so it is readily available.
Although Vatesus are good trail followers (Akre and Rettenmeyer,
1968) , our observations indicate that adults apparently fly to their host
f.nts. All specimens of Vatesus clypeatus and V. goianus collected in the
field had their wings broken off along fold lines and could not fly. How-
ever, several pupae of V. clypeatus had fully developed wings indicating
that freshly emerged individuals have functional wings. These wings
must be broken off sometime after the Vatesus locate their host ants. A
similar case occurs in the Heleomyzidae (Diptera). The type series of
Lutomyia hemiptera (Curran) have all but the basal portion of the wing
lorn off (Gill, 1962). This mutilation of the wings apparently occurs
citer the flies locate the rodent burrows in which they live. The adult
Vatesus lay eggs during the statary phase of the ants’ cycle. When the
OCTOBER 1969] AKRE & TORGERSON — VATESUS BEHAVIOR
281
ants emigrate the nearly grown larvae move with the ants until they
pupate. Since Vatesus are excellent trail followers they should have no
difficulty in locating their host by species odor; especially Vatesus cly-
peatus that has several host species. V. goianus probably flies to find its
host since E. dulcius crassinode seems to have regular periods of no-
madic-statary phases as E. hamatum and E. burchelli. Lahidus praedator
does not seem to have regular cycles and V. praedatorius may be able to
complete its development without being left behind by the ants. How-
ever, it has functional wings at all times and should be able to locate its
host ants readily if it cannot complete its development while the ants
are statary.
Literature Cited
Akre, R. D. 1964. The behavior of insects associated with army ants with emphasis
on chemical trails. Ph.D. Dissertation. Kans. State Univ. 119 p.
1968. The behavior of Euxenister and Pulvinister, histerid beetles associated
with army ants (Coleoptera: Histeridae; Hymenoptera: Formicidae: Dory-
linae). Pan-Pac. Entomol., 44: 87-101.
Akre, R. D. and C. W. Rettenmeyer. 1966. Behavior of Staphylinidae associated
with army ants, (Formicidae: Ecitonini). J. Kans. Entomol. Soc., 39: 745-
782.
1968. Trail-following by guests of army ants (Hymenoptera: Formicidae: Eci-
tonini) . J. Kans. Entomol. Soc., 41 : 165-174.
Borgmeier, T. 1961. Beitrag zur Kenntnis der Ecitophilen Staphyliniden-Gattung
Vatesus Sharp, Nebst Beschreibung von Sieben Neuen Arten (Col. Staphy-
linidae). An. Acad. Brasil. Cienc., 33: 189-202.
Gaines, J. C. and F. L. Campbell. 1935. Dyar’s rule as related to the number of
instars of the corn earworm, Heliothis obsoleta (Fab.) collected in the
field. Ann. Entomol. Soc. Amer., 28 : 445-461.
Gill, G. D. 1962. The Heleomyzid flies of America North of Mexico (Diptera:
Heleomyzidae) . Proc. U. S. Nat. Mus., 113(3465) : 495-603.
Ludwig, D. and W. F. Abercrombie. 1940. The growth of the head capsule of the
Japanese beetle larva. Ann. Entomol. Soc. Amer., 33: 385-390.
Paulian, R. 1941. Les premiers etats des staphylinoidea (Coleoptera). Etude de
morphologic comparee, Mem. Mus. Hist. Natur. Paris, (N.S.) 15: 361 p.,
2 pis., 1367 figs.
Rettenmeyer, C. W. 1961. Arthropods associated with Neotropical army ants with
a review of the behavior of these ants (Arthropoda: Formicidae: Dory-
linae) . Ph.D. Dissertation. Univ. Kans. 605 p.
Schneirla, T. C. 1933. Studies on army ants in Panama. J. Comp. Psychol., 15:
267-299.
Servers, C. 1958. A revision of the Vatesini, a tribe of Neotropical myrmeco-
philes (Coleoptera, Staphylinidae). Rev. Brasil Entomol., 8: 181-202.
1965. The systematics, evloution, and zoogeography of staphylinid beetles
associated with army ants (Coleoptera: Staphylinidae). Fieldiana, 47: 139-
351.
282
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
h Crossbreeding Study of the Cedar Tree Borer, Semanotus
ligneus amplus, and the Fir Tree Borer, S. litigiosus
( Coleopter a : Cerambycidae )
Boyd E. Wickman
Pacific NW Forest and Range Exp. Sta.
Forest Serv. USDA, Corvallis, Oregon
During a study of the biology of the fir tree borer, Semanotus litigio-
siis (Casey), a laboratory technique for inducing mating and oviposition
\ as devised (Wickman, 1968). Also during this work, one male cedar
t ee borer, 5. ligneus amplus (Casey), was captured in flight on 9 May
1965. This presented an opportunity to check on Craighead’s (1921)
I eport that these two species do not crossmate.
The male cedar tree borer was placed with one unmated female fir
tree borer. They mated immediately and did so repeatedly for 4 days.
On 10 May, the fir tree borer laid 20 eggs; on 11 May, 25 eggs; and on
12 May, 2 eggs. On 21 May, egg hatch began, and all eggs successfully
1 atched by 24 May. The larvae were immediately placed on an artificial
media (Lyon and Flake, 1966) . Most of the larvae died within 2 weeks,
1 ut six individuals did molt once and survived for a month.
The success of the first crossmating prompted further studies to deter-
mine the ease of crossbreeding and the possibility of progeny complet-
i ag their development to viable adults.
Techniques
A plentiful supply of fir tree borers was available for the study from
i\4ndthrown white fir, Abies concolor (Gordon & Glendenning) Lindley,
near Hat Creek, California. Cedar tree borers were harder to find, but
one windthrown incense-cedar, Libocedrus decurrens Torrey, in the area
j)rovided adults for the study.
On 4 March 1966, unmated adults of both species were dissected from
their overwintering pupal chambers in the wood. They were collected in
individual gelatin capsules, brought to the laboratory, located at Berke-
ley, California, and held at 2° C. for 10 days until the test was started.
A male and female were placed on a moist blotter paper on the bottom
of a petri dish (in four dishes one male was given two females), and
overhead fluorescent lights and room temperature of 20° to 23° C. pro-
^ ided satisfactory conditions for mating and ovipositing.
The Pan-Pacific Entomologist 45: 282-285. October 1969
OCTOBER 1969] WICKMAN CEDAR AND FIR TREE BORERS
283
On 15 March 1966, 12 breeding tests were started in petri dishes
(Table 1) with reciprocal pairings and checks.
Eggs were also used from crosses and checks for cytological studies.
Newly deposited eggs were smashed, then stained and fixed with aceto-
orcein. Slides were studied at ca. 900 X .
Results
Mating took place immediately and continuously during the first day
in all dishes. Oviposition started 2 days later in the dishes with female
fir tree borers and continued for 11 days, with peak egg production oc-
curring around the 4th and 5th days. Cedar tree borer females started
oviposition 5 days after mating and continued for 9 days, with peak
production occurring the 3rd to 5th days. However, two female cedar
tree borers (one check and one cross) produced no eggs.
Larvae began hatching 8 days after oviposition and continued for 7
additional days. Egg hatch varied from 54 to 100% and was considered
acceptable under the artificial conditions of rearing.
Larvae were transferred to artificial media in new dishes the day they
hatched. They all started feeding immediately, usually under the media
next to the dish. Their growth appeared to be satisfctory on the media
diet, and early mortality was similar to that experienced in previous
rearings of natural progeny on the artificial media (Wickman, 1968).
The first 14 days after hatching, mortality was about 10% — 31 of 306
(Table 1) .
Then, on 14-16 April 1966, a heat wave with unseasonably high tem-
peratures occurred in Berkeley. The temperatures in the rearing room
rose to about 33° C. and remained that way over the weekend. By Mon-
day morning, 18 April, all larvae were dead except one second instar in
dish 12; it died on 2 May.
The cytological studies showed similar chromosomal formulae and
configurations for the two species, but the work did not produce clear-
cut results. We could not provide enough eggs from the breeding and
rearing portion of the study to determine the chromosome numbers with
certainty.
Discussion
The crossbreeding test must be considered partially successful, since
viable eggs were produced and mortality in the first larval instar was
twice as high in the checks as it was in the crossbred insects. However,
it is unfortunate that uncontrolled high temperatures apparently killed
all but one of the test insects.
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
244
Table 1. Matings of the cedar tree borer (CTB) and fir tree borer
(j’TB) in petri dishes, Berkeley, California, March— April 1966
Eggs
Oviposition
Eggs
Eclosion
Larval
ish and Mating
laidi
Start Finish
hatched
Start Finish
mortality^
Num-
Per-
Number
Dates
Percent
Dates
her
cent
CTB
FTB
39
3/20 - 3/24
57
3/31 - 4/2
2
10
CTB
FTB
0
— ■ — •
—
— —
—
—
o
FTB
CTB
80
3/17 - 3/22
74
3/27 - 4/4
2
4
i- >
FTB
CTB
50
3/17 - 3/23
57
3/27 - 3/31
2
10
5
CTB
CTB
0
— —
—
— —
— '
—
()
CTB
CTB
38
3/21 - 3/28
90
4/2 - 4/7
11
32
7
FTB
FTB
13
3/24 - 3/24
100
3/31 - 3/31
2
15
8
FTB
FTB
29
3/17 - 3/25
79
3/27 - 4/14
0
0
9
CTB
FTB
59
3/21 - 3/27
100
3/30 - 4/14
3
6
CTB
10
FTB
CTB
63
3/17 - 3/27
100
3/24 - 4/2
1
2
FTB
1 .
CTB
FTB
23
3/20 - 3/29
91
3/27 - 4/5
2
10
CTB
12
FTB
CTB
14
3/22 - 3/27
50
4/2 - 4/7
6
100
FTB
1 Includes eggs used for cytological work.
2 Mortality of first instar prior to heat wave.
The tests have shown that the two species will readily crossmate, pro-
duce eggs, and the eggs will hatch. In nature, both species fly and at-
tack in the same timber stand at the same time of year and accidental
crosses seem possible. If so, do progeny result? So far there has been
ro success in rearing crossbred larvae to adults. Perhaps they cannot
develop to adults; and this, combined with their host specificity for
r lating and oviposition sites, keeps the two species distinct.
Craighead’s failure to mate the two species during his tests of Hop-
lins’ host selection principle remains unexplained. They mated readily
r nder the conditions described. This mating may lead to further interest-
i ig work on host selection and speciation if viable progency can be pro-
c uced.
Cytological studies would be valuable for ascertaining the genetic in-
tegrity of the two species and their potential for hybridizing.
OCTOBER 1969] WILLIAMS A NEW MEXICAN SYNTROPIS
285
Acknowledgments
I wish to thank Dr. Gerald N. Lanier of the Forest Entomology and
Pathology Laboratory, Canada Department of Forestry, Calgary, Al-
berta, for giving instructions on cytological preparations and examina-
tions; and Dr. R. W. Stark of the Department of Entomology, Berkeley,
California, for providing facilities and suggestions.
Literature Cited
Craighead, F. C. 1921. Hopkins host selection principle as related to certain
cerambycid beetles. J. Agr. Res., 22: 189-220.
Lyon, R. L. and H. W. Flake, Jr. 1966. Rearing Douglas-fir tussock moth larvae
on synthetic media. J. Econ. Entomok, 59: 696-698.
WiCKMAN, B. E. 1968. The biology of the fir tree borer, Semanotus litigiosus (Co-
leoptera: Cerambycidae) , in California. Can. Entomok, 100: 208-220.
A New Species of Syntropis from Baja California Sur, Mexico
with Notes on its Biology
(Scorpionida; Vejovidae)
Stanley C. Williams
San Francisco State College, California
In 1900 Kraepelin described a new genus and species of scorpion
which he named Syntropis macrura based on a single specimen. This
genus was of special interest because it was characterized by the
presence of a single, unpaired inferior median keel on the metasoma,
instead of the paired condition generally found in the family. Over
the years this genus acquired more and more interest since its unique-
ness within the family Vejovidae became better recognized and since it
still remained known by only the type specimen.
During the summer of 1968 extensive field work in the Baja California
peninsula of Mexico revealed much information about the genus
Syntropis. Two parties, one led by Mont A. Cazier, the other by myself,
together collected several dozen Syntropis macrura and over 1,000 speci-
mens of a new species of Syntropis, here described and named Syntropis
longiunguis.
The Pan-Pacific Entomologist 45: 285-291. October 1969
236
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Figs. 1 and 2. Syntropis longiunguis Williams, holotype male. Fig. 1. Dorsal
V ew. Fig. 2. Ventral view.
Syntropis longiunguis Williams, new species
(Figs. 1, 2)
Diagnosis. — Moderate sized, light yellow species easily placed in genus Syntropis
hy single median keel on inferior surface of metasoma. Differs from S. macrura as
follows: uniform light yellow in color (not rusty) ; fingers of pedipalp distinctly
shorter and with subtle internal scalloping; palm swollen; telson not bulbous but
s rikingly elongate; pretarsal claws very elongate, outer claw of each pair distinctly
longer; unguicular spine reduced; distal tarsomere densely hirsute and lacking
cefinite row of short spines; inferior border of fixed cheliceral finger with several
riinute denticles.
Holotype Male. — Coloration. — Carapace, mesosoma, metasoma and pedipalps of
i niform pale yellow ; walking legs and pectines similar but lighter ; only contrasting
OCTOBER 1969] WILLIAMS A NEW MEXICAN SYNTROPIS
287
color markings are; eyes black, teeth of chelicerae and chela reddish-brown, aculeus
dark reddish-brown, tips of pretarsal claws light reddish-brown. Most of cuticle
lustrous.
Carapace . — Anterior margin straight and narrow, set with three pairs of erect
reddish bristles. Lateral eyes three per group, these very tiny. Median eyes on
raised ocular tubercule; one pair of stout bristles on ocular tubercule medial and
posterior to median eyes; diad % carapace width at that point, and completely
on anterior half of carapace. Carapace surface with large granules; median groove
narrow and shallow anterior to median eyes, continues over ocular tubercule, be-
comes deeper posteriorly, ends as relatively deep pit near posterior border of cara-
pace. Carapace wide posteriorly and conspicuously narrows near anterior border.
Mesosoma . — All dorsal plates densely covered by large granules mostly located
on posterior part of each tergite, median and anterior areas relatively smooth;
subtle suggestion of obsolescent median keel on tergites 3 to 7; tergite 7 with two
pairs of distinct lateral keels set with large dentate to serrate granules; lateral
margin of segment 7 abruptly flattened horizontally and set with large dentate gran-
ules. Sternites relatively smooth, agranular and transparent; one pair of poorly de-
veloped lateral keels on last sternite, these smooth to crenulate. Stigma long and
slit-like.
Metasoma . — All dorsal and dorsolateral keels complete and well developed except
dorsals absent on segment V ; keels deeply serrate on segments I to IV and each
terminates posteriorly in enlarged spine; segment V with dorsolateral keels finely
serrate over anterior half but gradually becomes obsolete on posterior half; dorsal
keels gradually converge posteriorly on segments I to III. Lateral keels deeply den-
tate along length of segment I; keels represented as four granules along posterior %
of II and posterior % of III; absent on IV; crenulate on anterior % of segment V.
Inferior lateral keels present and complete on all segments, irregularly crenulate on
I to III, irregularly serrate on IV and V. Inferior median keel unpaired where
present; keel absent on segments I and II, smooth to obsolete on III, smooth to cren-
ulate on IV, irregularly serrate on V and not extending to posterior margin of
segment. Inferior intercarinal space of segment V with about eight pairs of stout
reddish hairs, surface very finely granular. Segment V with very conspicuous
gradual, posterior taper in width.
Telson . — Entire vesicle long slender and lance-shaped, with no bulbous appear-
ance; vesicle elongate, indistinguishable from aculeus except by beginning of red-
dish-brown pigmentation of aculeus. Ventral surface of vesicle with about 10 to 12
pairs of moderately long reddish hairs; entire surface of vesicle agranular, smooth,
lustrous ; no subaculear tubercule. Base of aculeus with lateral pair of finely barbed
strips.
Pectines . — Long and thick ; middle lamellae numerous, most subcircular, but prox-
imally several become more columnar; fulcra subcircular to triangular; 31/30 pec-
tinal teeth; teeth long; entire inferior surface of pectines (except teeth) densely
covered with short light brown hairs, each fulcrum with 10 to 15 of these hairs.
Genital Operculum . — Completely divided longitudinally; large distinct genital
papillae visible externally.
