Vol. 52 OCTOBER 1976 No. 4
THE
PAN-PACIFIC ENTOMOLOGIST
CLIFFORD—Observations on the life cycle of Siphloplecton basale (Walker)
(Ephemeroptera;/ Meiretopodidae)) 2) 0-0 aw se a 265
LINSLEY AND ROSS—Plant associations among adult Hippomelas (Coleop-
Leracm UPLestiducyi me. et get MAP WT ao, A a ee ee be Pe 272
ANDREWS—A new species of Alloxysta hyperparasitic on aphids associated
with South American Nothofagus forests (Hymenoptera: Cynipidae) . 286
DENNIS AND LAVIGNE—Oviposition of Cyrtopogon inversus (Diptera:
DAIS TMU ete ten = ot Bs Memes oh Medel fee athe neta ae 288
YOUNG—Studies on the biology of Heliconius charitonius L. in Costa Rica
wNymphalidacs Meliconiinae) 2242.6 4 fe ee 291
STONE—Notes on the biology of the introduced elaterid Conoderus exsul
(Shanp)> (@eleopteras. Blateridae) i008 304
MIDDLEKAUFF AND LANE—A new species of Apatolestes from California
GDinteramailabanidaeyy wt ee he eel be Re a Efe. “oil
EVANS—Bembicini of Baja California Sur: Notes on nests, prey and distri-
bution (iymenoptera: Sphecidae). -i8e 00008 314
POLHEMUS AND CHENG—A new Rheumatobates from Costa Rica
Chemupterac’ Germdae) co. 0 ee at al a ie ee SURO > eee 321
MARI MUTT—A new species of Heteromurus from the Solomon Islands
(Collembolaz Entomobryidae)! 326
BOHART AND VILLEGAS—Nesting behavior of Encopognathus rufiventris
Timberlake (Hymenoptera: Sphecidae) __ 331
GW YNNE—Observations on the nesting behavior of Ammophila nasalis
(Hymenopteras@ophecidac). = 8 ee ee ee ee 335
ANDREWS—A new species of Fuchsina Fall with notes on some California
Watnnidindac. <(@oleopteray' = © Wear. eg ae td es ae 339
SCIENEIFIC NOTES es -<Uec ch 271, 313, 323, 338, 347, 348
ITU TEDA GEST AS Teocas NCSI), ee Se et ea Nig A Oo oa te el oe 285
GE Nee ROACH Cie ee OE Foy ee 290, 330, 334
ZOOVO GICAL ANOMENCUATURE 20 29S ie Ne Ae 320
LID) a VO NC WIM 920 reas Leake es Le ot oe Oe AS 351
SAN FRANCISCO, CALIFORNIA e 1976
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
THE PAN-PACIFIC ENTOMOLOGIST
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STATEMENT OF OWNERSHIP
Title of Publication: The Pan-Pacific Entomologist.
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US. A
The Pan-Pacific Entomologist
VoL. 52 OcTOBER 1976 No. 4
Observations on the Life Cycle of Siphloplecton basale
(Walker )
(Ephemeroptera: Metretopodidae)
Hucu F. CLirrorp
Department of Zoology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E9
Metretopodidae is a small mayfly family placed in the major hepta-
geniod phyletic line, the metretopodids representing a special lineage
from the Siphlonurinae (Edmunds, 1972). Several workers, e.g. Berner
(1950), Burks (1953), have commented on metretopodids exhibiting
features in common with both Heptageniidae and Siphlonuridae.
Only two genera are now assigned to Metretopodidae. Metretopus, con-
taining a single North American species (M. borealis Eaton), is Hol-
arctic and mainly northern. Siphloplecton is Nearctic and contains
five species. Siphloplecton species occur east of the Cordillera
over much of North America, usually in slow-flowing streams but oc-
casionally in lakes. S. basale is the most widely distributed species, ap-
pearing to be most abundant in northern regions.
Detailed life cycle data for North American mayflies are rapidly ac-
cumulating, but there are apparently no complete studies for any
Siphloplecton species. This is especially surprising for the widely dis-
tributed S. basale, since fully grown nymphs are large, up to 20 mm,
and adults, being vigorous fliers with pronounced mottled wings, are
easily recognizable on-the-wing. The species is conspicuous enough to
be given a special name, Great Speckled Lake Olive, by fly fishermen
(Schwiebert, 1973). One reason for the paucity of information, espe-
cially nymphal data, is that S. basale populations are probably seldom
large. But another factor has to do with the habits of the nymphs. The
large streamlined nymphs are vigorous swimmers; they not only can
avoid standard netting devices, but if caught in anything other than
long deep nets they can swim against the current and out of the net. Col-
lecting medium and large size S. basale nymphs seems more comparable
Tue Pan-Paciric ENTomMoLoGiIst 52: 265-271. OctosBer 1976
266 THE Pan-PAciric ENTOMOLOGIST
to capturing small cyprinids than collecting typical bottom fauna. In
fact, Leonard and Leonard (1962) used a fine-meshed minnow seine to
collect S. basale.
Siphloplecton basale occurs in a small brown-water stream of Al-
berta, Canada, a stream that has been under continuous investigation
since 1966. Except for the initial exploratory study, in which a life
cycle approach was used to describe the fauna (Clifford, 1969), no
study was designed specifically to gather life cycle data on S. basale;
and only two S. basale nymphs were collected during the 1% years of
the initial study. However considerable information stemming from
other studies has now accumulated for S. basale. By treating the 10
years of S. basale data as an entity, S. basale’s life cycle can be described
throughout a complete year for the northern stream.
Stupy AREA
The North Fork of the Bigoray River, located in west-central Alberta
(53° 25’N, 115° 30’W), is part of the Arctic Ocean drainage. The
stream drains extensive muskeg-type terrain. The water color is dark
brown during the ice-free season, but the stream becomes quite
clear in late winter. At the sampling site, the stream is small (av-
erage base flows in winter and summer are 0.14 m?/s and 0.83
m?/s respectively), slow-moving (average gradient is 3.0 m/km) and
meandering. In late summer, aquatic macrophytes, especially Spar-
ganium, cover much of the substrate, which is mainly of sand and
silt with lesser amounts of small rubble. The Bigoray River can be
described as a chironomid-ostracod type stream, these two taxa col-
lectively comprising on a yearly basis about 49% of the total macro-
invertebrate fauna by numbers. Ephemeroptera is the third most abun-
dant taxon, making up 17% of the yearly bottom fauna. There are few
resident fish; but shortly after the ice goes out in spring, large num-
bers of white suckers, Catostomus commersoni (Lacépéde), make a
spawning run into the Bigoray River from the Pembina River.
LIFE CYCLE
Nymphs—Based on 5 years of bottom fauna samples, S. basale makes
up only a minor component of the total invertebrate fauna in the brown-
water stream. Nymphs made up 0.2% of the total yearly bottom fauna
and 1.3% of the total yearly mayfly bottom fauna. Siphloplecton
basale is a univoltine species and generations do not overlap (Fig. 1).
New generation nymphs first appear in July. They grow rapidly during
the remainder of the ice-free season. From July through November,
VoL. 52, No. 4, OctosperR 1976 267
=
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5 =
ow
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: ICE - FREE ee
JAN. FEB. MAR. APR. MAY JUNE JULY AUG. SEPT. OCT. NOV. DEC.
MONTH
Fic. 1. Growth features and emergence of S. basale as related to average 5-day
water temperatures and ice conditions. Range and mean size were compiled by
measuring nymphs (numbers above range) collected from 1969 through 1975 and
treated as monthly samples. Monthly sampling date is expressed as the average
of all collecton dates within the month.
nymphs increase in size at a rate of about 0.8 mm per week. Most
nymphs appear to be fully grown by November, when the stream be-
comes completely ice-covered. Nymphs grow very little, if at all, dur-
ing the long winter. Average ice break-up time at the sampling site is
during the second week in April, and shortly after this water tempera-
tures start to rise. Nymphs do not increase very much in total length
after the break-up; however development resumes, and the adults usu-
ally start emerging during the second or third week of May.
Nymphs smaller than 9 mm seemed to be distributed in a variety of
habitats including shallow, fast-moving water; whereas nymphs 9 mm
and larger were usually found only in slow-moving water having exten-
sive beds of aquatic macrophytes. Lyman (1956), Lehmkuhl (1970)
and Hilsenhoff et al. (1972) have all commented on S. basale nymphs
being most abundant in slow-water along the banks and usually as-
sociated with shoreline vegetation. In contrast, Leonard and Leonard
(1962) found S. basale in deep water having a strong current. In the
Bigoray River, Sparganium can extend from bank to bank, and the
268 THE Pan-PaciFic ENTOMOLOGIST
Table 1. Volume of stomach content items of S. basale nymphs. Vol-
umes were determined using a Model TA II Coulter Counter.