Sternum . — Distinctly pentagonal, length approximates width, with deep trian-
gular pit posteriorly.
Chelicerae . — Inferior border of movable finger completely lacking denticles,
border may appear subtly crenulate under high magnification, but essentially
238
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
siaooth. Inferior border of fixed finger with row of three or four small unpigmented
g anules.
Pedipalps . — Hand swollen, palm keeled, fingers of moderate length. Superior
a: id secondary superior keels set with rounded granules; inner marginal keel es-
sentially smooth; inferior keels at base of movable finger with dentate granules.
Carapace slightly shorter than movable finger but distinctly longer than fixed
fi ager. Grasping edge of fingers armed with one long continuous row of sharp ser-
iTte denticles, each row subdivided by the occurrence of eight to nine distinctly
hrger and spaced denticles. Principle row of denticles flanked inwardly by eight to
nine lateral denticles, no such denticles flank outer side of principal row. Fixed
fi Qger distinctly extends distally further than movable finger, each finger terminates
ir small, sharp, cone-shaped denticle. Both fingers with slight scalloping of their
g asping edges; teeth not extending to proximal end of either finger and small
o )en space remains in proximal end of chela when fingers closed.
W alking Legs . — Tarsomeres with abundant, long reddish hairs, most conspicu-
ous on distal one. Pretarsal claws very elongate and not symmetrical in relation to
length, outer claw always distinctly longer than inner claw (2.0 and 1.5 mm respec-
tiv'ely on hind walking legs) ; unguicular spine reduced in size; unguicular lobe very
elongate and densely set with long stout reddish hairs. Pedal spurs paired, elongate,
a id tipped with red pigment, but not serrate.
Allotype Female. — Essentially as holotype in coloration and morphology with
following exceptions: slightly smaller in total length; slightly wider in carapace
width; chela slightly longer and with distinctly more elongate fixed finger; pectines
siaaller and with slightly fewer teeth (27/26 instead of 31/30) ; no genital papillae;
genital operculum not completely divided longitudinally.
Standard Measurements . — Table 1.
Study of 690 paratopotypes (255 males, 325 females, 110 early instar
E/mphs) indicated little variation from the descriptions of the holotype
a id allotype. Males varied in total length from 14 to 60 millimeters while
f(;males varied from 14 to 66 millimeters. Adult males and females
tf;nded to have about equivalent hody sizes and proportions, the females,
h owever, had carapace slightly wider and longer, and with chela slightly
n lore elongate. Pectine tooth count varied from 24 to 31 in females and
f; om 28 to 35 in males. Many very young juveniles (probably second
a id third stage) were encountered in the samples and these were similar
to adults except for their smaller body size and more tan appearance.
\ ery little sexual dimorphism occurred. Adult and older subadult male
iiistars had genital papillae distinct while females lacked these organs.
Males had genital operculum more completely divided longitudinally and
h Eld larger pectines. Male pectines tended to have more teeth than female
pectines, but counts often overlapped, adult male pectinal teeth were
longer than those of the female.
The holotype male, allotype and 690 paratopotypes were collected on
tlie SAND DUNES OF SaN AnGEL, 13 MILES WEST OF SaN IgNACIO, BaJA
California Sur, Mexico, 27 June 1968 by S. C. Williams, M. A. Cazier
OCTOBER 1969 ] WILLIAMS A NEW MEXICAN SYNTROPIS
289
Table 1. Measurements (in millimeters) of Syntropis longiunguis
Williams, new species, holotype and allotype.
Holotype
(male)
Allotype
(female)
Total length
58.7
58.1
Carapace, length
7.2
7.8
width (at median eyes)
5.8
6.3
Mesosoma, length
13.0
13.0
Metasoma, length
28.1
27.3
segment I (length/width)
3.8/3.8
3.7/3.9
segment II (length/ width)
4.4/3.7
4.3/3.8
segment III (length/width)
4.5/3.6
4.5/3.7
segment IV (length/width)
5.7/3.5
5.6/3.5
segment V (length/width)
9.7/3.0
9.2/3.0
Telson, length
10.4
10.0
Vesicle (length/width)
7.4/1.8
6.5/2.0
depth
1.6
1.8
Aculeus, length
3.0
3.5
Pedipalp
Humerus (length/width)
5.6/2.0
5.7/2.1
Brachium (length/width)
5.9/2.8
6.3/2.8
Chela (length/width)
11.3/3.0
11.8/2.9
depth
3.8
4.0
movable finger, length
7.5
7.7
fixed finger, length
5.8
6.6
Pectines
teeth (left/right)
31/30
27/26
and party. The holotype and allotype are permanently deposited in the
California Academy of Sciences.
This species is named “longiunguis” because of its highly modified
pretarsus in which the ungues are conspicuously elongate and the ungui-
cular spine is greatly reduced.
In addition to the holotype, allotype and 690 paratopotypes, an addi-
tional 793 paratypes were available for study. All specimens were col-
lected in the following three localities in Baj a California Sur, Mexico ; 3
miles south of Rancho Tablon, elevation 1,500 feet, 23 June 1968 (S. C.
Williams, M. A. Cazier and party) , 29 males, 40 females; 5 miles west of
San Angel, 28 June 1968 (S. C. Williams, M. A. Cazier and party), 69
males, 92 females, 37 young juveniles; approximately 5 miles north of
La Laguna on east shore of Laguna de San Ignacio, 29 June 1968 (S. C.
Williams, M. A. Cazier and party), 229 males, 216 females, 81 young
juveniles.
250
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Comment. — This species was found along the west coast of the Baja
California peninsula from Rancho Tablon to near La Laguna. In this
a rea it showed definite preference for habitats characterized by unpacked
dime type soils. These habitats, appeared to be secondarily created by
sand transported in by the prevailing winds. Throughout this area this
s md takes the form of totally unstabilized and moving sand dunes. These
d unes are large and very extensive in places such as San Angel but may
te very small and relatively isolated in areas like Rancho Tablon. In the
h abitats of occurrence this species was conspicuously the numerically
dominant scorpion. It had a nervous and responsive temperament, was
intolerant to other scorpions of the same or different species and showed
vary aggressive behavior toward any local disturbance in the nearby
sand.
The morphological modifications of this species for sand dwelling are
s riking. The pretarsal claws are extremely elongate while the unguic-
rlar spine is very reduced in size. The entire surface of the last tarso-
riere is densely clothed in long reddish hairs which permit rapid locomo-
tion on loose sand without lack of traction. This species is even capable
cf traveling up the steep leeward side of large sand dunes at a fast speed.
The streamlining of the metasoma and telson is perhaps an adaptation
f ir escaping from being buried in the loose sand.
At night the surface of the sand dunes were abundantly covered with
individuals of this species. The population was densest on sloping re-
§ ions of the dunes rather than on the crests or basins. The scorpions came
c ut on the sand surface at dusk and sat perfectly still. A moving speci-
rien was a rare observation. When disturbed on the side of a dune an
iidividual would escape by gliding rapidly over the sand and coming to
1 3st in a lower location about 10 to 15 feet away. They always escaped
i 1 a downward direction.
Most of the individuals appeared to be sexually mature, and males
spproximated females in number. Very young juvenile instars were also
s bundant. These were probably second and third instar nymphs. Inter-
nediate aged subadults were rare on the surface of the dune. Dissection
i idicated that the adult females had already given birth to their young
for the season. The great similarity of the reproductive systems of these
c issected females indicated that time of birth must be synchronized to
some extent within the population and that probably only a single litter
is produced by a female during the year.
Based on the single inferior median keel of the metasoma, Syntropis
j iacrura would be the closest known relative of this species. However,
tbese two species of Syntropis do not appear to be closely related, and
OCTOBER 1969] ALEXANDER — A NEW CRANE FLY
291
even appear superficially very unrelated. In secondary morphological
adaptation, ecology and behavior, S. longiunguis appears more like Ve-
jovis mesaensis (Stahnke) than any other North American scorpion.
These two species should, therefore, be considered as ecological counter-
parts, S. longiunguis occurring in dune communities of the Viscaino
desert, V . mesaensis occurring in dune communities of the Sonoran and
Colorado deserts. Syntropis macrura on the other hand appears more
like Ve jovis minckleyi Williams in secondary morphological adaptation,
ecology and behavior, V. minckleyi occurring in talus deposits and along
rocky cliffs in Coahuila while S. macrura lives in similar habitats in the
Sierra de la Giganta.
Acknowledgments. — The field and laboratory research involved in
the discovery, collection and study of this species was supported by the
systematics branch of the National Science F oundation by research grant
number GB 7679 and by a Faculty Research Leave from San Francisco
State College. Thanks are due C. F. Williams for clerical assistance in
the preparation of this manuscript and to Mont A. Cazier for cooperative
aid in the field.
A New Species of Crane Fly Associated with the Plant Genus
Lopezia in Mexico
(Diptera; Tipulidae)
Charles P. Alexander
Amherst, Massachusetts
During the course of his studies on the pollination of plants of the
onagraceous genus Lopezia in Mexico, Dr. Dennis E. Breedlove collected
about a dozen species of Tipulidae of which one proved to represent a
new species. In order to make the name available for his use the fly is
described herewith. The type of the novelty, as well as the other crane
flies collected, are preserved in the California Academy of Sciences. I
am indebted to Dr. Paul H. Arnaud and Dr. Breedlove for the privilege
of examining this interesting material.
Tipula (Trichotipula) breedlovei Alexander, new species
Mesonotal praescutum with four nearly confluent grayish brown stripes, posterior
sclerites of notum variegated brown and yellow, pleura chiefly light gray; wings
light brown, stigma and costal border slightly darker; outer wing cells with strong
The Pan-Pacific Entomologist 45: 291-293. October 1969
292
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Fig. 1, Tipula (Trichotipula) breedlovei Alexander. Details of male hypopyg-
iiim. Symbols: d — dististyles; t — ninth tergite.
black trichia, abundant in cell Rs; abdominal tergites brown, bases more yellowed,
sternites brown, yellow laterally; male hypopygium with tergal lobes bearing
abundant spinoid setae; outer margin of beak of inner dististyle with about a dozen
long powerful yellow bristles, face of style near base with a slender darkened rod
a ad a broad-based blackened plate that narrows into an acute spine.
Male. — Length about 13 mm; wing 13 mm .
Frontal prolongation of head obscure yellow, nasus short; palpi broken. Antennae
Y ith proximal three segments yellow, remainder broken. Head brownish gray above,
front, orbits and sides of the low vertical tubercle obscure yellow.
Pronotal scutum brownish gray, scutellum and pretergites yellow. Mesonotal
praescutum with four nearly confluent grayish-brown stripes, the very narrow inter-
s >aces darker brown; scutal lobes brownish gray; scutellum brownish yellow, para-
seutella clearer yellow; mediotergite light yellowish brown, yellowed laterally, pleu-
rotergite clear light yellow, its ventral border light brown. Pleura chiefly light gray,
propleura restrictedly brown; dorsopleural membrane broadly clear light yellow,
nietapleural region more obscured. Halteres with stem brownish yellow, base nar-
rowly yellowed, knobs broken. Legs with coxae grayish yellow; trochanters yellow;
femora yellow, tips infuscated; tibiae obscure yellow, the outer parts and the tarsi
broken. Wings light brown, stigma, prearcular field and cells C and Sc slightly
darker; veins brown. Strong black macrotrichia in outer ends of cells Rs to Mi,
abundant in cell Rs; a few similar trichia in proximal half of stigma. Venation:
Sci ending about opposite one-third Rs, Sc 2 near its tip; petiole of cell Mi nearly
t vice m; m-cu on vein Mi some distance beyond base, the latter subequal in extent
to Ma + i.
Abdominal tergites chiefly brown, bases more yellowed; sternites brown, yellow
laterally; hypopygium light yellow. Male hypopygium (fig. 1) with posterior border
of tergite, t, deeply notched, the resulting lobes with abundant black spinoid setae,
OCTOBER 1969] BLACKWELDER NATURE OF TAXONOMIC DATA
293
those at midline long and slender, the most lateral ones small and peglike. Outer
dististyle, d, relatively small, about equal in length to the inner style, setae pale;
inner style on outer margin of the beak with about 20 long powerful yellow spinoid
bristles, the remaining setae shorter, black; face of style in the position of the lower
beak bearing a long slender darkened rod, above this with a broad blackened plate
that narrows to an acute spine. Aedeagus very long and stout, as in the subgenus.
Holotype male (broken), taken about 10 road miles northeast of
Madera, Chihuahua, Mexico, at 7300 feet, 29 September 1966, on
flowers of Lopezia gracilis (Onagraceae) by D. E. Breedlove.
I am pleased to name this distinct fly for the collector. Dr. Dennis E.
Breedlove. Approximately 35 regional species of the subgenus now are
known, as discussed in the Crane flies of California (Alexander, C. P.,
Bulletin of the California Insect Survey, volume 8: 56-60, 1967) and in
various other papers that concern the species of Mexico. All of these
differ from the present fly in hypopygial characters, most evidently in
the long powerful bristles on the beak of the inner dististyle, a unique
character in this subgenus but occurring in various members of the sub-
genus Indotipula Edwards in the Oriental and eastern Palaearctic regions.
The Nature of Taxonomic Data
Richard E. Blackwelder
Southern Illinois University, Carbondale
The data of taxonomy is a subject that would scarcely have been dis-
cussed even a few decades ago. There was no need to talk about it,
because every taxonomist knew what data were relevant in his field, or
thought he did, and the only thing to discuss was whether or not he was
right — whether there might be a better choice of data in that particular
case.
The idea of discussing taxonomic data in general occurred to taxono-
mists only as a result of the claims and challenges of some non-taxono-
mists, who thought they had discovered serious faults in the taxonomic
system and great failures of taxonomists. These challenges were a surprise
to most taxonomists, who were so busy with their endless work that they
had never stopped to think about their data in any theoretical sense.
Many of them never have understood what all the shouting is about,
because they’re generally unable to use the supposedly “new” data urged
on them by the outsiders, and they’ve seen only occasional need to do so.
The Pan-Pacific Entomologist 45: 293-303. October 1969
294
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Zoological and also botanical classifications are based entirely on data
- -data about the attributes of the organisms. This general word attribute
is used rather than character to emphasize that “characters” don’t have to
I:e structural (what in entomological circles is usually called morphologi-
cal) . To be useful in classification, the data must be comparative in na-
t are — either the organisms agree in possessing the attribute, or they differ
i i its possession or extent. The differences may be such as to attract
c ur attention through any one of the senses, but most commonly through
c irect vision. On the other hand, the difference may be such as to appear
cnly after the application of special techniques — microscopy, experi-
rientation, analysis of measurements, chemical analysis, instrumental re-
cording and interpretation of invisible attributes, and so on.
It should be clear that any fact that can be known about an individual
era kind-of-organism is potentially comparative. It can at some future
t me be compared with the corresponding fact about some other individ-
i al or kind. The corresponding features will then be found to be the
same or different, to unite or separate the two individuals or groups.
Not all comparative data will be taxonomic. At the species level it
c ften will not be. Knowledge of the biology of the organisms will enable
tie taxonomist to eliminate sex-likened differences, pathological differ-
ences, differences due to age, developmental stage, caste, and so on. But
tele remaining comparative features are automatically taxonomic. They
£ re the things in which in taxa are alike or different.
Not only are all taxonomic data comparative in nature, by definition,
but, turning this around, all comparative data may be taxonomic, and
ttiey must all be studied to determine whether they are variation within
tlie species or diversity among kinds. (By study, is not meant anything
^ery formal; background knowledge usually makes it unnecessary to
£ pend much time on this. )
Taxonomy, especially in entomology, was once almost exclusively
‘ morphological” ; it used principally data of structure, especially external
structure. For many years taxonomy has been making use also of other
binds of data whenever they are available. In many parts of the Animal
1 iingdom these newer data have not been needed to any great extent and
£o have been little used. But taxonomy can use any comparative data,
£ nd all comparative data are automatically of taxonomic interest.
The use of comparative data isn’t new, even if we now have some sorts
of comparative data that weren’t available at all a few decades ago. But
^ ery few sorts of data available today in any usable quantity are entirely
new. Genetic data were used long ago in the form of breeding compatibil-
ity. Biochemical data of a rough sort were used more than half a
OCTOBER 1969] BLACKWELDER NATURE OF TAXONOMIC DATA
295
century ago. X-ray observation and infrared spectroscopy had to await
invention of the machines, but they have not yet produced data in suf-
ficient quantity to be really useful in classification.