Average Volumes (mm*) per stomach
Average Size
Month Numbers (mm) Detritus Diatoms Sand and Silt Total
Feb.-March 4 16.0 0.66 0.02 0.01 0.69
May 9 16.5 0.62 0.14 0.02 0.78
large nymphs appeared to be distributed across an entire width transect,
but the nymphs are such vigorous swimmers that it was difficult to de-
termine accurately their preferred habitat. Lyman (1955) observed
S. basale migrating shoreward just prior to emergence in a Michigan
lake. During an April-May study of Leptophlebia cupida’s upstream
migration (Hayden and Clifford, 1974), we found no S. basale nymphs
moving upstream along the banks or even congregating along the shore.
During a year’s study of the drifting organisms (Clifford, 1972),
S. basale nymphs comprised a total yearly drift density of 304 per 100
m?® of water filtered. This was 4% of the total yearly mayfly fauna
found in the drift. Only 25% of the S. basale nymphs were larger than
11 mm. Eighty percent of all drifting S. basale nymphs were collected
between sunset and sunrise; hence S. basale does exhibit a night-active
drift pattern.
Nymphs are herbivore—detritivores (Table 1). The stomach analy-
sis, although based on a small sample size, indicates that by late win-
ter the nymphs are almost entirely detritivores; but during the ice-free
season prior to emergence, they are partly herbivorous, ingesting living
diatom cells. White suckers usually make their spawning run up the
Bigoray River at about the time S. basale starts emerging, but the
nymphs are not a major food item of the suckers (Bond, 1972).
Nymphs occurred in the stomachs of 9.5% of the suckers moving up-
stream from 28 April to 26 May 1969. These nymphs accounted for
less than 1.0% of the total number of food items, but 3.9% of the total
weight of the stomach contents. There were no S. basale nymphs in
stomachs of post-spawning suckers moving downstream between 24:
May and 4 June 1969, a period when S. basale is still emerging.
Adults—Siphloplecton basale is one of the earliest emerging may-
flies of the Bigoray River. Subimagos start emerging usually during
the second or third week of May, when the water, although still quite
cold, is warming rapidly (Fig. 1). Cumulative emergence percentages
of Figure 1 were compiled from data of a 1973 emergence study (Boerger
VoL. 52, No. 4, OcToBErR 1976 269
and Clifford, 1975), in which 16 floating box traps, each 0.1 m? in
area, were arranged in four transects across the stream; the traps were
operated continuously from 25 April to 23 October 1973. Total yearly
emergence for S. basale was 44.5 (18.6 males and 25.9 females) per
square meter of substrate, and this accounted for 13% of all mayflies
emerging during 1973. Assuming the emergence traps were not some-
how biased in favor of collecting disproportionately large numbers of
S. basale, this figure indicates that S. basale is more abundant than
would be indicated by bottom fauna samples (1.3% of the total yearly
mayfly fauna) or drift samples (4% of the total mayfly fauna).
Adults are present for about 1 month, most of the adults having
emerged by the middle of June. The timing of this phenophase in the
brown-water stream agrees well with S. basale emergence timing in
other northern regions: from 23 May to 12 June in Ontario (Clemens,
1915), from 12 May to early June in a Michigan lake (Lyman, 1955),
and in early May in Wisconsin (Hilsenhoff et al., 1972).
In the Bigoray River, S. basale emerges during full daylight, usually
around midday. In Michigan, Lyman (1955) observed a nymph trans-
forming at 1100 hrs and its subimago stage lasting 48 hours. Males
have been observed swarming in full daylight of the afternoon and
early evening (Clemens, 1915; Lyman, 1955) and full daylight of the
morning (present study). Males swarm above water, usually at a height
of 3 to 6 meters. Lyman (1955) gives a description of the male’s hover-
ing and darting type flight. Leonard and Leonard (1962) noted that
once the female enters the swarm and is siezed, the pair rises to tree-
top level. Spieth (1940) determined that the S. basale pair normally
remains coupled for no more than 1% minutes; however when captured
in a net, the pair might remain in copula for up to 7 minutes. I could
not confirm Leonard and Leonard’s (1962) observation that ovipositing
females sometimes drop their eggs while in flight. In the Bigoray River,
all ovipositing females appeared to release a few eggs at a time by mo-
mentarily resting on the water’s surface or less frequently by dipping
the abdomen into the water.
Koss and Edmunds (1974) have described S. basale’s egg. For Bigo-
ray River adults, average number of eggs per subimago (average size:
15.5 mm) was 2063 (Clifford and Boerger, 1974). Total potential fe-
cundity generally varied with the female’s size. For example, average
number of eggs per millimeter size class of subimagos and imagos (that
had not oviposited) combined were 1832 (13 mm), 1633 (14 mm),
2043 (15 mm), 2168 (16 mm), 2499 (17 mm), and 3019 (18 mm).
Females have a high total potential fecundity relative to that of most
270 Tue Pan-Paciric ENTOMOLOGIST
mayflies, but this is because S. basale is a large mayfly. Average num-
ber of subimago eggs per millimeter of total length was 133, which is
about in the middle of the range of all Bigoray River mayflies. The ovoid
shaped eggs had an average length of 0.21 mm and average width of
0.14 mm.
I conclude from the 10 years of data that S. basale is probably more
abundant in streams and lakes than would be indicated by bottem fauna
samples, since the large fishlike nymphs can often avoid standard col-
lecting devices. In terms of the Bigoray River’s total yearly mayfly
fauna, S. basale made up only 1.3% of the bottom fauna and 4% of the
drift fauna, but the subimagos accounted for 13% of all mayflies
emerging.
ACKNOWLEDGMENTS
I would like to thank Dr. Lewis Berner, University of Florida, for
his review of the manuscript and Mr. Hal Hamilton, University of Al-
berta, for determining the stomach content volumes. Parts of this study
were supported through grants from the National Research Council of
Canada.
LITERATURE CITED
Berner, L. 1950. The mayflies of Florida. Univ. of Florida Press, Gainesville.
267 pp.
Boercer, H. anp CiirForp, H. F. 1975. Emergence of mayflies (Ephemeroptera)
from a northern brown-water stream of Alberta, Canada. Verh. Internat.
Verein. Limnol. 19: 3022-3028.
Bono, W. A. 1972. Spawning migration, age, growth, and food habits of the
white sucker, Catostomus commersoni (Lacépéde), in the Bigoray
River, Alberta. M.Sc. thesis, Dept. of Zool., Univ. of Alberta. 125 pp.
Burks, B. D. 1953. The mayflies, or Ephemeroptera, of [linois. Bull. [l. Nat.
Hist. Surv. 26: 1-216.
Ciemens, W. A. 1915. Rearing experiments and ecology of Georgian Bay Eph-
emeridae. Contrib. to Can. Biol., Sessional Paper, 396: 113-128.
Currrorp, H. F. 1969. Limnological features of a northern brown-water stream,
with special reference to the life histories of the aquatic insects. Am.
Midl. Nat. 82: 578-597.
Cuirrorp, H. F. 1972. A year’s study of the drifting organisms in a brown-
water stream of Alberta, Canada. Can. J. Zool. 50: 975-983.
CuirForb, H. F., anp Borercer, H. 1974. Fecundity of mayflies (Ephemeroptera) ,
with special reference to mayflies of a brown-water stream of Alberta,
Canada. Can. Entomol. 106: 1111-1119.
Epmunps, G. F., Jr. 1972. Beogeography and evolution of Ephemeroptera. Ann.
Rey. Entomol. 17: 21-42.
Haypen, W. anp Currrorp, H. F. 1974. Seasonal movements of the mayfly
Leptophlebia cupida (Say) in a brown-water stream of Alberta, Can-
ada. Am. Midl. Nat. 91: 90-102.
VoL. 52, No. 4, OcToBER 1976 271
Hitsennorr, W. L., Loneripvce, J. L., Narr, R. P., Tennessen, K. J., anp WAL-
ton, C, P. 1972. Aquatic insects of the Pine-Popple River, Wiscon-
sin. Tech. Bull. 54, Dept. Nat. Res., Madison. 42 pp.
Koss, R. W. ano Epmunps, G. F., Jr. 1974. Ephemeroptera eggs and their con-
tribution to phylogenetic studies of the order. Zool. J. Linn. Soc. 55:
267-349,
LeumkunL, D. M. 1970. Mayflies in the South Saskatchewan River; pollution
indicators. The Blue Jay 28: 183-186.
Leonarp, J. W. anp F. A. Leonarp. 1962. Mayflies of Michigan trout streams.
Cranbrook Inst. of Sci., Bloomfield Hills. 139 pp.
Lyman, F. E. 1955. Seasonable distribution and life cycles of Ephemeroptera.
Ann. Entomol. Soc. Amer. 48: 380-391.
Lyman, F. E. 1956. Environmental factors affecting distribution of mayfly
nymphs in Douglas Lake, Michigan. Ecology 37: 568-576.
ScHWIEBERT, E. 1973. Nymphs. A complete guide to naturals and their imita-
tions. Winchester Press, New York. 339 pp.
SpieTH, H. T. 1940. Studies on the biology of the Ephemeroptera. II. The nup-
tial flight. J. New York Entomol. Soc. 48: 379-390.
SCIENTIFIC NOTE
Leucopsis klugii (Hymenoptera, Chalcidoidea) reared from Xylocopa
brasilianorum (Hymenoptera, Apoidea) in Costa Rica.—A nest containing
19 Xylocopa brasilianorum (Linnaeus) was found in a slender fence post on 24
January 1975 near the Palo Verde Research Station, Comelco Property, about 15
km SW Bagaces, Guanacaste Province, Costa Rica. The nest entrance led to a
burrow 1.5 cm in diameter that extended lengthwise in the post for 7.4 cm in one
direction and 10.4 cm in the other. In the latter section were two completed cells
and an incomplete loaf of pollen. In the former section was a pollen loaf with egg,
but no pith partition.
The bee larva in the first cell constructed by the female bee had been completely
consumed by 11 parasitic larvae. Judging by the amount of bee feces in the cell
and the absence of pollen, the bee larva was evidently fully grown before death.
Later 59 9,224 4 of Leucopis klugii Westwood successfully emerged. The wasps
have been deposited in the British Museum and their identity kindly confirmed
by Z. Boucek. This is the first host record for L. klugii. Host bees were previously
known for 3 of the 9 species in the Neotropical hopei group to which L. klugii
belongs (Bouéek, 1974, Bull. Brit. Mus. (Nat. Hist.), Suppl. 23: 44). These
include species of Megachile (Megachilidae) and Xylocopa (Apidae).—Howetu
V. Daty, Department of Entomological Sciences, University of California, Berkeley,
California 94720.
Plant Associations Among Adult Hippomelas
(Coleoptera: Buprestidae)
E. Gorton LINSLEY
Division of Entomology and Parasitology
University of California, Berkeley, California 94720
AND
Epwarp 8S. Ross
Department of Entomology
California Academy of Sciences, San Francisco, California 94118
Although the genus Hippomelas contains some of the largest and
most striking buprestids in North America, relatively little is known of
their biology. Yet at times the adults are very abundant and may be
seen by the hundreds flying about shrubby plants in the desert or semi-
arid southwestern United States and northern Mexico. However, they
are usually most active in the warmest parts of the day and their readi-
ness to take flight and agility in moving around the stems of larger
plants makes feeding, mating and other activities difficult to observe
except in the late afternoon or early evening.
The adults are known to be flower visitors and because many species
have a yellow “pollinose” character to the integument it was long be-
lieved that this was simply pollen inadvertently brushed up from flowers.
However, the powdery pattern, although variable, is too consistent, the
beetles too large and the distribution over the surface too even to be
inadvertent. Furthermore, some of the species which are most heavily
““pollinose” visit the same flowers as those that are among the least
heavily “‘pollinose.” Vogt (1949), in commenting upon the Buprestidae
of the Lower Rio Grande Valley of Texas, has not only helped to clarify
this situation in Hippomelas but also where pulverulent character ap-
pears elsewhere in the Buprestidae by pointing out that it is a secretion
(see below). His findings will probably be confirmed in future studies
of the family.
The majority of the Buprestidae are believed to be host-specific in the
larval stage, at least at the generic or family level (see, for example,
Barr, 1971; Chamberlin, 1936; Knull, 1925; Vogt, 1949). Further-
more, among those species which visit flowers in the adult stage, there
is a marked tendency among some of them to visit blossoms of the plant
Tue Pan-Paciric ENntomowocist 52: 272-285. Ocroser 1976
Voi. 52, No. 4, OcToBER 1976 273
species which serve as hosts for the larvae especially in such genera as
Acmaeodera, although this is by no means a general rule (cf. Beer, 1940,
1944; Linsley and Ross, 1940). As a result, it might be reasonable to
suppose that the same correlation applies to some of the species of Hip-
pomelas. However, the evidence on this point is mixed. In the hope of
stimulating further studies on the subject we are presenting a summary
of the published information known to us, together with a few original
observations and adult plant associations derived from data on speci-
mens in the collection of the California Academy of Sciences, San Fran-
cisco (CAS), the California Insect Survey, Essig Museum of Entomology,
University of California, Berkeley (CIS) and the Department of Biology,
Arizona State University, Tempe (ASU).
Appreciation is expressed to W. F. Barr and Gayle H. Nelson for read-
ing the manuscript and providing helpful suggestions and to the latter
for kindly identifying much of our material. D. S. Verity provided
some useful records from his collection and Mont A. Cazier arranged
to have specimens from Arizona State University made available. Juan-
ita M. (Mrs. E. G.) Linsley ably assisted in making field observations
and collections.
HipPOMELAS (GYASCUTUS) CAROLINENSIS (Horn)
The original description of this species is correctly cited by Cham-
berlin as Horn (1883:288). The type locality given by Horn was Wil-
mington, North Carolina and was undoubtedly based upon erroneously
labeled material. It was first recorded from the West (Grand Canyon,
Arizona) by Townsend (1895).
The records at hand for plant associations for H. (G.) carolinensis
are as follows: from the collection of Arizona State University (all in
the San Simon Valley, Cochise Co., Arizona or the adjacent Chiricahua
Mountains, det. Nelson): two miles northeast of Portal, June 18, 1961
(M. A. Cazier) and 8 miles NE of Portal in Turkey Creek Wash, June
10, 1960 (M. A. Cazier) on Acacia greggii and 2.5 miles northeast of
Portal, September 6, 1960 (M. A. Cazier) and 5 miles northeast of Por-
tal, July 13, 1965 (J. H. Davidson, J. M. Davidson and M. A. Cazier)
on Larrea tridentata; from the California Academy of Sciences (det.
Helfer), Cathedral City, Riverside Co., California, July 16, 1950, four
on Larrea [tridentata], one on Acacia greggii (J. W. MacSwain), same
locality, July 13, 1951, three on Acacia greggii (J. W. MacSwain and
Ray F. Smith), Whitewater, Riverside Co., California, July 9, 1950, five
on Larrea tridentata (T. R. Haig, P. D. Hurd, and H. M. Graham), Palm
Springs, Riverside Co., California, July 16, 1950, five on Larrea [triden-
274 THe Pan-Paciric ENTOMOLOGIST
tata] (P. D. Hurd), Palms to Pines Highway, Riverside Co., Califor-
nia, elev. 1000 ft. May 28, 1940, one on Larrea tridentata (W. L.
Swisher), Banner, [San Diego Co.], California June 25, 1953, cut from
Croton root (Helfer collection), and Borego, [San Diego Co.], Califor-
nia, May 29, 1950, reared from Encelia root (Algert).
Records supplied by D. S. Verity include one example 3 miles west
of Beaumont, Riverside Co., California, July 9, 1961 on Eriogonum fas-
ciculatum, four at 2000 ft. elev., along highway 74, Santa Rosa Mins.,
Riverside Co., California, June 17, 1961, on Tamarix, one same area
3000 ft. elevation, May 21, 1962, one 5 mi. south of Palmdale, Los An-
geles Co., California, July 6, 1957, five 2 mi. southwest of Phelan, San
Bernardino Co., California, June 14, 1959, and five specimens with the
same data, July 16, 1960. These last four collections were all taken
from Juniperus californicus.
HIpPpOMELAS (GYASCUTUS) CASTANEUS HELFER
Described originally as a subspecies of H. insularis, H. (G.) castaneus
is treated as a full species by Barr (1970). The type series consisted of
three specimens from Angel de la Guardia Island, Gulf of California,
taken on Larrea mexicana |=tridentata], June 30, 1921, by E. P. Van
Duzee.
HIPPOMELAS (GYASCUTUS) DIANAE HELFER
Helfer (1954) described this species from a long series of adult speci-
mens from Palm Springs and Whitewater, Riverside Co., California
mostly in July, stating that they were consistently taken on Ephedra.
Most of the types and other specimens studied by him are housed in
collections of the Essig Museum of Entomology (CIS) and the Cali-
fornia Academy of Sciences (CAS). Confirming a preference for
Ephedra, we have examined material in the collection of Arizona State
University as follows: 9 miles west of Winterhaven, Imperial Co., Cali-
fornia, June 3, 1968 (R. Nevelyn) on E. trifurca 18 miles southeast of
Parker, Yuma Co., Arizona, May 14 and 18, 1966 (J. H. Davidson, J.
M. Davidson, and M. A. Cazier) on E. trifurca, and 67 miles north of
San Felipe, Baja California, Mexico, June 6, 1968 (N. Leppla, J. Bige-
low, M. A. Cazier and J. Davidson) on E. trifurca. In addition, the ASU
collection contains two specimens each from Algodones Dunes, Imperial
Co., California on Larrea tridentata, June 3, 1970 (M. A. Cazier, O.