There are four words basic to this discussion. They are : comparison,
resemblance, difference, and correspondence.
Classification implies that there are “classes” of things. If we start
with individual things and compare one with another, we can unite sim-
ilar ones in classes based at first thought on similarities. If we start with
many things at once and divide them into classes and subclasses, we will
have used differences as our first criterion. These two approaches are the
two basic methods of classifying. Both of them are based on comparison
of the objects or their attributes. In the first we compare to find what
there is that’s similar. In the second we compare to find what there is
that’s different. The key word and concept is comparison.
To group, or classify, things there must also be some similarities
among them. This is resemblance. We know that resemblance may be
produced by several processes or circumstances (heredity, convergence,
or chance), but it’s the existence of resemblance that allows comparison
to recognize groups, and it’s the existence at the same time of differences
between organisms that allows us to classify these groups by comparing
them among themselves.
Two things can be compared even if they have nothing in common,
but there may seem to be no purpose to the comparison. If you had a
bottle in which there was a big cerambycid beetle, would you find it use-
ful or even entertaining to compare the bottle with the beetle? It would
be pointless, because there’s no correspondence of features to make the
comparison meaningful. There must be correspondence before we can
classify by comparing. There are various ways in which things can cor-
respond: in structure, in shape, in position, in embryological develop-
ment, in function, and so on. All classification is based on the belief
that we can find useful correspondence, what we call homology, in a com-
bination of structure and development and location. We believe that this
correspondence is due to the fact that the animals had a common ances-
try.
We’ve been considering taxonomic data so far as if we had already dis-
cussed what this expression means. Let’s go hack now and establish the
relationship between taxonomic data (that is, taxonomic features or
taxonomic evidence) and the diversity of animals.
The diversity of animals is simply endless. It would be impossible to
catalog what we know of it completely because that is even now increas-
ing steadily. We tend to forget the extent of the diversity, because we
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
2^6
can make our distinctions with reference to only a small part of it. The
minute we start to talk about the extent of diversity, we get into trouble,
be cause there’s no effective way to classify or describe it. F or example,
W3 think of the digestive tract as being quite similar in different groups
ol insects, differing in details of size, shape, and such things, but show-
ing no diversity within a family or genus, and therefore serving as a
distinctive character only at high levels. We may assume that this is
ahrays true. We forget that there are groups of animals in which the
individuals in one interconnected colony may differ in possessing a di-
gestive tract or not. Diversity exists at all levels, and there is no theoreti-
Cfl way to tell what level will be exemplified by any particular sort of
f e ature.
What we, as taxonomists, are interested in is that some of the diversity
is useful to us and some is not. The part that is not useful can’t be f orgot-
te n, however, as it must be recognized in order to be avoided. The useful
part is what we just called comparative data. When we compare, the sim-
ilarities allow us to group like things; the differences allow us to separate
uidike things. But because taxonomists seldom are interested in group-
irg or distinguishing individuals, the features must be ones that don’t
vary within the species (or other group) .
That individual variation is not a simple matter, a single sort of diver-
si ly, is suggested by this abbreviated list of the causes of diversity within
tl e species :
1) Normal developmental processes
2) Sexuality
3) Colonial division of labor
4) Genetic recombination and ploidy
5) Mutations
6) Response to environment
7) Combinations of these
If your reading takes you outside of the field of entomology (and I
hope it does), you may sometime note that G. G. Simpson once wrote
tl at one cannot classify individuals, but only species. There’s a sense in
which this is true; for example, if all the species are known, as in the
h gher vertebrates, then one classifies only species and groups of species.
In most of the Animal Kingdom, however, we must start with individ-
uals and determine which belong to one species and which to another.
^ ' e would like to do this by knowing enough about them to say that they
are the members of an interbreeding population and thus are genetically
related. It’s extremely rare that we can do this, because a species is not
a population but a complex of local populations which we never can
actually see.
OCTOBER 1969] BLACKWELDER — NATURE OF TAXONOMIC DATA
297
What the taxonomist does is to assign to one species all the specimens
which his comparative studies lead him to believe are related because
they are as alike as their offspring or their hereditary relatives within a
few generations. Many biologists find this definition unsatisfying, but
it’s the actual basis of most taxonomic work. It has been worded even
more simply, that a species consists of whatever individuals the taxono-
mist thinks belong to it, but this fails to make clear that he thinks this
because he believes that their similarities are due to the fact that they are
genetically related. He puts them together because of the comparative
features, but with the assumption that these reflect actual kinship.
There have been several attempts in the past 30 years to change the
philosophical or methodological basis of taxonomy. There has been
much discussion of these, but it’s questionable whether they have had
much effect on the practicing taxonomist. The first of these, and the
one most bitterly fought over, was the claim that taxonomy is a branch
of the study of evolution; that its principal purpose is to unravel the
course of evolution ; and that its classifications are based on phylogeny.
This is The New Systematics as defined by Mayr; Huxley and his col-
laborators in the original New Systematics book did not make this claim.
Nowadays, no biologist denies that animals have undergone changes,
that these changes coming in succession have produced new kinds, that
this “evolution” has produced all the kinds we now know, and that the
history of the kinds is phylogeny, the record of the successive changes.
No entomologist uses phylogeny to a significant extent in his taxonomy
because there isn’t enough known about the phylogeny of insects. It is
curious, however, that we come to think of phylogeny as something more
than a theoretical concept. It’s quite possible that if we knew the phylog-
eny of every species, we could classify them on this basis. (It’s more
likely that we’d find it unnecessary or even impossible to classify this
way.) But we don’t know the phylogeny of even one species. We have
merely worked out a scheme, in such a case as the horse, which we think
may have been its phylogeny, and which we hope was, to bolster our ego.
Where does this supposedly phylogenetic data come from? There are
two sources: First, the sequence of fossils in successive layers of rocks
gives a direct clue to succession of these forms in time. But this is highly
subjective, and it won’t work at all until taxonomic comparisons of the
specimens convinces the paleontologist that the successive forms are re-
lated.
If you found a bone in a relatively old stratum and a spiral shell in
a younger stratum in the same sequence, as you very likely could do, you
2^8
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Wouldn’t conclude that the snail evolved from the horse. You’d know that
tl ey are unrelated and not comparable.
The second source is direct comparison of the animals, or their fossil
remains, in the usually subjective taxonomic manner, to see which one
has features that could have been derived from corresponding features
o. the other. In either case the result is not the phylogeny of the animal
but a supposed phylogeny. If the work has been done well, it may be a
very convineing supposed phylogeny.
But, if the phylogenies are based essentially on comparative data, then
a classification supposedly based on phylogeny is actually no different
ii basis from one based directly on comparative data. There simply are
no real distinct data of phylogeny, although the age of a fossil and its
p ace in the series of rocks is a datum of great value. (It’s another sort
o : comparative datum.)
Thus, the only real phylogenetic data are derived either directly or in-
directly from comparison of features of specimens. You’ll probably
n 3ver meet a person who can show you a classification hosed on phylog-
e iy. In a few groups, well represented in the fossil record, you may see
some classifications that were influenced by speculations on phylogeny.
4 hus, this attempt to change taxonomy has not been successful.
The most recent effort to change taxonomy is the one that is least
a ctively pressed. Strangely enough, it’s the one that has the most to offer
to taxonomy. This is the belief by bioehemists that their data on the kinds
of animals should be used by taxonomists. Much of this data is truly com-
p arative ; as such it is unquestionably of taxonomic interest.
The only difficulty is that some biochemists seem to think that all
classification should be re-done on the basis of electrophoresis patterns
cr serological reactions. This is unrealistic. The data of biochemistry
3 re automatically stored by taxonomy, but they are used in classification,
1 ke most other kinds of data, only when they are needed. So far, this
r eed has been shown in very few cases, because biochemical data almost
i ivariably confirm prior taxonomic conclusions.
Unfortunately, when the biochemical data do not confirm, the bio-
chemists usually assume that their data must be wrong. For example.
Eta conference on bioehemical taxonomy at Lawrence, Kansas, a couple
c f years ago, most speakers compared their biochemical results with the
standard taxonomic classifications. If the results agreed, they were grati-
fied, but if the results differed from the accepted classifications, they
E Iways assumed that their results were wrong. If the results always agree,
tiren we don’t need them. If we don’t believe them when they disagree,
then they’re useless.
OCTOBER 1969] BLACKWELDER — NATURE OF TAXONOMIC DATA
299
There will be few cases where taxonomists will discard the older data
and switch to biochemical data, but the new data will always be interest-
ing and sometimes of conclusive value. (Here’s a case where the new data
can theoretically work, but the taxonomist would rather fight than
switch. )
The third idea recently “forced” into the consideration of taxonomy
is called Numerical Taxonomy. This is a misnomer, because whether
or not it constitutes taxonomy, it’s much more than just numerical; it’s
statistical. It’s not interested in counting the number of setae but in
statistically analyzing the variation in the number of them. Numerical
taxonomy has been the most active of the challenging fields, as well as
the most illogical; but only a suggestion can be given here of the falla-
cies or the nature of the attack.
Numerical Taxonomy is one of the new fields of systematics. At least,
its protagonists describe it as new. It has been practiced in fairly sophis-
ticated form for more than a hundred years. The only really new thing
about it is its machinery, the computers. This field has been the sub-
ject of more publications than most, summarized in the book by Sokal
and Sneath, Principles of Numerical Taxonomy. It is curious that the
publications in this field seem to ignore all criticisms, as if its internal
growth made each criticism irrelevant. This would be an absurd miscon-
ception.
This was called a new field of systematics ratlier than of taxonomy,
because it’s not a form of taxonomy but a technique useful in several
branches of the more inclusive field of systematics.
Systematics is here used as the broad term for all the fields studying
the diversity of organisms, the causes of the diversity, and its history.
Taxonomy, then, is the actual work of distinguishing kinds and grouping
them into taxa — ^the work that actually occupies most of the time of the
people who call themselves taxonomists.
I call it one of the new fields because it’s listed as one of several “new
methods” in taxonomy. Such as : numerical methods, comparative serol-
ogy, chromatography, electrophoresis, infrared spectroscopy, and cyto-
taxonomy; and a miscellaneous group including: chemistry, electron
microscope cytology, behavior, ecology, histology, and parasitology.
Leaving aside for the moment the first of these, all the rest produce data
about the organisms, data which are at least potentially comparative. As
such, these data are automatically taxonomic. They can be recorded in
the taxonomic system, and they can be directly employed in that system
if there’s need to do so.
The newer forms of data offered to taxonomy by these fields can be
300
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
ei tirely valid taxonomic data of as much value as any older forms of data.
But it’s a mistake to expect to have them used widely. This is simply
because we have already accomplished the erection of an elaborate sys-
te m using the older data. We can’t now switch to the newer forms, dis-
carding much of the older work, unless there are real advantages. There
usually are none, because the really new conclusions from the new data,
on the rare occasions when they are different, can be integrated into the
old system without starting all over again.
Even so, the verification and adjustment of the older conclusions are
valuable and desirable. It’s part of the normal process of growth of
t£ xonomy.
On the other hand, the first item on this list. Numerical Taxonomy,
doesn’t claim to produce new data. It analyzes the old and new data
a ike. This is fine. Much can be gained by such analysis of variation.
But the numericists go much farther and claim that they are doing
t£ xonomy, that the machines produce classifications. In the sense that a
t£ xonomist is likely to understand this, the claim is completely false.
In order to justify the statistical and computer work which they claim
is taxonomy, the numericists make a bitter attack on the classical
n ethods of taxonomy. Unfortunately, they prove to be so ignorant of
w hat taxonomy is that their charges against taxonomy are ridiculous.
JNiumerical writers admit that “numerical taxonomy would have no
c aim to the serious attention of biologists unless it could overcome some
o I the faults found in conventional taxonomic procedure.”
This seems to be fair enough. But the faults which are then cited are
a curious lot. The general critical conclusions of Sokal and Sneath give
some idea of the quality of this attack. They say that taxonomy is a ne-
g ected discipline which has made little progress in this century (or
longer) , is unstable in its results, is inadequate in theory and practice,
aid is subject to many ills such as unreliability, failure to recruit high
c aliber students, circular reasoning, and the admission of work done by
a inateurs. It’s said that “little work has been directed toward the con-
ceptual basis of classification and indeed, the taxonomy of today is but
liltle advanced from that of a hundred or even two hundred years ago.”
If these things are true, then there is indeed much room for numericists
or someone to lead us into improved methods and results. But are they
t ue? Here is the curious thing: several of these charges are completely
t ue ; what is false is the implication that it would be better if it were not
s 3. The last criticism cited, for example, about there being no change in
p rocedures for two hundred years. This is true. The implication that this
is a bad situation is entirely false. The methods and logic of classifica-
OCTOBER 1969] BLACKWELDER NATURE OF TAXONOMIC DATA
301
tion, as developed by the ancient philosophers and adopted by Linnaeus
and his followers, works as well today as it did 200 years ago. Taxono-
mists have had very little reason to want to change it or even to discuss
it.
The fact is that the numericists are so far from understanding the na-
ture of taxonomy that their criticisms are simply irrelevant, and their
claims of replacing it with something better are meaningless.
This doesn’t mean that there is nothing of value to taxonomy in statis-
tical methods. It simply means that statistics offers taxonomy valuable
methods of analysis of variation and related things, but does not, because
it cannot, produce taxonomic data or classifications.
It seems to me to be the critical answer to all the sensible criticisms
that taxonomic results, which are in the nature of hypotheses, must be
testable, and, if the later worker finds them inadequate, that he be able
to emend them, to correct them. This is the basis of conventional tax-
onomy ; it’s not an obvious feature of numerical taxonomy.
The work of a taxonomist consists of three things, besides the routine
matters of collecting, curating, and describing species; First, the selec-
tion of characters to be used for discrimination ; second, the comparative
analysis of the characters in the specimens, to find out if they’re variable,
and so on ; and third, the assignment of the resulting groups to levels in
the taxonomic hierarchy, deciding whether the group is a subfamily or
a distinct family, for example. Numerical studies are able to help us with
the second ; they have no usefulness in the first or third.
You’ll find certain expressions common in the taxonomic literature of
our times. The oldest is The New Systematics. Also biosystematics, bio-
species, biological species concept, natural classification, and others.
These are supposed to reflect modern approaches or new developments
in taxonomy. If you will examine them carefully you’ll find that they
are in an odd position: They are useless without the conventional tax-
onomy as a starting point. Furthermore, they have no actual workable
basis other than the comparative data of the conventional taxonomy.
They may be useful in theoretical discussions, but they’re almost useless
in taxonomic work.
It is well to reiterate what is meant by the expression modern conven-
tional taxonomy. All successful zoological and botanical taxonomy is
based on selected comparative data drawn from individual organisms.
The primary feature of this system is the use of all available data of what-
ever sort, so far as necessary. By “available data” is meant comparative
data of any particular sort that is available for all the objects or taxa
3C2
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
being classified. We can’t classify unless we have this particular data for
al the included items.
For instance, if you have ten species of insects among which blood
aralysis gave two clear-eut groups except for one species in which the
in dividuals were so small that they had too little blood for analysis, you
cc uldn’t use this distinction in your classification of the ten species. Or,
if you had records of the distinctive food habits of all but one, you
cculd use this datum only as supporting evidence, not as basis for the
cl issification.
A further qualification was that we use all available data so far as nec-
essary. This means that when a satisfactory classification is produced
we stop looking for other data. We use the most available data first and
g( ‘ on to use other data only when the obvious ones fail to give an eff ec-
tr e scheme. So we would use biochemical data, for example, only
when external structure is inadequate.
By data, we mean, of course, anything that can be known about the
oiganisms, whether it’s morphological, biochemical, reproductive, or
behavioral. Anything that can be compared from one kind to another.
In this system, the extensive background of the taxonomist enables
hi ra to pass over, almost without conscious thought, all the non-varying
features of the organisms, as well as the ones due to sex, age, pathology,
ai id so on, and to use a workable number of features that are evidently
cc rrelated with many unmentioned ones. All features of the organisms
a] e considered, so far as they are available, but only those are employed
w aich are neeessary to show the groupings and distinctions that occur.
When we find eases in which developmental patterns, or behavioral
patterns, or biochemical components, are not correlated with more read-
ilv observed features, then will be the time to use these less obvious fea-
tr res in the classification. There have already been cases of this. They’re
ex ceptional and apparently rare. In the ordinary situation, use of the
visible and usually structural features involves many unseen but direetly
cc rrelated other features, so that even a “morphological” system is in
reality not exclusively based on one type of feature.