Francke and L. Welch) and 16 miles west of Winterhaven, Imperial
Co., California, July 12, 1966 (J. M. Davidson and M. A. Cazier) on
Eriogonum deserticola. D. S. Verity (in litt.) collected seven specimens
Vou. 52, No. 4, OcToBER 1976 215
of this species on Acacia at Palm Springs, Riverside Co., California,
June 22 and 23, 1957 and seven more 2 miles south of Palm Desert,
Riverside Co., June 30, 1957.
HIPPOMELAS (GYASCUTUS) COMPACTA CASEY
A specimen in the collection of the California Academy of Sciences
identified as this species by E. C. Van Dyke, was collected on mesquite
|Prosopis| at San Pedro Bay, Gulf of California, July 7, 1921 by E. P.
Van Duzee.
HIPPOMELAS (GYASCUTUS) INSULARIS HELFER
Although Helfer (1953) does not mention the fact, some of the speci-
mens in the type series (CAS) from San Marcos Island [anglicized to
“Marcus Island” by Helfer], Gulf of California, June 19, 1921, collected
by E. P. Van Duzee, bear the label “ex Larria (sic) mexicana” |=Larrea
tridentata].
HIPPOMELAS (GYASCUTUS) FULGIDA BARR
Barr (1969, 1971) has reported finding larvae and a dead adult of H.
(G.) fulgida Barr in the roots of shadscale, Atriplex confertifolia in El-
more Co., Idaho. However, adults were also found on foliage of willow
(Salix sp.), sweet clover (Melilotus alba) and Russian thistle (Salsola
kali). These were regarded by him as visitation rather than true host
records (although he did not speculate on the reasons for the visitation,
it is possible that in one or more of the instances cited it may have been
for the purpose of adult feeding). Barr also added the observation that
many specimens had been taken from Japanese beetle traps, providing
the first record of the response of a buprestid beetle to this type of chemi-
cal attractant.
HIPPOMELAS (GYASCUTUS) JUNIPERINUS (WICKHAM)
Described from three specimens beaten from juniper in July on Chad-
burn’s Ranch in the foothills of the Pine Valley Mountains, about 22
miles from St. George, Utah, at an altitude of about 4500 feet. We are
not aware of further published plant associations nor have we seen speci-
mens bearing host data.
HIPPOMELAS (GYASCUTUS) OBLITERATA (LECONTE)
Horn (1866, 1868) reported that this species occurs rather abun-
dantly during the summer “on the low willows that are so plentiful in
the Owens Valley [California].” This observation was apparently veri-
276 THe Pan-Paciric ENTOMOLOGIST
fied by J. W. MacSwain who captured nearly 250 specimens on willow
(Salix sp.) at Antelope Springs, Inyo Co., California on July 17, 1953
(CIS, Helfer det.). However, Nelson (in litt.) who examined the Mac-
Swain specimens and also collected a long series of specimens at this
locality himself does not regard them as obliterata, the type of which he
has seen, but as close to or part of the H. fulgida-H. lariversi Barr com-
plex.
Fall (1901) and Van Dyke (1942) have recorded the occurrence of
the beetles on flowers of mesquite (Prosopis juliflora) at Banning,
California, in July. Chamberlin (1926) lists this plant as the host for
the species and Van Dyke (1942) reiterates that it is normally found
about mesquite (Prosopis). Wickham (1905), without being specific
stated that it was found on several desert shrubs. Kunze (1904) ob-
served adults on little-leaf palo-verde (Cercidium microphyllum) (listed
as Parkinsonia) in Arizona. He remarked that this buprestid, “heavily
dusted with yellow powder, and the ground or body color being of a
ereenish-gray, is found on young “Palo Verde,” the stems of which are
bright green. Under a glaring hot sun in May or June this beetle is
very alert, and a silken net is soon torn to pieces by the long thorns of
this shrubby tree. The bark and thorns of very young “Palo Verde”
much resemble obliteratus.”
All of these records may require revision when the taxonomic status
of H. (G.) obliterata is clarified.
HIPPOMELAS (GYASCUTUS) PACIFICA CHAMBERLIN
Adults of this species were captured on Atriplex polycarpa in late June
and early July at a locality 5 miles south of Kettleman City, Kern Co.,
California (Nelson, 1962). The species had been reported previously
as having been collected on “sagebrush” (Chamberlin, 1938). A “co-
type” bearing this label is in the collection of the California Academy
of Sciences.
HrppoMELas (GyascuTus) PLANICOSTA (LECONTE)
The first plant association record for this species known to us is that
of Townsend (1893) who in a report on some insects of New Mexico
stated “This large buprestid was found July 8 on mesquite bushes
(Prosopis juliflora) and later on flowers of the same. On July 17 great
numbers ake seen on flowers of Larrea mexicana |=L. tridentata] or
Creosote Bush. When found on the flowers they are covered with pol-
len, sivinel them a rich yellow color.’’ In 1895, he reiterated that the
species was common at Las Cruces, New Mexico on Larrea and mes-
VoL. 52, No. 4, OcToBER 1976 pare
quite. Wickham (1905) stated that it frequents especially the bushes
of Larrea [tridentata]. Smyth (1934) also confirmed the association
with Larrea, stating that they usually carried pollen upon the thorax
and forepart of the elytra and commented that when alarmed they some-
times flew straight up into the air to a height of a hundred feet or more,
then took off with the wind. Hurd and Linsley (1975) reported that it
was the most abundant species of Hippomelas encountered by them at
flowers of Larrea in southeastern Arizona and New Mexico during the
summer-fall blooming period (after the arrival of the summer rains).
They also reported that in southern California it has been taken in June
and July on Larrea when the plants are not in bloom and thus they are
believed to be the larval host. They added that during this off-season,
J. W. MacSwain took long series from Atriplex at Blythe, Riverside
County, California, although these may ultimately prove to be a differ-
ent species.
Plant associations listed on specimens at hand are as follows:
Larrea tridentata: New Mexico (Granite Gap, Peloncillo Mts., Hidalgo Co.,
July 2, 1965, J. H. and J. M. Davidson and M. A. Cazier) (ASU); Arizona (Dia-
mond Creek and Colorado River, Mojave Co., June 18, 1972, M. Kolner (ASU) ;
9 miles SE of Bell Butte, Tempe, Maricopa Co., June 20, 1972, M. Kolner (ASU) ;
6 miles S of Parker, Yuma Co., July 22, 1967, J. H. and J. M. Davidson and M. A.
Cazier; vicinity of Portal, Cochise Co., various dates in June, July, August and
September, M. A. Cazier, E. G. and J. M. Linsley) (CIS) ; Nevada (Overton, August
22, 1930, E. W. Davis) (CAS); California (Midway Well, Death Valley, Inyo Co.,
June 11, 1970, M. A. Cazier, O. Francke, L. Welch (ASU); 5.2 miles S of Sara-
toga Springs, Inyo Co., June 12, 1970, M. A. Cazier, O. Francke, L. Welch (ASU) ;
Indian Wells, Riverside Co., June 6, 1970, M. A. Cazier and O. Francke (ASU) ;
Oasis Station, Riverside Co., June 20, 1956, M. Wasbauer (CIS); Saltdale, River-
side Co., June 8, 1940, K. S. Hagen (CIS); 8 miles NW Palm Springs, Riverside
Co., July 28, 1956, M. Wasbauer (CIS); Whitewater, Riverside Co., July 9, 1950,
P. D. Hurd, Jr. (CIS) ; 2 miles S of Bard, Imperial Co., September 15, 1951, W. L.
Swisher) (CIS); Baja California, Mexico (67 mi N of San Felipe, June 6, 1968,
N. Leppla, J. Bigelow, M. A. Cazier, J. Davidson) (ASU).
Prosopis juliflora: Arizona (9 miles SE Bell Butte, Tempe, Maricopa Co., June
21, 1972, Donna and Martin Kolner) (ASU); California (Winterhaven, Imperial
Co., July 12, 1966, J. M. Davidson and M. A. Cazier) (ASU).
Atriplex (various species): New Mexico (White Sands, National Monument,
Otero Co., June 29, 1947, C. P. Strand (CAS); Arizona (2 miles NE Portal, Co-
chise Co., August 5, 1965, J. Hand, J. M. Davidson and M. A. Cazier) (ASU);
California (Blythe, Riverside Co., July 4, 1951, J. W. MacSwain and R. F. Smith)
(CIS) ; Palo Verde, Riverside Co., July 11, 1966, J. M. Davidson and M. A. Cazier)
(ASU).
Tamarix pentandra: Utah (Lime Creek, 8 miles N Mexican Hat, San Juan Co.,
June 29, 1974, L. Draper, O. Francke, M. A. Cazier) (ASU); Arizona (1 mile SW
Marble Canyon, Coconino Co., July 9, 1967, J. H. and J. M. Davidson and M. A.
278 Tue Pan-Paciric ENTOMOLOGIST
Table 1. Plant associations.
Plants
Cupressaceae
Juniperus californicus
Juniperus sp.
Ephederaceae
Ephedra sp.
Ephedra trifurca
Salicaceae
Salix spp.
Polygoniaceae
Eriogonum deserticola
Eriogonum fasciculatum
Chenopodiaceae
Atriplex sp.
Atriplex confertifolia (larval host)
Atriplex polycarpa
Salsola kali
Leguminosae
Acacia greggii
Acacia constricta
Acacia vernicosa
Acacia sp.
Mimosa sp.
Mimosa dyscocarpa
Prosopis sp.
Prosopis juliflora
Prosopis pubescens
Cercidium microphyllum
Cercidium floridum
Melilotus albus
Dalea spinosa
Beetles
Hippomelas carolinensis
H. juniperinus
H. dianae
H. dianae
H. imperialis
. cuneata
. fulgida
. obliterata
mez
H. dianae
H. imperialis
H. carolinensis
. planicosta
. fulgida
. pacifica
fulgida
by my be
carolinensis
. planicauda
caelata
caelata
. planicauda
dianae
. planicauda
. planicauda
compacta
cuneata
. obliterata (?)
caelata
cuneata
. planicosta
sphenica
cuneata
. obliterata (?)
caelata
fulgida
. planicosta
py BB BR
Vou. 52, No. 4, OcTroBER 1976 279
Table 1. (Cont.)
Plants Beetles
Zygophyllaceae
Larrea tridentata H. carolinensis
H. castanea
H. cuneata
H. dianae
H. insularis
H. planicosta
H. sphenica
Euphorbiaceae
Croton sp. (larval host) H. carolinensis
Tamaricaceae
Tamarix pentandra H. planicosta
Tamarix sp. Hf. carolinensis
Compositae
Encelia sp. (larval host) H. carolinensis
Cazier (ASU); Cliff Dwellers Lodge, Coconino Co., July 8, 10, 1967, J. H. and
J. M. Davidson and M. A. Cazier) (ASU).
Dalea spinosa: California (16 miles S of Vidal, Riverside Co., July 14, 1966, J.
M. Davidson and M. A. Cazier) (ASU).
In terms of numbers, 16 collections from Larrea are represented by 62 indi-
viduals, 3 collections from Atriplex by 58 individuals, two collections from Prosopis
by. 4 individuals, two collections from Tamarix by 5 individuals and the collection
from Dalea by a single specimen.
HipPpoMELAS (HIPPOMELAS) PLANICAUDA CASEY
Although we have not found plant associations for this species in
the literature, of 23 specimens before as bearing plant records 18 are
from Mimosa, 3 from Prosopis and two from Acacia. The collection
data as follows: Tucson, Pima Co., Arizona, “on cat claw [Acacia greg-
gii|, October 5, 1935 (O. Bryant) (CAS); Santa Rita Experiment Sta-
tion, Pima Co., Arizona, elev. 4400 ft., Mimosa dyscocarpa, September
5, 1970, Martin Kolner (ASU); Madera Canyon, Santa Rita Mts., Ari-
zona, elev. 4500 ft., Mimosa sp., September 3, 1964, W. Turner (ASU) ;
Santa Rita Mts., Arizona, Sept. 4, 1934, Mimosa, D. K. Duncan (ASU) ;
and one mile E of Portal, Cochise Co., Ariz., Acacia vernicosa, August
18, 1957, J. A. Chemsak and B. J. Rannells (CIS).
Nelson (in litt.) confirms that in his experience this species has been
consistently taken on Acacia. The adults superficially resemble those of
Hf. (H.) sphenica.
280 THE Pan-PactFic ENTOMOLOGIST
HipPpoMELAS (HIPPOMELAS) SPHENICA (LECONTE)
(Figure 2)
The first record of a plant association for Hippomelas sphenica is
apparently that of Griffith (1900), who, in discussing the Coleopterous
fauna of the Salt River Valley, Arizona, reports finding the species in
November, stating that “so closely does it resemble the branches of the
mesquite that it requires close searching to find them.” However, the
most informative statement about the habits of the beetles is that of Vogt
(1949) who observed them in the Lower Rio Grande Valley of Texas.
He reported as follows: “very common on mesquite, especially on dec-
adent and fresh cut trees where brush is being cleared. Females were
often seen ovipositing in dead mesquite snags. These beetles seem to
feed freely on mesquite foliage as evidenced by their copious green-
colored fecula.’ Fresh emerged specimens lack the characteristic yellow
pulverulence which seems to be a secretion accumulating with age. Evi-
dently pulverulence, as it generally occurs among the Buprestidae, is
of this nature; therefore, the variations in the definition of the pulveru-
lent markings in this family. It is pointed out, however, that the pubes-
cence associated with such markings is apparently unaffected by age
but rather by abrasion.” Hurd and Lindsley (1975) while confirming
that the species is ordinarily associated with mesquite in the San Simon
Valley of Arizona and New Mexico report that it also feeds occasionally
at flowers of Larrea tridentata.
Garnett (1918) records the capture of a wasp (probably Cerceris)
near Barstow, California, carrying a small specimen of H. sphenica,
although the identification of the beetle is doubtful (Verity, in litt.).
Although we have collected this species from mesquite (Prosopis
juliflora) on a number of occasions, it was not until an hour before
sunset on September 2, 1975, at the San Xavier Mission, near Tucson,
Pima Co., Arizona that we encountered them in such large numbers that
it was possible to make a few observations on adult behavior. The sky
was broken to the west providing intermittent sun and the temperature
ranged from 22 degrees C to 20 degrees C. The beetles were flying to
low growing living mesquite plants, mostly less than 8 ft. tall in a low
area adjacent to cultivated fields, mostly planted to cotton. The beetles
ranged from one or two per plant to 12 or 15. Both sexes were repre-
sented and were mostly resting on stems or feeding on foliage, confirm-
ing the conclusion of Vogt based upon the color of their feces. Nearly
all of the plants exhibited extensive feeding damage. During the brief
periods when the rays of the sun were unobscured the beetles were more
VoL. 52, No. 4, OcToBER 1976 281
Fic. 1. Female of Hippomelas caelata on stem of living Acacia vernicosa, nr.
Fairbank, Arizona, July 1965 (E. S. Ross).
Fic. 2. Mating pair of Hippomelas sphenica on stem of living Prosopis juli-
flora, at San Xavier Mission nr. Tucson, Arizona, after sunset, September 3, 1975
(E. S. Ross).
282 THE Pan-Paciric ENTOMOLOGIST
active in moving over the stems and leaves, flying about the plants and
were more easily disturbed. However, when they were more quiescent
they were difficult to see unless silhouetted against the sky because of
their cryptic coloration and the fact that they would move to the op-
posite side of the branch when approached. Although most individuals
observed were solitary, some mating was taking place (Fig. 2). Males
would crawl over individuals of either sex testing receptivity by probing
with the aedeagus.
Specimens available with plant association data are as follows: 2.5
miles NE Portal, Cochise Co., Arizona, Prosopis juliflora, July 29, 1959,
E. G. Linsley (CIS); 2 miles NE Portal, Arizona, Prosopis juliflora,
August 18, 1965, J. H. and J. M. Davidson and M. A. Cazier (ASU); 1
mile E Portal, Arizona, Acacia vernicosa, August 18, 1957, J. A. Chem-
sak and B. J. Rannells (CIS) ; Portal, Arizona, Prosopis juliflora, August
15, 1958, E. G. Linsley (CIS); San Xavier Mission, Pima Co., Arizona,
Prosopis juliflora, September 2, 1975, E. G. and J. M. Linsley and E. S.
Ross (CA5, C15) ; Madera Canyon, Santa Rita Mts., Pima Co., Arizona,
mesquite [Prosopis], September 2, 1950, W. L. Swisher (CAS); and 9
miles NNW Santa Rita Experiment Station, elevation 4150, Prosopis
juliflora, August 23, 1971, Martin Kolner (ASU).
HIPPOMELAS (STICTOCERA) CAELATA (LECONTE)
(Figure 1)
Chamberlin (1926) records the host of this species as Palo Verde
(Cercidium floridum) and Van Dyke (1942), apparently following
Chamberlin, states that “it breeds in a number of desert trees like the
Palo Verde (Cercidium floridum).’ However, this association is cer-
tainly doubtful and we regard it as regularly attached to Acacia. Cazier
(1951) reported the species rather common on Acacia at a number of
localities in north central Mexico and we have consistently found it on
Acacia constricta and A. vernicosa in southeastern Arizona. In addi-
tion to our own collections, representative data from material before
us is as follows: Granite Pass, Peloncillo Mts., Hidalgo Co., New Mex-
ico, Acacia constricta, August 6, 1963 and August 27, 1969, A. Raske
(ASU); same locality and host plant, August 31, 1965, J. H. and J. M.
Davidson and M. A. Cazier (ASU) ; same locality, Acacia greggii, July
21, 1965, A. Raske (ASU); Lake Cienega, Hidalgo Co., New Mexico,
Prosopis juliflora, July 24, 1964, J. M. Pickle, M. Mortensen and M. A.