This conventional system, then, is an all-seeing or all-considering sys-
tem. It does not pretend to use all features, because it decides against the
ur>e of some. It knows of the probable existence of other features which it
doesn’t have access to. It stores all these data, whether or not it uses
tiem.
This has been called the omnispective system. It has classified a mil-
lion animals and more than a third of a million plants, so effectively that
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
303
every little failure irks us and leads some of us to say that the system is
inadequate. It’s merely not yet complete.
It’s based entirely on comparative data, the only thing yet discovered
on which it could be based. We still need taxonomists of varied interests
to continue the work of assembling these data, and especially for the work
of classifying — ^translating the comparative data into groups which we
call taxa, so that we can come closer to complete knowledge of the ap-
parently unlimited diversity of the living world.
Records and Descriptions of Mexican and Central American
Tillomorphini
(Coleoptera: Cerambycidae)
John A. Chemsak
University of California, Berkeley
The tribe Tillomorphini is comprised of a rather homogenous group
of species in the Nearctic and Neotropical regions. Apparently this group
is closely related to the Anaglyptini with one of the major differences
between the tribes being the presence of eburneous elytral fasciae in the
Tillomorphini. Linsley (1964) consiAeieA Tilloclytus^aXes^Cyrtophorus
LeConte, and Microclytus LeConte as anaglyptine while Euderces Le-
Conte, Tetranodus Schaeffer, and Pentanodes Schaeffer were placed in
the Tillomorphini. Also at this time Linsley synonymized Eplophorus
Chevrolat with Euderces. It becomes obvious after examining a rela-
tively large amount of material that this synonymy is justified. The
characters of the antennal spines, palpal apices, and basal elytral gib-
bosities vary greatly in the available species. However, for purposes of
convenience, both generic names will be utilized here. While no attempt
is made at this time to critically analyze the tribal relationships of these
groups, the following New World genera may be considered as tillo-
morphine : Euderces, Eplophorus, Cleozona Bates, T etranodus, Pentano-
des, Tillomorpha Blanchard, Lamproclytus Fisher, and Calliclytus Fisher.
The other genera listed in Blackwelder (1946) are probably Anaglyptini
or other.
Most Tillomorphini are rare in collections although the adults may be
collected on flowers. Four genera are known from Mexico and Central
America while the remainder are South American and West Indian.
The Pan-Pacific Entomologist 45: 303-317. October 1969
3)4
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Pentanodes has been recorded only from Brownsville, Texas, but cer-
t£ inly must occur in Mexico.
This study was undertaken during the course of National Science
Foundation Grant GB-4944X on North American Cerambycidae. Celeste
G reen is gratefully acknowledged for the preparation of the illustrations.
Key to the Mexican and Central American Genera
OF Tillomorphini
1 Legs with femora suddenly clavate ; elytra with ehurneous fasciae transverse. 2
Legs with femora gradually clavate; elytra with a double pair of oblique
ehurneous fasciae. Larger species Cleozona
2(1) Elytra with ehurneous fasciae narrow, usually not extending completely
across elytra ; antennae of males without tumid segments 3
Elytra with a single pair of broad ehurneous fasciae extending from
lateral margins to suture; antennae of males with segments three to six
tumid T etranodus
3(2) Antennae with third segment bearing a long apical spine; elytra with basal
gibbosities usually well developed; palpi with apical segments usually
not broadly dilated Eplophorus
Antennae with third segment unarmed or bearing a short spine; elytra
with basal gibbosities absent or very low; palpi with apical segments
often broadly dilated Euderces
Genus Cleozona Bates
Cleozona Bates, 1874, Trans. Entomol. Soc. London, 1874:223; Bates, 1880, Biologia
Centrali-Americana, Coleoptera, 5:60; Linsley, 1935, Trans. Amer. Entomol.
Soc., 61:87.
Type species. — Cleozona pulchra Bates (monobasic).
This monotypic genus is characterized by gradually clavate femora,
s] lined antennae, and a double pair of oblique ehurneous fasciae on the
elytra.
Cleozona pulchra Bates
Cleozona pulchra Bates, 1874, Trans. Entomol. Soc. London, 1874:223; Bates, 1880,
Biologia Centrali-Americana, Coleoptera, 5:60, pi. 6, fig. 10; Chemsak, 1967,
J. Kans. Entomol. Soc., 40 :75.
Cleozona pulchra yar. rufipes Bates, 1885, Biologia Centrali-Americana, Coleoptera,
5:307; Chemsak, 1967, J. Kans. Entomol. Soc., 40:75.
Cleozona rufipes, Linsley, 1935, Trans. Amer. Entomol. Soc., 61:89.
The variety rufipes may constitute a good subspecies but sufficient
n aterial is laeking at this time to analyze the populations. All Mexican
n aterial available has reddish legs while specimens from Guatemala to
]) icaragua possess black legs.
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
305
Type locality. — of pulchra, Chontales, Nicaragua; rufipes, Tehuan-
tepec, Mexico.
Material examined. — Mexico: 1^,8 miles W. Alamos, Sonora, 9 August 1957
(R. L. Westcott) ; 1 ^ , 5 miles W. Alamos, 1-2 August 1953 (F. S. Truxal) ; 2 $ ,
Rio Cucliuhaqui, 7 miles S. Alamos, 25 July-7 August 1953 (F. S. Truxal) ; 2 $ ,
3 ? , Minas Nuevas, Sonora, 7 August 1952 (C. & P. Vaurie) ; 1 $ , 8 miles S. Elota,
Sinaloa, 2 July 1963 (F. D. Parker, L. A, Stange) ; 1 Los Mayos, Sinaloa, 24
July 1952 (J. D. Lattin) ; 1 $,1 miles S. Guamuchil, Sinaloa, 28 July 1966 (J. A.
Chemsak) ; 2 ^,1 2,5 miles N. Mazatlan, Sinaloa, 11 August 1965 (G. H. Nel-
son) ; 24 ^,19 2, Jesus Maria, Nayarit, 26 June and 6 July 1955 (B. Malkin) ; 2
$ ,7 2,3 miles N.W. Arroyo Santiago, near Jesus Maria, Nayarit, 5 July 1955 (B.
Malkin); 1 ^,3 miles N.W. Santa Maria del Oro, Nayarit, 27 June 1963 (J.
Doyen) ; 1 S,2 2, 10 miles W. Colima, Colima, 1 August 1954 (Cazier, Gertsch,
Bradts) ; 3^,32, Tecolopa, Colima, 31 July 1954 (Cazier, Gertsch, Bradts) ; 1^,
5 miles E. Apatzingan, Michoacan, 19 July 1954 (Linsley, MacSwain, and Smith) ;
1 2 , 11 miles W. Hidalgo, Michoacan, 12 July 1963 (Parker, Stange) ; 1 2 , 6 miles
S. Rio Mexcala, Guerrero, 5 August 1965 (G. H. Nelson) ; 1 2, 3 miles W. Tehuan-
tepec, Oaxaca, 2 August 1965 (G. H. Nelson). Linsley (1935) records one speci-
men from Tejupilco, Mexico, July (Hinton).
Guatemala: 1 2, Guatemala, ex. Guajacum officinale.
Genus Tetranodus Linell
Tetranodus Linell, 1896, Proc. U. S. Nat. Mus., 19:396; Linsley, 1935, Trans.
Amer. Entomol. Soc., 61:87; Linsley, 1964, Univ. Calif. Publ. EntomoL, 22:185.
Tetranodes, Schaeffer, 1904, J. N. Y. Entomol. Soc., 12:223.
Based upon the type species, Tetranodus niveicollis Linell, and on
available material, this genus appears to be a rather uniform unit. In all
cases, the ivory fasciae of the elytra are broad and extend across the
width of the elytra, the eyes are pointed behind, and the males have seg-
ments three to six of the antennae incrassate. Four species are presently
known.
Key to the species of Tetranodus
1 Elytra with basal punctures coarse, dense, without broad longitudinal
spaces between rows of punctures 2
Elytra with basal punctures fine, sparse, arranged in rows separated by
broad longitudinal glabrous spaces; color shining black, appendages
paler. Length, 5 mm. Chiapas to Guatemala reticeps (Bates)
2(1) Elytra behind fasciae separately punctate, not scabrous 3
Elytra behind fasciae deeply, closely punctate, scabrous appearing; color
reddish, elytra dark behind ivory fasciae. Length, 4.5 mm. Sinaloa
Tugipennis n. sp.
3(2) Pronotum cylindrical, sides not angulate at middle; elytra with basal
punctures dense, not linearly arranged; males with segments three to
six of antennae strongly incrassate. Length, 3.5-5 mm. Texas to Oaxaca
niveicollis Linell
336
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Pronotum with sides somewhat expanded, angulate; elytra with hasal
punctures linearly arranged, lines separated; males with segments
three to six slightly expanded. Length, 3-4 mm. Honduras
angulicollis n. sp.
Tetranodus RETICEPS (Bates), (new combination)
luderces reticeps Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5:59.
The type is a female from Guatemala (near the city) . It fits well into
Tetranodus by the broad elytral fasciae and eyes pointed behind. A fe-
riale is available from 4 miles N.W. Pueblo Nuevo, River Bajada, Chi-
apas, Mexico, 15 July 1965 (G. H. Nelson).
Tetranodus rugipennis Chemsak, new species
Female.- — Form small, cylindrical; integument reddish, elytra dark behind
eburneous fasciae. Head reticulate punctate, long erect hairs sparse; palpi with
c pices dilated; eyes entire, pointed behind; antennae filiform, shorter than body,
tiird segment longer than scape, fourth shorter than first, segments from seventh
sliort, segments unarmed, sparsely pubescent. Pronotum longer than broad, base
sharply constricted, apex shallowly impressed at sides; disk longitudinally striate;
long erect hairs sparsely interspersed over surface, base with a fine band of pale
cppressed pubescence; prostemum impressed, apical one-third glabrous, hasal two-
t birds sparsely punctate; mesosternum with epimera clothed with white appressed
]iubescence; metasternum with a band of appressed pubescence posteriorly. Scutel-
lim densely clothed with white appressed pubescence. Elytra almost 2% times
linger than basal width, wider behind middle, transversely impressed before fasciae;
eburneous fasciae antemedian; disk rather densely, coarsely punctate before fasciae,
]iosterior portion densely, rugosely punctate, more finely toward apex; pubescence
ery sparse, erect; apices rounded. Legs with tibiae bicarinate; pubescence sparse,
btbdomen very sparsely punctate; apex of last sternite rounded. Length, 4.5 mm.
Holotype female (Canadian National Collection) from 5 miles N.
Mazatlan, Sinaloa, Mexico, 24 July 1964, on flowers of Buddleia
ivrighlii (H. F. Howden) .
The rugose appearing elytra behind the eburneous fasciae distinguish
this species.
Tetranodus niveicollis Linell
'^etranodus niveicollis Linell, 1896, Proc. U. S. Nat. Mus., 19:396; Linsley and
Martin, 1933, Entomol. News, 44:181 (habits) ; Linsley, 1935, Trans. Amer.
Entomol. Soc., 61:91; Linsley, 1964, Univ. Calif. Publ. Entomol., 22:185.
Cetranodes niveicollis, Schaeffer, 1904, J. N. Y. Entomol. Soc., 12:223 (habits).
Tetranodus mexicanus Linsley, 1935, Trans. Amer. Entomol. Soc., 61:90. (New syn-
onymy.)
The characters used to separate mexicanus from niveicollis vary in the
series at hand. The color differences in the elytra may be geographic
3ut more material will be required to assess the situation.
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
307
Type locality, — of niveicollis, Brownsville, Texas; of mexicanus,
Tejupilco, Mexico, Mexico.
Material examined. — Mexico-. 1^, Penuela, Veracruz, 15 July 1941 (H. Dy-
bas) ; 1 Puente Nacional, Veracruz, 3 July 1941 (H. Dybas) ; 1 $,29 miles E.
Cordoba, Veracruz, 12-25 July 1964 (E. Fisber, D. Verity) ; 2$ , Cotaxtla Experi-
ment Station, Cotaxtla, Veracruz, 30 June 1962 (D. H. Janzen) ; 1 $ , 56 miles N.W.
Tehuantepec, Oaxaca, 27 July 1963 (J. Doyen).
Tetranodus angulicollis Chemsak, new species
Male.— Form small, cylindrical; integument dark reddish brown, antennae paler.
Head reticulate punctate, sparsely clothed witli long erect hairs; palpi with apices
strongly dilated; eyes large, pointed behind; antennae longer than body, segments
three to six slightly expanded, third segment longer than scape, fourth shorter than
first, segments unarmed, sparsely pubescent. Pronotum a little longer than broad,
sides expanded, angulate; disk longitudinally striate; long erect hairs sparse, base
with a band of appressed pubescence ; prosternum impressed, apical one-third trans-
versely wrinkled, remainder glabrous; mesosternum with epimera clothed with ap-
pressed pubeseenee; metasternum with a band of appressed pubescence posteriorly.
Scutellum densely clothed with appressed pubescence. Elytra over 2% times as
long as broad, slightly broader behind middle; base with vague, flat gibbosities;
disk impressed before fasciae; eburneous fasciae antemedian; punctures well sep-
arated, slightly smaller at apex; pubescence sparse, hairs erect, long and short;
apices rounded. Legs sparsely pubescent, tibiae bicarinate. Abdomen subglabrous;
apex of last sternite broadly rounded. Length, 3-4 mm.
Holotype male (United States National Museum) and one male para-
type from Tegucigalpa, Honduras, 12 May 1912 (F. J. Dyer) .
The angulate pronotum and weakly inflated antennal segments readily
separate this species from the others.
Genus Eplophorus Chevrolat
Eplophorus Chevrolat, 1861, J. EntomoL, 1:248; Linsley, 1935, Trans. Amer. Ento-
mol. Soc., 61 :87.
Apelocera Chevrolat, 1862, Ann. Soc. Entomol. Fr., (4)2:61, Thomson, 1864, Sys-
tema Cerambycidarum, p. 195.
Apilocera Chevrolat, 1862, Ann. Soc. Entomol. Fr., (4)2:535; Pascoe, 1866, Trans.
Entomol. Soc. London (3)5:295; Bates, 1880, Biologia Central!- Americana,
Coleoptei'a, 5:60.
Blackwelder (1946) lists 18 species of Eplophorus distributed from
Mexico to Colombia and Brasil. The Brasilian species, waltli Chevrolat,
is doubtfully congeneric because of its distribution and at least two of the
other species are synonyms. Since most of the species are known only
from the type, only those species at hand will be listed below.
Type species. — Tillomorpha spinicornis Thomson (by original desig-
nation).
338
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Eplophorus boucardi (Chevrolat)
Apilocera boucardi Chevrolat, 1862, Ann. Soc. Entomol. Fr., (4)2:535; Bates, 1880,
Biologia Central!- Americana, Coleoptera, 5:60.
Type locality. — Coban, Vera Paz, Guatemala.
Material examined. — Mexico-. 1^, 15 miles W. Las Cruces, Chiapas, 27 July
1352 (Gilbert, MacNeill) ; 1 $ , La Trinitaria, CJiiapas, 17 June 1965 (Burke,
IV.eyer, Schaffner) ; 1 , 5 miles N.N.W. Tuxtla Gutierrez, Chiapas, 22 July 1954
( A. A. Alcorn) ; 1 ^,1 $, Tuxtla Gutierrez, 22 June 1965 (Burke, Meyer, Schaff-
ner) ; 1 $, Sumidero Canyon, Tuxtla Gutierrez, 21-22 July 1963 (E. Fisher) ; 1 2,
10 miles W. Tuxtla Gutierrez, 21 June 1955 (R. B. & J. M. Selander) ; 1 5,2 2,
4 miles N.W. Pueblo Nuevo, River Bajada, Chiapas, 15 July 1965 (G. H. Nelson) ;
1 5 , El Suspiro, Berriozabal, Chiapas, 21 June 1955 (R. B. & J. M. Selander) .
I onduras: 1 S , Comavagua, 10 July 1964 (G. A. Axtell).
Eplophorus bicinctus Linsley
Iplophorus bicinctus Linsley, 1935, Trans. Amer. Entomol. Soc., 61:89; Linsley,
1942, Proc. Calif. Acad. Sci., (4)24:56.
L uderces balli Knull, 1935, Entomol. News, 46:192; Linsley, 1964, Univ. Calif. Publ.
Entomol., 22:191. (New synonymy.)
Iplophorus bicinctus peninsularis Linsley, 1942, Proc. Calif. Acad. Sci., (4)24:56.