Cazier (ASU); 3 miles E of Portal, Cochise Co., Arizona, Acacia con-
stricta, August 16, 1965, J. H. Davidson and M. A. Cazier (ASU);
Tombstone, Cochise Co., Arizona, Acacia vernicosa, August 17, 1957,
VoL. 52, No. 4, October 1976 283
J. A. Chemsak and B. J. Rannells (CIS) ; same locality, Acacia, August
13, 1940, W. L. Swisher (CAS); and one mile south of Santa Ana,
Sonora, Mexico, on Acacia constricta, September 14, 1953, N. A. Lewis
(CAS).
HIPPOMELAS (PRASINALIA) CUNEATA (Horn)
The first published plant association for this species known to us is
that of Smyth (1934) who reported collecting a series of forty examples
in an hour’s time from bull’s horn Acacia (probably A. cornigera) near
Salina Cruz, Isthmus of Tehuantepec, Mexico in August. Subsequently,
Nelson (1959) recorded finding adults abundant on Prosopis chilensis
[juliflora] and Acacia greggii at Morong Valley, San Bernardino
County, California in July. Material at hand yields the following addi-
tional information: Thermal, Riverside Co., California, on dead Proso-
pis, June 18, 1940, W. F. Barr (CAS); Borrego Valley, California,
Prosopis, June 6, 1946, Donald Davis (CAS); 2 miles S. of Bard, Im-
perial County, California, on mesquite | Prosopis], September 22, 1951,
W. L. Swisher (CAS); 0.9 miles SE of Bell Butte, Tempe, Maricopa
Co., Arizona, Prosopis juliflora, July 21, 1972, Donna and Martin Kol-
ner(ASU) ; 3 miles S of Parker, Yuma Co., Arizona, 3 examples on dead
Prosopis juliflora, July 9, 1966, J. M. Davidson and M. A. Cazier
(ASU); same locality, date and collectors, Prosopis pubescens; same
locality, one example on dead Prosopis juliflora, July 14, 1966, David-
son and Cazier (ASU); and 6 miles south of Parker, Arizona, Larrea
tridentata, July 21, 1967, J. H. and J. M. Davidson and M. A. Cazier
(ASU). D. S. Verity (in litt.) reports collecting 25 examples of this
species on Salix at Lost Hills, Kern Co., California, July 2, 1956, four
at Barstow, San Bernardino Co., California, July 7, 1963, and six at
Victorville, San Bernardino Co., July 13, 1963.
Nelson (in litt.) states that the most abundant he has ever seen this
species was in Kern Co., 4 miles E of Lost Hills, on June 25, 1961 where
he collected 80 specimens from one Prosopis juliflora tree that was ap-
proximately a quarter of a mile from any other such trees. These speci-
mens were Collected within 45 minutes.
HIPPOMELAS (PRASINALIA) IMPERIALIS BARR
When describing this species, Barr (1969) recorded two occasions on
which adults were collected from plants of Eriogonum deserticola on the
eastern slopes of the Imperial Valley, California. Subsequently, Walters
(1975) has found it on this same plant 5 miles west of Glamis, Imperial
County, California in mid-June. Verity (in litt.) collected an example
284, Tue Pan-PacitFic ENTOMOLOGIST
on Ephedra, 4 mi. west of Gordons Wells, Riverside Co., California on
July 8, 1972.
LITERATURE CITED
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EDITORIAL NOTICE
With this issue of the Pan-Pacific Entomologist, the editorship
changes hands. We are certain that the members of The Pacific Coast
Entomological Society and other suscribers to the journal join us in
thanking the retiring editor, John Doyen and his staff, consisting of
John Chemsak, Assistant Editor, and Rollin Coville, Editorial Assistant.
For three years John voluntarily served us in a truly professional man-
ner, producing our publication while employing his own high standards
of quality and scientific expertise. We are sincerely grateful to John
and his staff for their efforts on our behalf.—Editors.
286 THE Pan-Paciric ENTOMOLOGIST
A new species of Alloxysta hyperparasitic on aphids associated
with South American Nothofagus forests
(Hymenoptera: Cynipidae)
FreD G. ANDREWS
California Department of Food and Agriculture, Sacramento, 95814
The following description is done as part of an effort to catalog
arthropods associated with species of Nothofagus (Fagaceae). The
material was collected by E. I. Schlinger in 1966 and 1967 while survey-
ing for insects on Nothofagus in the southernmost areas of Chile and
Argentina. The aphids Neuwquenaphis edwardsi Laing and N. schlingeri
Ris-Lambers were collected by beating, and held until mummies were
noticed. The mummies were individually isolated. Several hundred
aphids were parasitized by a species of Pseudephedrus (Hymenoptera:
Aphidiidae) ; six of these aphids were hyperparasitized by a single,
previously undescribed Alloxysta species.
The presence of a primary and secondary parasite complex on New-
quenaphis on Nothofagus was previously unknown. This was in part due
to lack of collecting and in part due to the low density and high disper-
sion of the hosts. No other hyperparasites, either cynipid or chalcidoid,
were reared, although a single specimen of an undescribed species in
the charipine cynipid genus Phaenoglyphis was collected on Nothofagus
foliage.
The high dispersion of the aphid host suggests that this new Alloxysta
has great host-finding capabilities.
This species of Alloxysta is immediately distinguishable from all other
New World Alloxystinae by its bicolorous state. The equal length of the
third, fourth and fifth antennal segments in both male and female and
the lack of curvature in any of the male segments is also diagnostic.
Alloxysta nothofagi, n. sp.
Adult female——Head above antennal insertion pale reddish-brown; frons, genae,
mouthparts and antennae straw yellow. Pleural area including pronotum straw
yellow. Mesonotum and abdomen shiny castaneous brown. Entire leg and wing
veins pale lemon yellow.
Head as wide as high; frons sparsely setose; genae, occiput and vertex glabrous.
Antennae 13-segmented, filamentous, cylindrical; segments 1-3 smooth, 4-13 longi-
tudinally ridged; segments 3-5 in ratio of 19:19:19 (19=0.1 mm) (Fig. 2);
segments 6-13 slightly wider than 3-5. Pronotum behind occiput densely clothed
with transparent decumbent setae. Mesoscutum moderately convex, longer than
wide by ratio of 17:15, glabrous. Scutellum with long decumbent transparent setae
Tue Pan-Pactric ENtomouocist 52: 286-287. Octoser 1976
Vout. 52, No. 4, Ocroser 1976 287
Fics. 1-4. Alloxysta nothofagi Andrews. 1. Antennal segments 3-5, male. 2.
Antennal segments 3-5, female. 3. Wing, female. 4. Wing, male.
e
on lateral posterior margins, glabrous above. Wings exceeding body as 88:57 (88 =
1.6 mm). Radial cell, elongate, 1.6 times longer than wide (Fig. 4). Ratio of
radial cell length and width to wing width 2.6 and 7.0, respectively; r-1 straight;
r-2 evenly arcuate.
Adult male.—As in female, except antennae 14-segmented, with 1-3 smooth,
4-14 longitudinally ridged, segments 3-5 without bow and in ratio of 16:16:15
(Fig. 1); radial cell 2.3 times longer than wide, ratio of radial cell length and
width to wing width 2.5 and 6.1, respectively (Fig. 3).
Types.—Holotype male, 3.7 mi. S. Puerto Moreno, Rio Negro Prov., ARGEN-
TINA, XI-17-1966, E. I. Schlinger (E66-11-17a) (Nothofagus antarctica/Neuquen-
aphis edwardsi/Pseudephedrus sp.). To be deposited in University of Chile, Santi-
ago, Chile.
Paratypes——2 males, same data as holotype. 1 female (E67-2-10g), 2 males
(E67-2-10b), 18 km W. Angol, Malleco Prov. CHILE II-10-1967, E. I. Schlinger
(Nothofagus obliqua/Neuquenaphis schlingeri/Pseudephedrus sp.). Three para-
types to be deposited in USNM and two to remain in author’s collection.
288 THE Pan-Paciric ENTOMOLOGIST
Oviposition of Cyrtopogon inversus'
(Diptera: Asilidae)
D. STEVE DENNIS
Environmental Sciences Division, Stearns-Roger, Inc.,
Box 5888, Denver, Colorado 80217
AND
RosBert J. LAVIGNE
Plant Sciences Division, Univ. of Wyoming
There is a paucity of information concerning the oviposition behavior
of asilids belonging to the genus Cyrtopogon Loew. Bromley (1946)
observed that, like other members of the subfamily Dasypogoninae,
Cyrtopogon females possess spines on the ovipositor which are used to
brush soil over the eggs following oviposition.
Of the 98 described species of Cyrtopogon (Hull, 1962), detailed
observations have been made for one species in Sweden, Cyrtopogon
lateralis Fallen and a species in Russia, Grypoctonus (Cyrtopogon)
daimyo Speiser.