This species is very distinctive by the double pair of elytral fasciae.
Knull’s (1935) description of Euderces balli fits the long series at hand
f rom Mexico very well.
Linsley ’s (1942) designation of the subspecies peninsularis from San
Domingo, Baja California, is probably valid.
Type locality. — of bicinctus, Los Mochis, Sinaloa, Mexico; of balli,
Baboquivari Mts., Arizona.
The following specimens examined are all assignable to Eplophorus
bicinctus bicinctus.
Mexico: 10 5, 10 2, Minas Nuevas, Sonora, 7 August 1952 (C. & P. Vaurie) ;
1 5,1 2 , Alamos, Sonora, 13 August 1965 (G. H. Nelson) ; 1 2 , 12 miles N. Her-
iiosillo, Sonora, 14 August 1965, on Olneya tesota (G. H. Nelson) ; 1 5 , 5 miles W.
iLlamos, 1-2 August 1953 (F. S. Truxal) ; 1 5, 18 miles W. Alamos, 30 July 1957
(R. L. Westcott) ; 1 5,2 2, 10 miles E. Navajoa, Sonora, 13 August 1959 (W. L.
Hutting, F. G. Werner); 1 2 , 12 miles N.E. El Fuerte, Sinaloa, 12 July 1962
(Sleeper, Anderson, Hardy, Somerby) ; 1 2, Culiacan, Sinaloa, 21 July 1959 (H. E.
Evans) ; 1 5,2 2,7 miles S. Guamuchil, Sinaloa, 28 July 1966 ( J. A. Chemsak) .
Eplophorus longicollis Linsley
Eplophorus longicollis Linsley, 1935, Trans. Amer. Entomol. Soc., 61:88, pi. 2,
fig. 4.
This species also has a double pair of eburneous elytral fasciae but the
Eiiterior pair is reduced to two sutural dots. There is variation in the
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
309
amount of red on the elytra. Some specimens have the basal half reddish
while others are infuscated.
Type locality. — Bejucos, Mexico, Mexico.
Material examined. — Mexico: 2 2, 8 miles S. Elota, Sinaloa, 2 July 1963
(Parker, Stange) ; 33^, 42 2, Jesus Maria, Nayarit, 26 June 1955 (B. Malkin) ;
7 2,3 miles N.W. Arroyo Santiago, near Jesus Maria, 5 July 1955 (B. Malkin) ;
1 ^ , 55 miles W. Tequila, Jalisco, 28 June 1963 (J. Doyen) ; 12, Xalitla, Guerrero,
4 June 1946 (J. & D. Pallister) ; 1^,5 miles E. Apatzingan, Michoacan, 19 July
1954 (Linsley, MacSwain, Smith); 12, Jalastoc, Morelos, 22 May 1954 (J.
Pacheco) ; 1 2 , Lake Tequesquitengo, Morelos, 5 June 1959 (H. E. Evans) ; 1 ^ ,
Alpuyeca, Morelos, 8 July 1951 (P. D. Hurd) ; 1 2 , Acatlan, Puebla, 19 July 1955
(P. & C. Vaurie) ; 1 (J , 10 miles S.E. Pitago, Puebla, 27 June 1955 (R. B. & J. M.
Selander) ; 1^, Cacaloapan, Puebla, 26 April 1962 (F. D. Parker).
Eplophorus spinicornis (Chevrolat)
(Fig. 1)
Clytus spinicornis Chevrolat, 1835, Coleopteres du Mexique, fasc. 4, no. 7 ; Thom-
son, 1860, Classification des cerambycides, p. 229.
Eplophorus spinicollis, Chevrolat, 1861, J. Entomol., 1:248 (error for spinicornis).
Apelocera spinicornis, Chevrolat, 1862, Ann. Soc. Entomol. Fr., (4)2:62.
Apilocera spinicornis, Chevrolat, 1862, Ann. Soc. Entomol. Fr., (4)2:535; Bates,
1880, Biologia Central!- Americana, Coleoptera, 5:60.
Eplophorus spinicornis, Blackwelder, 1946, U. S. Nat. Mus. Bull., 185:583.
Clytus elegans Laporte and Gory, 1835, Monographic du genre Clytus, PI. 20.
In addition to the Biologia material, one specimen is at hand from
Jalapa, Veracruz, Mexico.
Eplophorus hoegei (Bates)
Apilocera Hogei Bates, 1885, Biologia Central!- Americana, Coleoptera, 5:305.
This species is characterized by the very long spine of the third an-
tennal segment, very prominent basal gibbosities of the elytra, and bi-
colored elytra.
Type locality. — Cerro de Plumas, Veracruz, Mexico.
Material examined. — 2 2 , 4 miles N.W. Pueblo Nuevo, River
Bajada, Chiapas, Mexico, 12-14 July 1965 (G. H. Nelson).
Eplophorus auricomis Chemsak, new species
Male. — Form moderate sized, cylindrical; color black, mouthparts, antennae,
legs, mesosternum, and lateral posthumeral spots on elytra pale; pubescence golden,
mostly appressed. Head ■with front vertical, very finely, densely punctate; pubes-
cence dense, appressed, obscuring surface; eyes divided, upper lobe small;
antennae about as long as body, scape paler, third segment % longer than scape,
almost 3 times longer than fourth segment, remaining segments short, eleventh
tapering at apex, spine of third segment blunt at apex, as long as fourth segment.
310
THE PAN-PACIFIC ENTOMOLOGIST
[vOL. 45, NO. 4
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
311
segments from fourth densely clothed with short appressed pubescence except
below. Pronotum about 1.3 times longer than broad; base constricted; disk longitu-
dinally striate, densely clothed with appressed pubescence except for apical margin
and an arcuate submedial band, long erect hairs numerously interspersed; pro-
sternum deeply impressed, apical one-third glabrous, basal two-thirds coarsely
deeply punctate, densely pubescent; mesosternum shining, testaceous, episternum
with a band of white appressed pubescence at margin which extends across
middle coxae; metasternum finely punctate and pubescent, posterior margin with
a band of dense white pubescence, episternum covered by elytra. Scutellum
moderately golden pubescent. Elytra about twice as long as broad; basal gib-
bosities highly elevated, arcuate, extending over basal one-third of elytra; each
side with a single ivory fascia behind basal gibbosities extending from lateral
margin but not to suture; posthumeral area pale, coarsely, separately punctate;
area behind ivory fasciae black, very densely punctate, opaque, appressed pubes-
cence becoming denser to glabrous transverse lines at apical one-third, apical one-
third densely clothed with appressed golden pubescence which obscures the surface,
long erect hairs present at base and apex; apices subtruncate. Legs with femora
clavate, each femur with a fine band of pubescence on outside surface, hind pair
extending beyond elytral apices; tibiae carinate. Abdomen minutely punctate,
stemites becoming more pubescent to apex; apex of last sternite rounded. Length,
7 mm.
Holotype male (Chicago Museum of Natural History) and one male
paratype from Turrialba, Costa Rica, June-July 1949 (K. W. Cooper) .
Eplophorus aspeiricollis Chemsak, new species
Male. — Form moderate sized, rather slender; color black, mouthparts and
appendages very dark rufous, two antemedian fasciae eburneous; pubescence
whitish. Head very densely, shallowly punctate, pubescence sparse, long, erect;
eyes divided; antennae about as long as body, third segment twice as long as
scape, fourth segment less than half as long as third, spine of third segment about
half as long as fourth segment, fourth segment minutely dentate, apical segments
finely clothed with very short pubescence, underside of segments three to six with
short erect hairs, each segment to seventh with a very long erect hair at apex.
Pronotum about 1.4 times longer than broad; base constricted; disk very finely
asperate, sparsely clothed with short appressed pubescence, more densely at base, long
erect hairs numerous; prosternum glabrous at apical third, basal two-thirds
coarsely punctate, moderately densely pubescent; mesosternum with a dense patch
of white appressed pubescence at sides; metasternum densely pubescent poste-
riorly. Scutellum finely pubescent. Elytra over twice as long as broad; basal
gibbosities obtuse, short, coarsely, deeply punctate; ivory fasciae antemedian,
extending from lateral margins but not to suture; median area very deeply
densely punctate, opaque, apical one-third shining, almost impunctate; pubescence
Figs. 1-4. Typical elytral patterns of: Fig. 1. Eplophorus spinicornis Chev-
rolat. Fig. 2. Euderces parallelus LeConte. Fig. 3. Eu. ciibratus Bates. Fig. 4.
Eu. howdeni Chemsak.
3.2
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
wiitish, short, depressed, extending over basal one-third from humeri arcuately
to suture just before ivory fasciae, a broad band present at apical one-third ex-
teading back from suture to lateral margins, apical portion sparsely clothed
w th short pubescence, long erect hairs present over basal one-third; apices
tiLincate. Legs with femora clavate, with a pubescent hne along outside and
ir side; tibiae carinate. Abdomen finely punctate at edges; first sternite pubescent
a1 sides; apex of last sternite rounded. Length, 6 mm.
Female.- — Antennae shorter. Prosternum very finely punctate before coxae.
Abdomen with apex of last sternite rounded. Length, 7 mm.
Holotjpe male (Cornell University) from Tuxpan, Michoacan, Mex-
ico, 6 July 1959, 6000 ft. (H. E. Evans). Allotype from 3 miles N.
Tzitzio, Michoacan, Mexico, 5500 ft., 29 July 1962 (U. Kansas Mex.
E Kped. ) .
This species somewhat resembles E. cribellata (Bates) but lacks the
pale areas of the elytra, the rugosely punctate pronotum, and long blunt
s]>ine of the third antennal segment.
Genus Euderces LeConte
E’xderces LeConte, 1850, J. Acad. Natur. Sci. Philadelphia, (2)2:30. (For com-
plete bibliography see Linsley, 1964, Univ. Calif. Publ. Entomol., 22:180.)
As here restricted, this genus includes only those species which lack
aitennal spines or possess only short spines on the third segment.
Type species. — Callidium picipes Fabricius (monobasic) .
Eight species are recognized here from Mexico and Central America.
Key to the Mexican and Central American
Species of Euderces
1 Elytra with apices rounded, emarginate or subtruncate 2
Elytra with apices acutely produced; pronotum wdth sides expanded
before middle; integument black, elytra with a single pair of eburneous
fasciae. Length, 3 mm. Guatemala acutipennis Bates
2(1) Elytra with a double pair of eburneous fasciae 3
Elytra with a single pair of eburneous fasciae 4
3(2) Elytra with posterior pair of ivory fasciae extending to suture; anterior
pair very close to posterior pair, area between fasciae rufous. Length,
3.5-5 mm. Baja California del Sur parallelus LeConte
Elytra with posterior pair of fasciae not attaining suture, anterior pair
far removed; elytra rather uniformly dark rufous to black. Length,
4 mm. Oaxaca to Guatemala cribratus Bates
4(2) Pronotum with disk finely asperate, not longitudinally striate 5
Pronotum with disk longitudinally striate 6
5(4) Elytra shining, rather uniformly punctate; pronotum and underside
densely clothed with white appressed pubescence, long white flying
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
313
hairs numerous; integument uniformly dark reddish-brown. Length,
7 mm. Durango howdeni n. sp.
Elytra very densely, opaquely punctate over basal one-half, apical half
almost impunctate; pronotum sparsely clothed with long erect pubes-
cence, surface dull; integument reddish, apical half of elytra dark.
Length, 4-5 mm. Texas to Tamaulipas reichei exilis Casey
6(4) Antennae without spines, third segment minutely dentate at most; elytra
not basally gibbose 7
Antennae with segments three to five distinctly spined at apices; elytra
with prominent gibbosities at base; color reddish, elytra darker at
apex. Length, 4.5 to 6 mm. Sinaloa nelsoni n. sp.
7 (6) Elytra with pubescence sparse, hairs long and erect, appressed pubescence
lacking; integument imiformly all black. Length, 3.5 to 5 mm. Naya-
rit to Morelos cribripennis Bates
Elytra with oblique bands of densely appressed pubescence at apical
one-third; integument bicolored, elytra red and black, pronotum vari-
able. Length, 4-6 mm. Puebla to Oaxaca laevicauda Bates
Euderces acutipennis Bates
Euderces acutipennis Bates, 1885, Biologia Centrali-Americana, Coleoptera, 5:305,
pi. 21, fig. 15.
This species is known only from the unique type from Panajachel,
Guatemala. It is very distinctive among the known species of Euderces by
the strongly produced apices of the elytra. Because of this character,
the cup-shaped pronotum, and deeply inserted head, E. acutipennis may
prove to be non-congeneric with the other species.
Euderces parallelus LeConte
(Fig. 2)
Euderces parallelus LeConte, 1873, Smithson. Misc. Collect., 11(264) :202; Leng,
1887, Entomol. Amer., 3:24, 44; Horn, 1894, Proc. Calif. Acad. Sci., (2)
4:339; Linsley, 1942, Proc. Calif. Acad. Sci., (4)24:57.
The double pair of ebumeous fasciae which extend to the elytral
suture make this species distinctive. The anterior pair of fasciae may be
greatly reduced on occasion.
It is known only from the Cape Region of Baja California.
Type locality. — Lower California.
Material examined. — Baja California: 1 2, San Domingo, 19 July 1938 (Michel-
bacher and Ross) ; 1 $ , Triunfo, 13 July 1938 (Michelbacher and Ross) ; 1 , 1 $ ,
Santa Rosa (Beyer) ; 1 $ , 26 miles W. La Paz, 10 August 1966, on flowers of
Yucca (J. A. Chemsak) ; 12, Todos Santos, 13 August 1966 (J. T. Doyen) ; 12,
1 mile S.W. Punta Palmilla, 13 September 1967 (J. Chemsak, A. & M. Michel-
bacher) .
314
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Euderces cribratus Bates
(Fig. 3)
El derces cribratus Bates, 1885, Biologia Centrali-Americana, Coleoptera, 5 :304.
Although the type locality is San Joaquin, Guatemala, a single female
spjcimen from 10 miles S.E. Huajuapan de Leon, Oaxaca, Mexico, 27
June 1965 (Burke, Meyer, Schaffner) is at hand which appears to be
assignable to this species. The type has rufous antennae and femora
and the elytra are now brownish although Bates stated the color as
black in the original description. The anterior ivory fasciae are reduced
to dots and well removed from the other pair. The specimen at hand is
al] black with somewhat rufous appendages.
Euderces howdeni Chemsak, new species
(Fig. 4)
lVIale. — ^Form moderate sized; integument shining, unifor ml y dark reddish-
brown; each elytron with an antemedian eburneous fascia; pubescence thick,
wl lite, appressed. Head with front densely clothed with appressed white pubescence
and long erect hairs; eyes divided, upper lobe small; antennae about as long as
body, third segment longer than scape, fourth segment three-fourths as long as
th rd, fifth subequal to third, remaining segments shorter than fifth, subequal,
basal segments sparsely clothed with long erect hairs beneath, all segments clothed
with very short appressed pubescence. Pronotum slightly longer than broad,
feebly impressed transversely behind apical margin; disk shining, irregularly
as derate punctate ; short depressed pubescence more dense at sides, long erect
hairs numerously interspersed; prosternum with apical margin glabrous, re-
minder coarsely, deeply punctate, densely clothed with appressed pubescence;
m^so- and metasternum densely pubescent broadly at sides. Scutellum densely
clothed ■with white appressed pubescence. Elytra over 2% times longer than broad,
net impressed at sides; antemedian eburneous fasciae not extending to suture
ncr lateral margins; short, white, appressed pubescence forming a line down
suture and a broad, oblique fascia at apical one-third; short depressed hairs and
very long erect hairs scattered over surface; punctures coarse, irregular, becoming
fil er at apex; basal gibbosities low; apices rounded. Legs clothed ■with long
flying hairs. Abdomen rather densely pubescent; apex of last sternite slightly
er larginate. Length, 7 mm.
Holotype male (Canadian National Collection) from 24 miles W.
Li Ciudad, Durango, Mexico, 20 June 1964, 7000 ft. (H. F. Howden).
This is probably the most easily recognizable species of Euderces.
No other species has the shining integument and dense appressed pubes-
ce nee.
I take pleasure in dedicating this species to H. F. Howden for his
ti] eless efforts in collecting Cerambycidae.
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
315
Euderces reichei exilis Casey
(Fig. 5)
Euderces exilis Casey, 1893, Ann, N. Y. Acad. Sci., 7:591; Linsley and Martin,
1933, Entomol. News, 44:191 (habits) ; Vogt, 1949, Pan-Pac. EntomoL, 25:144
(habits) .
Apelocera exilis, Aurivillius, 1912, Coleopterorum Catalogus, 39:420,
Euderces reichei exilis, Linsley, 1964, Univ. Calif. Publ. Entomol., 22:182.
The rather dull, finely asperate disk of the pronotum characterizes
this species.
Type locality. — Texas.
Material Examined. — Mexico : 2 $ , Santa Teresa, Tamaulipas, 15
May 1952 (Cazier, Gertsch, Schranunel) ; 1 2 , 9 miles S. Victoria,
Tamaulipas, 18 May 1952 (Cazier, Gertsch, Schrammel) ; 1 2 , Llera
Mesa, Tamaulipas, 1800 ft., 7 June 1961, on flowers of Acacia wrightii
(U. Kansas Mex. Exped.) .
Euderces nelsoni Chemsak, new species
(Fig. 6)
Male. — F orm small, cylindrical; integument reddish, elytral apices, abdomen,
and hind femora often dark. Head with front longitudinally striate, non pubes-
cent; eyes divided, upper lobe very small; antennae extending to about apical
one-third of elytra, segments three to five distinctly spined at apices, third seg-
ment about 1% times longer than scape and fourth segment, segments from fifth
gradually diminishing in length, outer segments minutely pubescent. Pronotum
longer than broad; disk longitudinally striate, sparsely clothed with fine appressed
pubescence, long, erect hairs numerous; prostemum glabrous apically, remainder
coarsely, deeply punctate, sparsely pubescent; mesosternum with a band of ap-
pressed pubescence over epimeron ; metastemum pubescent posteriorly toward sides.
Scutellum sparsely pubescent. Elytra over twice as long as broad, sides impressed
before middle; each elytron with an antemedian ebumeous fascia extending from
lateral margin but not attaining suture; basal gibbosities rather prominent;
punctures very deep, close, subopaque to apical one-third, apical area non
punctate; pubescence at apical one-third dense, fine, appressed, long, erect hairs
sparsely interspersed toward base and apex; apices rounded to shallowly emargi-
nate. Legs very sparsely pubescent. Abdomen sparsely pubescent; apex of last
sternite narrowly rounded. Length, 5-6 mm.
Female. — Form similar. Prosternum subglahrous throughout, not punctate.
Length, 5-6 mm.
Holotype male (California Academy of Sciences), allotype and 5
paratypes {2 $ , 3 2) from 5 MILES N. Mazatlan, Sinaloa, Mexico,
11 August 1965, on dead limbs (G. H. Nelson). An additional female
paratype from same locality, 24r-29 July 1964 (H. F. Howden) .
The spined apices of the elytra, single pair of ebumeous fasciae, and
dull reddish color readily separate this species. The type series is very
uniform in coloration.
316
[voL. 45, NO. 4
THE PAN-PACIFIC ENTOMOLOGIST
OCTOBER 1969] CHEMSAK TILLOMORPHINE CERAMBYCIDS
317
E. nelsoni is named for G. H. Nelson, who made most of the type
series available for study.
Euderces cribripennis Bates
(Fig. 7)
Euderces cribripennis Bates, 1892, Trans. Entomol. Soc. London, 1892:165, pi. 6,
fig. 14; Linsley, 1935, Trans. Amer. Entomol. Soc., 61:89.
This species appears very close to E. laevicauda but differs in being
all black and lacks the fine appressed pubescence at the apical one-third
of the elytra.
Type locality. — Chilpancingo, Guerrero.
Material examined. — 2 $ , 5 miles S. Tuxpan, Michoacan, 4 August
1962 (D. S. Verity) ; 1 2 , 3 miles N.W. Santa Maria del Oro, Nayarit,
27 June 1963 (J. Doyen) ; 1 ^ , Cuernavaca, Morelos, 12-19 July 1961
(R. & K. Dreisbach). Linsley (1935) lists two specimens from Teju-
pilco, Mexico, July (Hinton and Usinger) .
Euderces laevicauda Bates
(Fig. 8)
Euderces laevicauda Bates, 1885, Biologia Centrali-Americana, Coleoptera, 5:304.
Type locality. — San Geronimo, Guatemala.
Material examined. — Mexico: 2 $ , 2 $ , Tehuacan, Puebla, 6 July
1941, 5500 ft. (H. Dybas) ; 2 $ , 45 miles N. Acatlan, Puebla, 30 July
1963 (J. Doyen) ; 1 ^ , 18 miles N.W. Totolapan, Oaxaca, 29 July 1963
(J. Doyen) ; 2 $ , 10 miles S.E. Haujuapan de Leon, Oaxaca, 27 June
1965 (Burke, Meyer, Schaffner) .
Literature Cited
Blackwelder, R. E. 1946. Checklist of the coleopterous insects of Mexico,
Central America, the West Indies, and South America. U. S. Nat. Mus.
Bull., 185(4): 551-763.
Knull, j. N. 1935. New Coleoptera. Entomol News, 46:189-193.
Linsley, E. G. 1935. Studies in the Longicornia of Mexico. Trans. Amer.
Entomol. Soc., 61: 67-102.
1942. Contributions toward a knowledge of the insect fauna of Lower Cali-
fornia. No. 2. Coleoptera: Cerambycidae. Proc. Calif. Acad. Sci.,
(4)24: 21-96.
1964. The Cerambycidae of North America. Part 5. Taxonomy and classi-
fication of the subfamily Cerambycinae, tribes Callichromini through
Ancylocerini. Univ. Calif. Publ. Entomol., 22: 1-197.
<-
Figs. 5-8. Typical elytral patterns of: Fig. 5. Euderces reichei exilis Casey.
Fig. 6. Eu. nelsoni Chemsak. Fig. 7. Eu. cribripennis Bates. Fig. 8. Eu.
laevicauda Bates.
318
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Thinoseius spinosus Found in New and Unusual Habitat
(Acari: Eviphididae)
Dale Van Volkinburg
Oregon State University, Corvallis
There are only four species of the genus Thinoseius according to
E ans (1962), three from beaches of N. Europe and one, T. hrevistern-
alis (Canaris) from the Pacific Coast, supposedly the only representative
of the genus in N. America. The three European species T. fucicola
(.lalbert), T. spinosus (Willmann) and T. acuminatus (Evans) have all
been reported from tidal debris (Evans, 1954) collected from the
be aches of N. Germany, Great Britain and Iceland. Thinoseius spinosus,
ir particular, has been reported from the supratidal debris of N. Ger-
many and Reykjavik, Iceland (Sellnick, 1940).
Thinoseius hrevisternalis, the Pacific kelp mite (suggested common
name) , has been reported in the literature only as a phoretic associate
oi several species of beach hoppers of the genus Orchestoidea (Crus-
ts cea: Amphipoda), although I have found them free-living in all life
stages on partially decayed kelp. This mite is neither truly marine nor
wliat is normally considered terrestrial, but inhabits the beach-zone
li ter between high tide and the terrestrial border.
I was surprised to learn therefore that we had specimens of the
European species T. spinosus in our acarology collection which were
collected in 1966 from chicken litter and manure at the campus chicken
fs rm in Corvallis, Oregon. Collecting at the same place in February,
1969 confirmed this. Thinoseius spinosus was found in great numbers
and in breeding condition, all life stages being noted. Identification of
tl e species was confirmed by Dr. G. W. Krantz of this department. I
was also successful in collecting them from a commercial chicken farm
15 miles east of Corvallis. It should be noted that both these locations
a e well inland, 50-65 miles from the Pacific Coast.
It is singularly peculiar, not so much that T. spinosus should be dis-
cc'vered in N. America, but that it is found in what appears to be such
a:i entirely different type of microhabitat. If present in this country at
all, it seems one would expect to find it along our beaches occupying a
fscet of the coastal strand community similar to its “niche” in Europe;
however, in three years of investigating Pacific Coast debris I have not
encountered it.
Concerning the presence of T. spinosus in this unexpected locality, I
Tse Pan-Pacific Entomologist 45: 318-3'19. October 1969
OCTOBER 1969] VAN VOLKINBURG NEW MITE HABITAT
319
first assumed that it had been “accidentally” imported either with fish
meal, which is a common nutritional supplement used by poultrymen,
or with kelp itself, which is occasionally incorporated into chicken feed.
Two types of fish meal, one Canadian and one Norwegian, used here
in Corvallis, were checked for mites with negative results. Both the meal
and the chicken feed have such low humidities in comparison to either
manure or tidal debris that it is difficult to believe that these mites
could survive long or breed in either fish meal or chicken feed.
The fact that T. spinosus is so abundant and breeding in this “new”
microhabitat suggests that its presence here has been one of much
longer duration and raises a major question as to whether coastal debris
or chicken litter is the preferred and original habitat for T. spinosiis. It
would be interesting to learn if T. spinosus is also found typically in
chicken houses of Europe and farther inland in this country. Recent
collecting in mink litter and manure has not shown the mite to be
associated with that particular microhabitat.
If T. spinosus is not a typical inhabitant of chicken houses in Europe
and continues to be collected only from tidal debris there, then its
presence here should pose some new and provocative problems concern-
ing its dispersal and apparent adaptability to this seemingly new and
atypical ecological situation.
Acknowledgments
I wish to thank Rodger Shoemake, Whitworth College, Washington,
for specimens of Thinoseius spinosus and James Wernz, Oregon State
University, for assistance in final editing during my absence. Funds
for this research were provided by N.I.H. pre-doctoral fellowship
#5-F01 GM-32, 975 03.
Literature Cited
Evans, G. 0. 1954. A revision of the genus Thinoseius Halbert 1920 (Acari:
Laelaptidae) . Ann. Mag. Natur. Hist., (12)7, 615-622.
1962. The systematic position of Gammaridaearus brevis ter ncdis (Canaris)
(Acari: Mesostigmata) . Ann. Mag. Natur. Hist., (13)5, 395-399.
Sellnick, M. 1940. Die Milbenfauna Islands. Goteborgs Vetensk-Samh. Handl.
(5) 6B 14, 102-104.
320
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Distribution and Flights of Rare North American Desert
Termites of the Genus Amitermes^
(IsopteraiTermitidae)
W. L. Nutting
University of Arizona, Tucson
These notes are presented as an interim report to stimulate a new
interest in, and to further the search for, some of the rarer but none-
tieless important termites in the southwestern United States. Consider-
ing man’s increasing interest in termites from both the economic and
scientific points of view, it is disturbing to realize that, of approximately
38 species in the continental United States, all the castes of two species
Lave not yet been discovered. Only the worker and soldier castes are
Lnown for Amitermes snyderi Light, while only the alate is known for
/ mitermes pallidus Light.
Although Amitermes has a largely tropical distribution, eight species
provide a characteristic element of the fauna in the semi-deserts and
deserts of the American Southwest. They, and a few species in the
closely related genus Gnathamitermes, form a group commonly known
bare as desert termites. The eight species have been described since
die turn of the century, and seven, as we presently know them, by S. F.
light (1930a,b; 1932). Snyder (1949) presented detailed synonymies
and further references. Except for the biological and ecological notes
iicluded with the original descriptions almost nothing has been pub-
lished on them. Although at least two species, Amitermes minimus
light and A. wheeleri (Desneux), are of significant economic impor-
tance (Snyder, 1954), the challenging questions on their life histories,
labits and ecology which Light (1930a, p. 173) asked about the Amer-
ican species still remain unanswered. The reasons for this continuing
gap in our knowledge should be especially apparent to those who have
attempted to collect or study them. Most of the species occupy very
soarsely populated regions and, being cryptobiotic in the extreme, are
tlius rarely seen and even more rarely collected. As explained below,
many of them have unorthodox flight seasons and periods which
farther limit their notice by man (Fig. 1) .
The following distribution records, approximate elevations, and notes
are intended as a guide for those who may be interested in contributing
^ Journal Paper No. 1459 of the Arizona Agricultural Experiment Station. Supported in part by
S ate Research Project No. 461 and a research grant from the National Pest Control Association for
travel in Mexico.
T he Pan-Pacific Entomologist 45 : 320-325. October 1969
OCTOBER 1969] NUTTING — RARE DESERT AMITERMES
321
Amitermes J FMAMJ JASOND
Fig. 1. Limits of the flight seasons of North American species of Amitermes,
based mainly on records of the last ten years at Tucson, Arizona. Flights occur
during the winter and summer rainy seasons. Although the alate of A. snyderi is
unknown, its most probable flight season is suggested on the basis of negative
evidence. See notes in text under this species. Flight periods occur as follows;
D, day; E, evening twilight; M, morning twilight.
to the meager information on the group. Previous records have heen
taken from Light (1930a, b; 1931; 1932) and Snyder (1954). New
records, mainly distributional limits in Arizona, are added from col-
lections made by various members and friends of the Department of
Entomology at the University of Arizona.
Amitermes coachellae Light. Previous records. — California:
Riverside Co., upper end Coachella Valley, 1200-1800 ft. (Colorado
Desert) . Sandy desert and washes in dead roots and casual buried
wood; build carton-lined tunnels and carton-like underground nest.
Nevada: Clark Co., Boulder City, 2480 ft.; Las Vegas, 2030 ft. (Mojave
Desert). At Boulder City, under stone in sandy wash. Arizona: Pima
Co., suburban Tucson, 2400 ft. (Sonoran Desert) ; flight about noon,
20 February 1929.
New records. — Arizona: Yuma Co., Yuma, ca. 150 ft. (Sonoran
Desert) ; flight during day, 2 March 1959. Pima Co., Tucson. Speci-
mens collected from five flights, probably of this species, between 27
January and 25 May 1935—1968. Flights generally occurred on sunny
days following 0.1 inch or more of rain, and at least two of them were
concurrent with flights of A. enter soni Light.
Amitermes emersoni Light. Previous records. — California: River-
side Co., Coachella Valley, 240 ft. below sea level-640 ft. (Colorado
Desert). Varied situations in a winter corn patch, a fallow field, a ball
park; in roots of dead Bermuda grass, small dead tree trunks, desert
322
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
’willow mounds (Chilopsis) , and in and along dry river bed. Flights in
Jmuary and February on sunny days following about 0.1 inch or more
of rain.
New records. — Arizona: Yuma Co., Yuma, 4 mi. S.E., 200 ft. (So-
noran Desert). In sandy desert under cardboard (Emerson, 1960).
lima Co., Tucson, 2400 ft. Several colonies in suburban area — side-
walks, yards, etc. — which was originally creosote bush desert. Many
flights observed between 21 January and 17 April 1960-1968. Flights
usually took place on sunny days following 0.1 inch or more of rain.
Amitermes minimus Light. This is one of the most widespread and,
ill some places, the most abundant species. Previous records. — Califor-
nia: Several southeastern localities, mainly in the Colorado and Mojave
Eeserts. Nevada: Clark Co., near Las Vegas. Arizona: Pima Co.,
Tucson. Texas: Bexar Co., San Antonio, 660 ft., flight on 12 July at
0315 hrs.; Cameron Co., Brownsville, 40 ft. In a variety of situations
such as posts, poles, dead parts of shrubs, etc.; builds hard, crusty
g alleries over buried wood.
New records. — Arizona: Colonies have been found at several local-
ities in the Tucson area (Pima Co.), north to 6 mi. W. Oracle, 3800 ft.
(Pinal Co.) (Emerson, 1960) ; and south to 5.5 mi. W. Pena Blanca,
4400 ft. (Sta. Cruz Co.) . In Tucson flights have been recorded between
7 July and 4 September 1962-1968. They have usually occurred on
smny mornings following fairly heavy showers.
Amitermes pallidus Light. A colony has not yet been reported.
Irevious records. — Arizona: Pima Co., Sabino Can., Sta. Catahna Mts.,
2300 ft. (Sonoran Desert) ; numerous collections of alates taken at
1 ght, June-August.
New records. — Arizona: Pinal Co., Casa Grande, 1400 ft. ; Pima Co.,
Tucson. As many as 14 flights have been recorded in one season,
letween 26 June and 30 September 1933-1968. They occur during
evening twilight and are not always associated with rain. Mexico: 10
nd. S. Canipole, Baja California del Sur, in light trap, 28 August 1959;
]0 mi. E. Navojoa, Sonora, at UV light, 13 August 1959 (both Sonoran
Desert, low elevations).
Amitermes parvulus Light. Previous records. — Arizona: Pima
C o., Sabino Can., Sta. Catalina Mts., 2800 ft. ; three alates taken at lights
ill July. Texas: Bexar Co., San Antonio, a nest containing alates, 8
October 1921.