Cyrtopogon lateralis females deposit elongate whitish yellow eggs in
the soil (Melin, 1923). The presence of large numbers of spines on the
Ovipositor was presumed to be associated with its habit of ovipositing
in hard soil mixed with clay. Melin noted that C. lateralis used these
spines to brush soil over the eggs and to hide the oviposition hole. Lehr
(1966) made the following observations for Grypoctonus (Cyrtopogon)
daimyo (translated from the Russian) :
“Eggs are deposited in loose soil along pathways, under a grassy slope, or near
fruit trees in gardens. During oviposition, the female bends the abdomen and buries
it in the ground, often up to the very wings. Judging from the depth at which eggs
are found, one may assume that the abdomen extends slightly during oviposition.
There is one rosy-yellow egg laid in each place. During oviposition the female
crawls from place to place or flies over short distances.”
Lavigne (unpublished data) observed a Cyrtopogon sp. (probably
glarealis Melander) ovipositing at the base of a grass clump among
wood chips in soil and duff near Grassy Lake in Teton National Forest,
Wyoming. Oviposition occurred at 3:10 PM in the shade where the soil
temperature was 31°C.
The following observations were made of Cyrtopogon inversus on Elk
1 Published with the approval of the Director, Wyoming Agricultural Experiment Station, as Journal
paper no. JA-833. This research was supported in part by National Science Foundation Research
Grant GB-29617X.
Tue Pan-Paciric ENTOMOLOGIST 52: 288-290. OctosrerR 1976
Vou. 52, No. 4, OctoBER 1976 289
Fics. 1-2. Fig. 1. Coniferous forest clearing where Cyrtopogon inversus ovi-
positions were observed. Fig. 2. Elongate creamy white eggs of Cyrtopogon
inversus (52X).
Mountain near Elk Mountain, Wyoming. The area was located in a
coniferous forest clearing at an elevation of 2,920 m above sea level
(Fig. 1). Several complete ovipositions were observed between 12:00
noon and 2:30 PM between July 18th and 21st, 1972. All eggs were
deposited in the soil or in the soil litter.
Prior to oviposition Cyrtopogon inversus females usually “‘searched”’
for suitable sites to deposit their eggs. Females flew 45 to 60 cm above
the ground in a zigzag path and made two to three 60 to 72 cm circles
about a site before landing to oviposit. Once females landed they
crawled 2.5 to 5 cm into the shade of vegetation, small pieces of wood,
or rocks. As the females crawled along the ground, some were observed
to thrust their ovipositor from side to side as we have seen other asilids
do in their search for suitable oviposition sites.
Once in the shade the flies ordinarily pushed their ovipositors directly
into the substrate. However, two females dug their ovipositors into the
soil with rapid lateral sweeping movements. It is assumed that most fe-
males did not exhibit such behavior because of the ease with which they
could insert their ovipositors into the loose litter.
While ovipositing, females rested their legs on the substrate and kept
their wings folded over their body. The arched abdomen was sunk into
the substrate up to the posterior margin of the second abdominal seg-
ment.
The average time for a complete oviposition was 2 minutes, with a
range between 1 and 3 minutes. Once eggs were deposited, females
either withdrew their ovipositor with rapid lateral sweeping movements,
which covered the eggs and oviposition hole with soil or litter, or flew
off without exhibiting such behavior.
At the time of oviposition, substrate surface temperatures ranged
290 THE Pan-Paciric ENTOMOLOGIST
between 27 and 39°C. The temperature 3 to 4 mm under the surface,
at the approximate depth at which the eggs were laid, varied between
32.5 and 37°C.
The eggs of Cyrtopogon inversus are creamy white (Fig. 2). Two to
four eggs were laid per clutch and apparently “cemented” together.
Individual females were observed to oviposit up to three times before
being lost to sight.
The elongate eggs of this species ranged in length from 1.12 to 1.42
mm, with a mean of 1.31 mm. The average width was 0.48 mm, with a
range which varied between 0.42 and 0.56 mm. Eggs examined with a
scanning electron microscope, up to 2,000, showed no surface sculptur-
ing.
LITERATURE CITED
BRoMLEY, S. W. 1946. Guide to the insects of Connecticut. Part VI. The Diptera
or true flies of Connecticut. Third Fascicle. Asilidae. Conn. Geol. Nat.
Hist. Surv. Bull. 69: 1-51.
Hutt, F. M. 1962. Robber flies of the world. The genera of the family Asilidae.
Bull. U.S. Nat. Mus. 224(1 & 2): 1-907.
Leur, P. A. 1966. Biology and taxonomy of robber flies (Diptera, Asilidae) of
the genera Cyrtopogon Loew and Grypoctonus Speiser of Kazakhstan
and Central Asia. Collection of Scientific Articles by Graduate Students
and Researchers, Kazakh State Univ., Alam-Ata. p. 95-101. (In Rus-
sian.)
Meuin, C. 1923. Contributions to the knowledge of the biology, metamorphosis,
and distribution of the Swedish asilids in relation to the whole family
of asilids. Zoologiska Bidrag Fran Uppsala 8: 1-317.
RECENT LITERATURE
ToxicoLocy oF INsecTicipes. Fumio Matsumura. Plenum Publishing Co., New
York, N. Y. 1975. 10011. 504 pp., 89 illustrations, 85 tables. $27.50.
Brees AND BEEKEEPING. R. A. Morse. Cornell University Press, Ithaca, New York.
1975. 320 pp., 65 illustrations, $13.00.
BEES, THEIR VISION, CHEMICAL SENSES, AND LANGUAGE. Revised Edition. K. von
Frisch. Cornell University Press, Ithaca, New York. 1975. 176 pp., $8.75,
cloth; $3.45, paper.
Vou. 52, No. 4, OcToBER 1976 291
Studies on the Biology of Heliconius charitonius L.
in Costa Rica
(Nymphalidae: Heliconiinae)
ALLEN M. Younc
Division of Invertebrate Zoology, Milwaukee Public Museum,
Milwaukee, Wisconsin 53233
The heliconiine butterfly Heliconius charitonius L. (Nymphalidae),
is one of the most geographically widespread members of this subfamily
(Heliconiinae) throughout tropical America (e.g., Comstock and Brown,
1950; Brown and Comstock, 1952). The adult is strikingly different
in appearance from many other sympatric heliconiines by the distinct
black and yellow color pattern of the wings. Adults of this species are
known to roost communally (Jones, 1930; Poulton, 1931; Young and
Thomason, 1975), a sophisticated behavioral trait (Gilbert, 1975) it
shares with other Heliconius. Although the early stages of various Heli-
conius have been studied in Trinidad (Beebe, Crane, and Fleming, 1960;
Alexander, 196la, b) and Brazil (e.g. Brown and Mielke, 1972), little
is known about the biology of H. charitonius in Central America. The
apparent coevolutionary associations of Heliconius with certain plant
sroups in the tropics (Benson et al., 1975; Gilbert, 1975) warrants the
study of selected species. This paper summarizes studies on the adult
and juvenile biology of H. charitonius in Costa Rica.
Stupy AREA AND METHODS
Heliconius charitonius was studied near La Virgen de Sarapiqui (He-
redia Province) in northeastern Costa Rica; adults and early stages were
observed or collected from a strip of young secondary forest (5 years
old) bordering a dirt road that separates Finca La Tigre (property of
Compania Agricola Myristica, S.A.) and Finca de Oscar Arias. This
region of Costa Rica is a transitional zone between premontane and
lowland tropical wet forest; it is about 200 meters in elevation with a
short, erratic dry period during March and April. Although the area
includes primary wet forest habitat, much of it has been converted to
various successional stages of secondary forest and cultivated land.
Both field and laboratory studies on H. charitonius were carried out
from January 11 to February 15, 1976. During this period, field ob-
servations were conducted as follows: January 11-12 (2 days), Janu-
ary 15-17 (3 days), January 26—-30 (5 days), February 4-7 (4 days),
and February 11-15 (5 days). Field studies consisted of (1) searches
Tue Pan-Paciric Entomo.ocist 52: 291-303. Octroser 1976
292 THE Pan-Paciric ENTOMOLOGIST
for early stages, (2) observations on oviposition behavior, (3) observa-
tions on the mortality of early stages, (4) recording of adult feeding
sites, and (5) recording of other heliconiines active in the same habitat.
Laboratory studies dealt with the rearing of individuals from the egg
stage primarily for the description of the life cycle, and the estimation
of developmental time. Individuals were reared by confining eggs and
fresh clippings of the larval host plant into a clear plastic bag kept tightly
shut. Field observations were made at several different times during
the day.
RESULTS
Habitat, Oviposition, and Larval Host Plant.—Heliconius charitonius
occurs primarily in young secondary forest tropical communities, where
adults visit various inflorescences throughout sunny hours of the day.