New records. — ^Apparently none. Since this species has probably not
been taken again in Arizona, the original specimens may warrant re-
examination. It is possible that they have been confused with A. pal-
OCTOBER 1969] NUTTING RARE DESERT AMITERMES
323
lidus, the only other Amitermes known to fly in the evening in Arizona.
Amitermes silvestrianus Light. Previous records. — California:
Riverside Co., upper end Coachella Valley, 240-720 ft. (Colorado Des-
ert). Dry riverbed and washes, in dead wood on ground; build thick,
encrusting galleries over buried wood. Flight at Palm Springs, before
sunrise, 3 August two days after heavy rain.
New records. — Arizona: Yuma Co., near Yuma, 150 ft., colony under
corrugated cardboard (Emerson, 1960). Pima Co., Tucson and vicinity.
Several flights have been recorded between 17 July and 27 September
1961—1968. Flights always occurred within the hour before sunrise, on
mornings following heavy showers.
Amitermes snyderi Light. The winged form has never been re-
ported. Previous records. — California: Apparently this species has been
taken at only six localities between 1000-2000 ft., all in the Colorado and
Mojave Deserts of southeastern California. It was found in miscellane-
ous buried wood and the roots of trees and shrubs.
New records. — Arizona: Yavapai Co., Rock Springs, 2000 ft., 12
July 1962, had built much carton work into a stump in a dry wash.
Pima Co., Tucson, 2500 ft., March 1961, working manure buried for new
lawn. Molino Basin, Sta. Catalina Mts., 4400 ft., March, August and
October 1961, all under large, deeply set stones. Brown Can., Babo-
quivari Mts., 4050 ft., 4 August 1961; under buried wood. (All Sonoran
Desert.)
It is puzzling why no alate has yet been found as even a possible match
for the very large soldiers of this species. There are apparently not
even any likely candidates, for practically all of the Amitermes flying
in southern Arizona can be assigned to described species. Considering
the size of the soldiers, the alate might be expected to be larger than any
of the other southwestern species. It is therefore suggested that the alate
of A. snyderi may be flying concurrently with some large, common
termitid, such as Gnathamitermes perplexus (Banks), and is thus pass-
ing completely unnoticed. Collections from flights, supposedly of
Gnathamitermes^ should be carefully checked to see whether this might
be the case. These flights occur during or after summer rainstorms,
either in the daytime or early evening.
Amitermes wheeleri (Desneux). Previous records. — This species
has the widest distribution known to date, from Los Angeles Co., Cali-
fornia, south to Colima, Colima, Mexico, and east to Brownsville, Texas.
It is rather commonly found in cow chips, stumps, dead trees and desert
plants, as well as in posts, poles and the woodwork of buildings.
New records. — Arizona: Many colonies have been found on the
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
3:!4
desert and in moimtain canyons of the southeastern part of the state
between 1600 and 5600 ft. Flight records in this area extend from 3 July
to 21 August 1956-1968. All of the well-documented flights are known
tc have occurred within the hour before sunrise on mornings following
heavy showers. Many of them have been concurrent with flights of A.
silvestrianus. Specimens from flights of this species which have been
reported for other times of day should probably be re-examined. Mex-
ico: Among several other records, the following collections of colonies
si ow that this species is widely distributed over the central plateau:
49.5 mi. N.N.W. Gomez-Palacio, 3800 ft., Durango ; 32.4 mi. N. Aguas-
c< lientes, 6500 ft., Aguascalientes ; 32.8 mi. N.E. San Luis Potosi, 5250
fl,, S.L.P. ; and Ajijic, 5100 ft., Jalisco.
Discussion
It should be obvious from the slow accumulation of new records and
tl.eir fortuitous distribution that every opportunity should be taken to
olDtain members of this genus with complete biological information
ac companying each collection. Much more work needs to be done in
Aexico, since present evidence, particularly on the distribution of A.
pillidus and A. wheeleri, suggests that most, if not all, of the others will
li vely be found over many of the desert areas in the northwestern part
of that country. The most valuable records will be those made during
the flight season when workers, soldiers and winged forms may be
found together in the colonies.
Identification of soldiers and especially of alates is difficult unless one
it familiar with termites and has a reference collection available for com-
parisons. Most of the species are small, they are best handled in alcohol,
aid careful measurements must be used in conjunction with other
cliaracters. A well-illustrated, regional revision should simplify this
tfsk considerably. Toward this end it should be noted that approxi-
mately 200 collections of Amitermes are available for study in the in-
sect collection of the Department of Entomology at the University of
/ rizona. The author is willing to act as a temporary agent for receiving
further collections until such a task is undertaken. The time of flight
should prove extremely useful as a specific behavioral characteristic
and, with this in mind, a series of studies is planned, relating flight
activities of each species to seasonal meteorological conditions.
Literature Cited
Emerson, A. E. 1960. Personal communication.
Light, S. F. 1930a. The California species of the genus Amitermes Silvestri
(Isoptera). Univ. Calif. Publ. Entomol., 5: 173-214.
OCTOBER 1969]
BOOK REVIEW
325
1930b. The Mexican species of Amitermes Silvestri (Isoptera). Univ. Calif.
Publ. Entomol., 5: 215-232.
1931. The termites of Nevada. Pan-Pac. Entomol., 8: 5-9.
1932. Contribution tow^ard a revision of the American species of Amitermes
Silvestri. Univ. Calif. Publ. Entomol., 5: 355-414.
Snyder, T. E. 1949. Catalog of the termites (Isoptera) of the world. Smithson.
Misc. Collect., 112 : 490 pp.
1954. Order Isoptera, the termites of the United States and Canada. Tech.
Bull. Nat. Pest Control Assoc., New York. 64 pp.
BOOK REVIEW
Ecology of Aphidophagous Insects. Edited by Ivo Hodek and M. L. Dutkova.
1966. Dr. W. Junk Publishers, The Hague, The Netherlands. 360 p. + 10 plates.
Approximately $14.00.
The book presents the proceedings of a symposium held in Lihlice (near
Prague), Czechoslovakia from September 27 to October 1, 1965. The editor
states “In 1964, at the International Congress of Entomology, it was pointed out
that large Congresses fulfill other aims and that research workers engaged in
studying natural enemies of aphids, for example, should meet in smaller groups
to discuss their results and hypotheses.” The symposium was initiated to fulfill
this need. Specialists from 18 countries presented 74 papers during the course
of the five day symposium.
The book is divided onto 6 sections corresponding to those of the Symposium:
I, Food Ecology of Aphidophagous Insects; II, Voltinism and Arrest of Develop-
ment in Aphidophagous Insects; III, Behavior of Aphidophagous Insects and of
Aphids; IV, Distribution of Aphidophagous Insects in Habitats; V, Population
Dynamics of Aphids and Their Natural Enemies; VI, Aphidophagous Insects in
Biological and Integrated Control. The opening paper in each section is a review
of earlier published material or as in sections HI and IV, an introductory paper.
The last paper of each section presents a summary and general discussion of the
preceding papers to compensate for the subjective approach of individual authors.
The papers of the symposium have been condensed, overlapping sections deleted,
and data often summarized into tables and graphs in order that a concise manual
could be published. The editor reports that the personal views of the authors
have not been altered even if controversial or conflicting, but they receive
comments in the closing discussion paper of each section. Most of the papers
deal with aphid predators, especially coccinnelids, and to a lesser extent with
syrphids and chrysopids. There are areas where the coverage is rather scarce
(i.e. hemipterous predators), however, the editor points out that this is solely
a reflection of the activity of research workers attending the symposium. It is
also brought out that knowledge of voltinism and of Neuroptera is lacking. There
are occasional sentences whereby phrasing or wordiness may cause the reader
to falter, but these present no difficulty in interpretation.
The Pan-Pacific Entomologist 45: 325-326. October 1969
3^56
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
Each person, from the specialist to the beginning student that is concerned in
some respect with food ecology, behavior, distribution, population dynamics, and
the applied aspects of biological and integrated control of aphids, will find this
compendium an invaluable reference. One of the true values received from the
book is the cosmopolitan make up of the contributors and in the scope of the
material presented. The last section (VI) should be of particular interest to
thase workers concerned with biological and integrated control.
Dr. Ivo Hodek, the scientific editor, is to be commended for his efforts in
in tiating, directing the symposium, and summarizing the papers into the final
form. The foreign language editor, M. L. Dutkova, deserves far more than
passing praise for the translation of the many papers into English. Joseph
Pichal also contributed substantially to the proceedings by his space saving graphic
designs, enabling the reader to easily interpret data. Recognition is certainly
die to the Czechoslovak Academy of Sciences for the financial support of both
the symposium and the publishing of this book. — Robert G. Simpson, Colorado
Slate University, Fort Collins.
PAGE CHARGES
Due to increased costs of publishing the Pan-Pacific Entomologist
the normal page charge has been raised to $18.00. This new charge
w 11 go into effect 1 December 1969. All manuscripts received after
this date will be subject to the new charge. Authors without institutional
oi grant support are reminded that no page charges will be assessed if
they mark the appropriate box (Society grant desired) on the reprint
o] der form. — Editor.
OCTOBER 1969]
INDEX TO VOLUME 45
327
INDEX TO VOLUME 45
Addition, 80
Akre, Vatesus behavior, 269
Alexander, new crane fly subfamily, 51
a new crane fly, 291
Arachnida
Acari, Eviphididae, 318
Opiliones, Erebomastridae, 35
Opiliones, Ischyropsalidae, 73
Scorpionida, Vejovidae, 285
Arnaud, Edwin R. Leach — ninetieth
birthday, 66
Note in memoriam — Elwood Wendell
Molseed, 69
note on flight of Vanessa cardui, 69
note on Compsilura concinnata reared
from Agraulis vanillae incarnata,
77
note on Enlinia in the western Nearc-
tic, 79
recent literature, 243
Ashlock, Usinger bibliography and pro-
posed names, 185
Bibliography, Usinger, Robert Leslie,
185
Biography, Williams, Francis Xavier,
135
Usinger, Robert Leslie, 167
McKenzie, Howard Lester, Jr., 247
Blackwelder, nature of taxonomic data,
293
Blanton, new Nearctic Paradasyhelea,
97
Bohart, G. E., new Dufourea from
California, 57
Bohart, R. M., new Psenini, 216
Howard Lester McKenzie, Jr., 1910-
1968, 247
Book reviews, 241, 325
Borden, Notonecta ecology, 20
Briggs, a new phalangid family, 35
note on cave adaptation in phalangids,
73
Buckett, Tynomma revision, 204
Bugbee, new Eurytoma from California,
60
Butler, Cactobrosis development, 266
Caloptilia ovatiella, 260
diversilobiella, 262
Catachlorops (Psalidia) medemi, 149
Chemsak, Astromula nitidum habits, 81
Tillomorphine cerambycids, 303
Clark, note on new record of Labidura
rip aria, 64
Coleoptera
Bruchidae, 54, 237
Cantharidae, 119
Cerambycidae, 81, 282, 303
Nitidulidae, 100
Staphylinidae, 269
Colopterus testaceus, 101
Cordulegaster deserticola, 128
Correction and addition, 80
Cruden, a new Cordulegaster, 126
Cryptomaster leviathan, 41
Dailey, California cynipid synonymy,
132
Dermaptera
Labiduridae, 64
Dictya fontinalis, 222
Diplopoda
Chordeumida, Lysiopetalidae, 204
Diptera
Asilidae, 67
Bibionidae, 74
Cecidomyiidae, 16
Ceratopogonidae, 97
Dolichopodidae, 79
Sciomyzidae, 152, 222
Tabanidae, 147
Tachinidae, 77
Tipulidae, 1, 116, 229, 291
Trichoceridae, 51
Dufourea macswaini, 57
Edwards, book review, 241
Ellis, Notonecta ecology, 20
Ephemeroptera
Baetidae, 14
Leptophlebiidae, 103
Eplophorus auricomis, 309
aspericollis, 311
Erebomastridae, 37
Esenbeckia filipalpis divergens, 147
(Richardoa) potrix, 147
Euderces howdeni, 314
nelsoni, 315
Eurytoma foligalla, 60
flavifacies, 62
californica nana, 63
Fender, new California Malthodes, 119
Fisher, correction and addition, 80
two new Sepedon, 152
a new Dictya, 222
Gagne, R. J., review of genus Walsh-
omyia, 16
Gagne, W. C., note on distribution
Nearctic Acalypta, 68
Gardner, Tynomma revision, 204
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 45, NO. 4
3i!8
Gillogly, notes on Nitidulidae, 100
Gorelick, note on two species of Cal-
lophrys in California, 76
G issell, new Psenini, 216
Gustafson, note on rearing Coptodisca
arbutiella, 75
note on salt marsh insects, 78
Hinson, Utaperla revision, 26
H emiptera
Notonectidae, 20
Tingidae, 68
Horning, note on Callanthidium in
Idaho, 239
H )rmenoptera
Apidae, 87
Cynipidae, 132
Eurytomidae, 60, 112
Halictidae, 57
Megachilidae, 239
Orussidae, 73
Sphecidae, 216
Hmes, immature stages of Arctoconopa,
1
immature stages Gonomyodes, 116
immature stages Rhabdomastix, 229
IsDptera
Termitidae, 320
Jensen, A new Pseudocloeon from Idaho,
14
Johnson, C. D., Horn’s Bruchidae type-
material, 54
note on loeation of two bruchid holo-
types, 237
Johnson, P. H., Cactobrosis develop-
ment, 266
K awasemyinae, 51
Lane, note on salt marsh insects, 78
Lepidoptera, 74
Gelechiidae, 238
Gracilariidae, 259
Heliozelidae, 75
Lycaenidae, 76
Nymphalidae, 69, 77
Pyralidae, 266
Tortricidae, 70
Linsley, Robert Leslie Usinger, 1912-
1968, 167
Malthodes bissellarum, 119
arieticornus, 121
bifurculus, 122
immodestus, 123
angustifurcus, 124
Mayo, Thraulodes speciosus nymph, 103
Miller, Howard Lester McKenzie, Jr.,
1910-1968, 247
Nelson, Utaperla revision, 26
Nutting, rare desert Amitermes, 320
Obituary, see Biography
Odonata
Cordulegasteridae, 126
Opler, note on leaf-mining Lepidoptera
of California, 74
two new Caloptilia, 259
Orth, two new Sepedon, 152
a new Dictya, 222
Orthoptera
Gryllacrididae, 4
Page charges, 326
Paradasyhelea olympiae, 98
Philip, new Neotropieal Tabanidae, 147
Plecoptera
Chloroperlidae, 26
Powell, note on Cyrtopogon vanduzeei
behavior, 67
note on occurrence of Cacoeciamorpha
pronubana in Oregon, 70
note on oviposition behavior in Orus-
sus oceidentalis, 73
Proceedings, Pacific Coast Entomologi-
cal Society, 65
Pseneo leytensis, 220
irwini, 221
Psenulus mayorum, 216
aztecus, 219
Pseudocloeon edmundsi, 14
Ralston, note on new record of Labi-
dura rip aria, 64
Recent literatxire, 243
Rentz, Stenopelmatus synopsis, 4
Rotramel, note on orientation and eoupl-
ing in Dilophus orbatus, 74
Sepedon capellei, 157
pseudarmipes, 159
Simpson, book review, 825
Slobodchikoff, book review, 242
Society Notices, 244
Stenopelmatus nigrocapitatus, 12
Stevens, note on Exoteleia burkei in
Sierra Nevada, 238
Struble, gall wasps in pines, 112
Stypommisa (Styphocera) antennina,
151
Syntropis longiunguis, 286
OCTOBER 1969]
INDEX TO VOLUME 45
329
Tetranodus rugipeniiis, 306
angulicollis, 307
Theromaster, 49
Thorp, Bombus vandykei identity, 87
Tinkham, Stenopelmatus synopsis, 4
Tipula (Trichotipula) breedlovei, 291
Torgerson, Vatesus behavior, 269
Turner, note on oviposition behavior in
Orussus occidentalis, 73
Tynomma magnum, 207
Utaperla orientalis, 32
Van Volkinburg, new mite habitat, 318
Walshomyia cupressi, 19
Wickman, cedar and fir tree borers, 282
Wilkey, Howard Lester McKenzie, Jr.,
1910-1968, 247
Williams, a new Mexican Syntropis, 285
Wirth, new Nearctic Paradasyhelea, 97
Zimmerman, Francis Xavier Williams,
1882-1967, 135
Zoological Nomenclature, 56, 240
MAILING DATES FOR VOLUME 45
No. 1 16 April 1969
No. 2 18 July 1969
No. 3 14 October 1969
No. 4 8 December 1969
Published by the
Pacific Coast Entomological Society
in cooperation with
The California Academy of Sciences
VOLUME FORTY-FIVE
1969
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
1969 1970
H. V. Daly, Chairman D. D. Linsdale P. D.
W. E. Ferguson D. P. Furman R. F.
1971
Hurd, Jr.
Wilkey
San Francisco, California
1969
11
CONTENTS FOR VOLUME 45
itkre, R. D. and R. L. Torgerson
Behavior of Vatesus beetles associated with army ants 269
/dexander, C. P.
A new subfamily of winter crane flies 51
A new species of crane fly associated with the plant genus
Lopezia in Mexico 291
Arnaud, P. H., Jr.
Edwin R. Leach — ninetieth birthday 66
In memoriam — Elwood Wendell Molseed 69
A 1968 flight of Vanessa cardui Linnaeus in San Francisco and
Pacifica, California 69
Compsilura concinnata (Meigen) reared from Agraulis vanillae
incarnata (Riley) at San Leandro, California 77
Occurrence of the genus Enlinia Aldrich in the western Nearctic 79
Recent literature 243
Ashlock, P. D.
Robert L. Usinger bibliography and list of names proposed 185
llackwelder, R. E.
The nature of taxonomic data 293
Bohart, G. E,
A new species of Dufourea from California 57
Bohart, R. M. and E. E. Grissell
New species of Psenini 216
Briggs, T. S.
A new Holarctic family of Laniatorid Phalangids 35
Cave adaptations in phalangids of the genus Taracus 73
Buckett, J. S. and M. R. Gardner
Revision of the chordeumid milliped genus Tynomma Loomis
from California 204
Bugbee, R. E.
New species of the genus Eurytoma from California
60
m
Butler, G. D., Jr., and P. H. Johnson
The development of Cactohrosis fernaldialis and Albareda para-
bates in relation to temperature 266
Chemsak, J. A.
Notes on the habits of Astromula nitidum Chemsak and Lindsley
with descriptions of the immature stages 81
Records and descriptions of Mexican and Central American
T illomorphini 303
Clark, W. H. and G. L. Ralston
Occurrence of Labidura riparia (Pallas) in Baja California,
Mexico 64
Cruden, R. W.
A new species of Cordulegaster from the Great Basin region of
the United States 126
Dailey, D. C.
Synonymy of Dryocosmus attractans (Kinsey) and Callirhytis
uvellae Weld 132
Edwards, J. G.
Book Review: Ecology and biogeography of high altitude insects 241
Ellis, R. A. and J. H. Borden
Effects of temperature and other environmental factors on
Notonecta undulata Say 20
Fender, K. M.
New California Malthodes and distribution notes on other species
in California 119
Fisher, T. W.
Correction and addition 80
Fisher, T. W. and R. E. Orth
Two new species of Sepedon separated from S. armipes Loew
in western North America 152
A new Dictya in California with biological notes 222
Gagne, R. J.
A review of the genus Walshomyia including a new species from
Cupressus galls in California 16
IV
(ragne, W. C.
New distribution records of two Nearctic Acalypta Westwood 68
(iillogly, A. R.
Taxonomic notes on Nitidulidae of California 100
(jorelick, G. A.
Callophrys dumetorum (Bdv.) and Callophrys viridis (Edw.)
in California 76
Gustafson, J. F.
Rearing observations on the madrona shield bearer Coptodisca
arbutiella 75
Horning, D. S.
First recorded occurrence of the genus Callanthidium in Idaho
with notes on three nests of C. formosum (Cresson) 239
Hynes, C. D.
The immature stages of Arctoconopa carbonipes (Alex.) 1
The immature stages of Gonomyodes tacoma Alex 116
The immature stages of the genus Rhabdomastix 229
Jensen, S. L.
A new species of Pseudocloeon from Idaho 14
Johnson, C. D.
Horn’s Bruchidae type-material in the Ulke collection 54
The location of the holotypes of Bruchus cubiculus Casey and
Mylabris wheelocki Blatchley 237
Lane, R. S. and J. F. Gustafson
Salt marsh insects 78
Linsley, E. G.
Robert Leslie Usinger, 1912-1968 167
vlayo, V. K.
Nymph of Thraulodes speciosus Traver with notes on a symbiotic
Chironomid 103
Miller, D. R., R. M. Bohart, and R. F. Wilkey
Howard Lester McKenzie, Jr., 1910-1968 247
V
Nelson, C. H. and J, F. Hanson
The genus Utaperla 26
Nutting, W. L.
Distribution and flights of rare North American desert termites
of the genus Amitermes 320
Opler, P. A.
Status of knowledge of the leaf-mining Lepidoptera of California 74
Two new species of Caloptilia associated with Rhus in California 259
Page charges 326
Philip, C. B.
New or little-known Neotropical Tabanidae 147
Powell, J. A.
Prey hunting and courtship of Cyrtopogon vanduzeei Wil. & Mart. 67
Occurrence of Cacoeciamorpha pronubana (Hbn.) in Oregon 70
Powell, J. A. and W. J. Turner
Observations on oviposition behavior in Orussus occidentalis 73
Proceedings, Pacific Coast Entomological Society 65
Rotramel, G.
Orientation and coupling in Dilophus orbatus 74
Simpson, R. G.
Book Review: Ecology of aphidophagous insects 325
Slobodchikoff, C. N.
Book Review : Six-legged science 242
Society Notices 244
Stevens, R. E.
Occurrence of Exoteleia burkei in the Sierra Nevada 238
Struble, G. R,
Gall wasp infestations in forest trees, chiefly pines, of California 112
Thorp, R. W.
The identity of Bombus vandykei 87
VI
Tinkham, E. R. and D. C. Rentz
Notes on the bionomics and distribution of the genus Steno-
pelmatus in Central California with the description of a
new species 4
^i^an Volkinburg, D.
Thinoseius spinosus found in new and unusual habitat 318
Hickman, B. E.
A crossbreeding study of the cedar tree borer, Semanotus ligneus
amplus, and the fir tree borer, S. litigiosus 282
Williams, S. C.
A new species of Syntropis from Baja California Sur, Mexico
with notes on its biology 285
A^irth, W. W. and F. S. Blanton
A new Nearctic species of the genus Paradasyhelea Macfie 97
Zimmerman, E. C.
Francis Xavier Williams, 1882-1967 135
Zoological Nomenclature 56, 240
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Entomological Exchanges and Sales ix
Publications of the Society x
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problems for close-to-harvest uses.
Look for Cythion, the premium-grade malathion,
on pesticides for agricultural, home and garden,
pet, stored product, ULV and public health use-
in liquids, wettable powders, dusts, aerosols and
low-pressure bombs. Cythion is your assurance of
the highest quality.
AMERICAN CYANAMID COMPANY
PRINCETON, NEW JERSEY
Before using any pesticide^ stop and read the label.
A Service for the Field
Testing & Evaluation
of
Agricultural Chemicals
Under Western Conditions
Entomology
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Engineering
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Trojan Laboratories
Box 518
Montebello, California 90640
(213) 721-9574
IV
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. . . Kills grape leafhoppers.
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Write for technical information and recommendations.
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THE ALCOA BUILDING • ONE MARITIME PLAZA • GOLDEN
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Here s an effective way to control tomato
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Th iricide 90TS, the microbial insec-
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•THUE CIDE is a trademark of Bioferm Division,
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And on forest trees and ornamentals for
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If your dealer doesn’t stock Thuricide
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Los Angeles, Fresno, Bakersfield, Cali-
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Stauffer
Read the label, heed the label and GROW WITH STAUFFER CHEMICALS
VI
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COMMERCIAL AND RESEARCH
ENTOMOLOGISTS
Obtain more accurate population counts
with a light-weight motor-fan unit
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D-WAC CO.
P. O. Box 2095
RIVERSIDE, CALIFORNIA
Vll
ENTOMOLOGICAL
RESEARCH INSTITUTE
Box 375 Lake City, Minnesota 55041
IXATION FOR:
Schmitt-type Field Box
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FOAM PINNING BOTTOM
California Academy-type glass-topped drawer
U.S.N.M.-type glass-topped drawer
Philadelphia Academy-type glass-topped drawer
Cabinet to hold 12 of above types of drawers
SPECIAL SIZED DRAWERS, BOXES, CABINETS CUSTOM MADE
TO YOUR LIKING.
V^rite for information and prices.
INSECT MUSEUM SUPPLY
1021 8th Ave. So., Moorhead, Minnesota— Jack R. Powers
L OCATION FOR:
CUSTOM PRINTED PIN LABELS — printed with 4 pt. type on 100% cotton fiber
ledger paper (colored papers and PRICE CARD on request)
Elephant brand INSECT PINS — Excellent quality — Sizes 00, 0, 1,2, 3, 5 —
$4.25 per 1000
/V inutien nadein — $1.00 per 500 pkg.
Custom Printed GUMMED MICROSCOPE SLIDE LABELS — $4.50 per 1000
G ASS TOPPED INSECT DISPLAYS FOR TEACHING
DFIAWER LABELS, MAMMAL TAGS, ALCOHOL LABELS, Printed to your
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Vlll
Every product
bearing the Blue Bullseye
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You can count on it.
At Chemagro, test tube to test plot to market
is a never ending process. As makers of chemicals
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keep scientists very busy in the laboratories, testing,
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making the Blue Bullseye a meaningful trademark.
Look for it when you need an insecticide, miticide,
fungicide or herbicide. EB71 a
CHEMAGRO
CORPORATION
KANSAS CITY, MISSOURI 64120
ENTOMOLOGICAL EXCHANGES AND SALES
This page is available to members and non-members who wish to advertise
entomological 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
tl e Advertising Manager, W. H. Lange, Department of Entomology, University of
California, Davis, California 95616.
Besearch Request. — Am nearing completion a study of the Microlepidoptera as-
sociated with oaks in California. Would appreciate the opportunity to borrow any
n aterial associated with oaks or to receive any biological information on these
species. Appropriate acknowledgments accorded. P. Opler, Division of Ento-
n ology, University of California, Berkeley, Calif. 94720.
SvN Francisco on a Budget? Charming, centrally located. Hotel Beresford, 635
Satter Street.
Zdologist going to Central and South America undertakes all collectings.
Ji rite: M.de Munter 1040 Chaussee d’Alsemberg, Bruxelles 18 Belgium
L \RGID AND Pyrrhocorid Hemiptera — ^Will purchase or trade. T. F. Halstead, 1523
^ . Colter #25, Phoenix, Ariz. 85015.
Insect Pins. — Original Karlsbader — Brass Heads. Sizes 00-4 $5.20/1000 Post Paid.*
^Tite for Quantity Discount. Prompt Delivery. R. B. Hewitt, Insect Sales, 928-46th.
S ;., Sacramento, Calif. 95819. * Calif. Residents Please Remit 5% Sales Tax.
Help Wanted
Economic Entomologist — $829-1007 (starting salary commensurate with ex-
perience) Centrally located, San Joaquin County offers easy access to recreational.
Cl dtural and academic facilities and has the fifth highest county agricultural income
ir the nation. Excellent fringe benefits and diversified career growth opportunity.
Bequires college degree in entomology and three years economic entomology
e iperience. Apply to Wilden H. Tweedie, Personnel Dept., 121 East Weber Avenue,
S ockton, California 95202.
PUBLICATIONS
OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY
PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL
SOCIETY.
Vol. 1 (16 numbers, 179 pages) and Vol. 2 (numbers 1-9, 131
pages) . 1921-1930. Price $2.50 per volume.
PAN-PACIFIC ENTOMOLOGIST.
Vol. 1 (1924) to present. Price $6.00 per volume of 4 numbers,
or $1.50 per single issue.
MEMOIR SERIES.
Volume 1. The Sucking Lice by G. F. Ferris. A 320-page book
which summarizes the knowledge of the Anoplura of the world.
Published October 1951. Price $6.00. (Plus 350 postage.) *
Volume 2. The Spider Mite Family T etranychidae by A. Earl
Pritchard and Edward W. Baker. This worldwide treatment deals
with the systematics, identification, and economics of the “Red
Spiders” and includes descriptions of 33 new species. 472 pages.
Published July 1955. Price $10.00. (Plus 500 postage.) *
Volume 3. Revisionary Studies in the Nearctic Decticinae by David
C. Rentz and James D. Birchim. This 173-page revision of Nearctic
shield-back katydids includes descriptions of one new genus and
13 new species and subspecies. Published July 1968. Price $4.00.
(Plus 250 postage.)*
* (Add 5% sales tax on all California orders. Members of the Society -will
receive a 20% discount.)
Send orders to:
Pacific Coast Entomological Society
c/o California Academy of Sciences
Golden Gate Park
San Francisco, California 94118
J
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,
and distribution. Excess pagination must be approved and may be charged to the author.
Papers are published in approximately the order that they are received. Immediate publi-
cation may be arranged after acceptance by paying publication costs. Papers of less than
a printed page may be published as space is available, in Scientific Notes.
Manuscripts for publication, proof, and all editorial matters should be addressed to the
Editor, Robbin W. Thorp, Department of Entomology, University of California, Davis,
California 95616.
Typing. — All parts of manuscripts must be typed on bond paper with double-spacing and
ample margins. Carbon copies or copies on paper larger than 81/^ X H inches are not
accepted. Do not use all capitals for any purpose. Underscore only where italics are
intended in the body of the text, not in headings. Number aU pages consecutively and
put author’s name at the top right-hand corner of each sheet. References to footnotes
in text should be numbered consecutively. Footnotes should be typed on a separate
sheet.
First page. — The page preceding the text of the manuscript should include (1) the
complete title, (2) the order and family in parentheses, (3) the author’s name or
names, (4) the institution with city and state or the author’s home city and state if
not affiliated, (5) the shortened title (running headline) not to exceed 38 letters and
spaces when combined with the author’s last name or names, (6) the complete name
and address to which proof is to be sent.
Names and descriptions of organisms. — The first mention of a plant or animal should
include the full scientific name with the author of a zoological name not abbreviated.
Do not abbreviate generic names. Descriptions of taxa should be in telegraphic style.
References. — All citations in text, e.g., Essig (1926) or (Essig, 1958), should be listed
alphabetically under Literature Cited in the following format:
Essig, E. 0. 1926. A butterfly migration. Pan-Pac. Entomol.,
2 : 211 - 212 .
1958. Insects and mites of western North America. Rev.
ed. Tlie Macmillan Co., New York. 1050 pp.
Abbreviations for titles of journals should follow the list of Biological Abstracts, 1966,
47(21) : 8585-8601.
Tables. — Tables are expensive and should be kept to a minimum. Each table should be
prepared as a line drawing or typed on a separate page with heading at top and foot-
notes below. Number tables with Arabic numerals. Number footnotes consecutively
for each table. Use only horizontal rules.
Illustrations. — No extra charge is made for the line drawings or halftones. Submit only
photographs on glossy paper and original drawings (no photographs of drawings).
Authors must plan their illustrations for reduction to the dimensions of the printed
page (4% X 6% inches). If possible, allowance should be made for the legend to be
placed beneath the illustration. Photographs should not be less than the width of the
printed page. Drawings should be in India Ink and at least twice as large as the
printed illustration. Excessively large illustrations are awkward to handle and may
be damaged in transit. Include a metric scale on the drawing or state magnification
of the printed illustration in the legend. Arrange figures to use space efficiently.
Lettering should reduce to no less than 1 mm. On the hack of each illustration should
be stated (1) the title of the paper, (2) the author’s complete name and address, and
(3) whether he wishes the illustration and/or cut returned to him at his expense.
Cuts and illustrations not specifically requested will be destroyed.
Figure legends. — Legends should be typewritten double-spaced on separate pages headed
Explanation of Figures and placed following Literature Cited. Do not attach
legends to illustrations.
Proofs, reprints, and abstract. — Proofs and forms for the abstract and reprint order
will be sent to authors. Major changes in proof will be charged to the author. Proof
returned to the editor without the abstract will not be published.
Page charges. — All regular papers of one to ten printed pages are charged at the rate of
$18.00 per page. This is in addition to the charge for reprints and does not include the
possible charges for extra pagination or the costs for immediate publication. Private
investigators or authors without institutional or grant funds to cover this charge may
apply to the Society for a grant to cover the page charges. A mark in the appropriate
box (society grant desired) on the reprint order form will suffice as an application.
All articles are accepted for publication only on the basis of scientific merit
and without regard to the financial support of the author.
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ORTHO
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