The primary flowers visited include several Compositae, one or more
species of Curcubitaceae and Cephaelis tomentosa (Rubiaceae). The
Compositae visited have either white or yellow inflorescences while
the other plant families visited have deep orange or red inflorescences
It is very likely that adults feed on pollen (Gilbert, 1975),
although “pollen loads” on probosci are less frequently encountered in
this species relative to other microsympatric species such as H. hecale,
which bears notoriously large pollen loads. Gilbert (1972), however,
reports heavy pollen loads for H. charitonius. Adults generally visit
flowers throughout the day on clear sunny days.
Typical flower visitations by H. charitonius relative to other Heli-
conius species, occurred on the sunny morning of February 13, 1976.
In a forest edge clearing where one curcubit species and Cephaelis
tomentosa were in bloom, with their patches separated by only four
meters of bare ground, H. charitonius visited both. There were seven
flowers of C. tomentosa with full inflorescences (although 35 in total were
present, most of these had dropped the yellow corolla tubes), and eleven
flowers of the curcubit. One “fresh” adult visited a curcubit flower at
9:05 A.M. and a “worn” adult soon followed suit at 9:15 A.M. This
same individual then flew across and visited a Cephaelis flower by 9:18
A.M. Since no butterflies were marked, “traplining” or other move-
ment patterns could not be demonstrated. Two other species active in
this area visited both flowers: at least two different adults of H. hecale
(“fresh”; “worn’) together gave a total of four visits to curcubit flow-
ers and three visits to Cephaelis, while at least two adults of H. cydno
(““fresh”; “worn”) were scored twice on Cephaelis and four times on the
curcubit. All of these visits occurred between 8:40 and 9:55 A.M. A
similar pattern of visits for the same three species was seen the following
VoL. 52, No. 4, OcToBER 1976 293
Fic. 1. Egg positioning and the egg stage of Heliconius charitonius. Upper left:
three eggs are shown in a folded terminal bud of leaflets of T. lobata. Upper left:
three eggs are shown; note one egg adjacent to a young extrafloral nectary. Lower
left: top view of one egg nestled down in folded leaflets; lower right: the egg.
morning. Both H. hecale and H. cydno are more frequent visitors at
these particular flowers than H. charitonius.
Adults are concentrated in large numbers on a daily basis in patches
of young secondary forest where the primary larval host plant, Tetra-
stylis lobata (Passifloraceae) is found in close proximity to adult feed-
ing sites. Tetrastylis lobata is the primary plant used for oviposition by
294, THE Pan-PaciFIc ENTOMOLOGIST
H. charitonius in this region of Costa Rica, despite the fact that it often
occurs together with other passifloraceous vines such as P. vitifolia and
P. edulis. Adult vines of 7. lobata are very large and consist of large,
bulky patches growing over other vegetation. It generally flowers dur-
ing January and February, and H. charitonius lays eggs on young and
adult vines.
Although eggs are laid singly, a female may lay several eggs on the
yet folded terminal leaf buds of T. lobata (Fig. 1). One female laid
five eggs on a single terminal leaf bud within a 20-minute period one
morning. I observed a total of four ovipositions; the first occurred on
January 11, 1976 (3:00 P.M.). The female in this instance was very
worn and tattered, and she took about 20 seconds to lay one egg. During
the oviposition, the wings were fluttering at a steep angle to the body.
Several other individuals of H. charitonius were flying in the immediate
vicinity of this female, but this did not stop the oviposition. On another
day (January 16), a female (fresh) laid three eggs in quick succession
on a single terminal unit of folded leaf buds. I have also observed that
different females very likely lay their eggs on the same unfolded leaf
units or that the same female will return to lay eggs on the same leaf
unit on different days. Individual females, presumably searching for
Oviposition sites, spend a great deal of time selecting such sites. Some-
times the egg is nestled far down into the whorl of folded leaves, making
only the apical aspect visible (Fig. 1). In addition to eggs being at-
tached to folded leaf buds, they may also be found on the stem near
the leaf bud (Fig. 1). Oviposition on tendrils has not been observed.
On large vines, eggs are often laid near the top, where most of the folded
leaf buds are found; on young vines, eggs are often laid within a few
centimeters of the ground, and amidst tall grasses that cover them. As
with feeding, I have observed oviposition throughout the day in good
weather. Although several other heliconiines lay eggs on Passiflora in
the area, none of these lays eggs on 7. lobata, and H. charitonius does
not lay eggs on Passiflora. I searched a total of eight vines of P. vitifolia
and did not find any eggs or caterpillars of H. charitonius; I did, how-
ever, find caterpillars of erato, cydno and hecale. It is noteworthy that
at least six other heliconiines occur in the same habitat: Agraulis va-
nillae, Philaethria dido, Heliconius cydno, H. hecale, H. erato, and H.
sara. All heliconiine eggs found on 7’. lobata here and reared (a total
of 15 eggs collected) proved to be H. charitonius. Caterpillars of H.
charitonius feed on both young and old leaves of T. lobata.
The following observations on the early stages were made from indi-
viduals obtained from a set of eight eggs discovered in a single unit of
VoL. 52, No. 4, OcToBEer 1976 295
Fic, 2. Caterpillars of Heliconius charitonius. Upper left: first instar just above
a young extrafloral nectary; upper right: third instar. Lower left: fourth instar;
lower right: fifth instar.
folded leaf buds; of the eight eggs, four were reared successfully to the
adult stage in the laboratory.
“Disappearances” of Eggs.—Although several eggs were collected for
laboratory rearing studies, I left a total of seven additional eggs on vines
in the field to see if they hatched successfully. These eggs were on three
296 THE Pan-PaciFIc ENTOMOLOGIST
Fic. 3. The pupa of Heliconius charitonius. Upper two and lower left photo-
graphs: lateral, ventral, and dorsal views respectively. Lower right: a pupa in the
field being attacked and eaten by Crematogaster ants. Several individuals of a
small ant are shown on and along the abdominal region of the pupa; a single
individual of a larger ant (Ectatomma) is present on the pupa along the frontal
area of the wing case (barely in focus).
VoL. 52, No. 4, OcToBER 1976 297
different vines of T. lobata. They all appeared about the same age from
their color. Within two days, all were gone, and no first instar larvae
were found. I had searched a total of 12 different vines for Heliconius
eggs, but found only three with eggs in the same area of habitat.
Early Stages—tThe large, yellow egg (Fig. 1) is 1.4 mm high and
0.9 mm at the base; it is elongated and truncated at the top and bottom.
It lasts five days as measured on four eggs kept in the laboratory.
The first instar larva (Fig. 2) is 3-4 mm at hatching; it is light,
translucent orange with a glossy orange-yellow head capsule. It lasts
two days (as measured on four individuals).
The second instar bears the full complement of head and body scoli,
with the ground color of the body remaining light orange. The scoli are
black and branched. The head scolus is about 1.2 times the head height.
The dorsal prothoracic plate or crest is thin. This instar lasts four days
and attains a length of 7 mm.
The third, fourth, and fifth instars are virtually identical in appear-
ance (Fig. 2). The basic color of the body is bluish-white and the head
capsule is a glossy pale lemon-yellow. All scoli are black. The head
capsule possesses two sets of black blotches. All legs are dull orange;
the anal clasper is dull orange with a black spot dorsally. The dark spots
on the body segments (Fig. 2) are very dark reddish-brown. The head
scolus is about one-half the height of the head. The third instar lasts
four days and reaches a length of 13 mm; the fourth instar lasts four
days and reaches 30 mm, and the fifth instar lasts three days and reaches
41 mm. A pinkish prepupa (30 mm long) is active prior to pupation.
The highly sculptured pupa (Fig. 3) is about 30 mm long with the
basic color being brownish-orange. It is moderately patterned with dark
lines and reflective silver spots. The silver spots are paired and situated
dorsally in the thoracic region and first abdominal segment. Fine streaks
of white occur on the wing cases. The head bears broad, well developed
spatulate appendages that are 5 mm long. Dorso-lateral flanges occur
on the third and fourth abdominal segments (smaller on the fourth) and
bearing long, outward and downward-directed spines; shorter spines
occur on the fifth, sixth, and seventh segments. Short spines with tuber-
cles as bases occur on the metathorax and also on the first and second
abdominal segments. Very short spines occur along the antenna cases
(at forewing costal sutures) ; there are very slight tubercles on the wing
cases in future cells Ms-Cu, and Cu,;-Cus. As measured on four indi-
viduals, the pupa lasts 10 days for both sexes. From four pupae two
males and two females were obtained; three of these adults are shown
in Fig. 4.
298 THE Pan-PactFIC ENTOMOLOGIST
Fic. 4. Adult Heliconius charitonius. Upper: an adult female just after eclosion
(part of pupa case visible to the right). Lower left: three adults reared from the
egg stage in the laboratory; lower right: a young adult feeding on a curcubitaceous
flower (February 13, 1976, near La Virgen).
Pupae as Prey.—I searched the vicinity of one vine of 7. lobata on
January 12 and discovered four pupae that later matched the description
of the pupa of H. charitonius obtained in the rearing studies. All of these
were affixed to plants very close to the vine. Two of the pupae were being
attacked by ants (Fig. 3). One pupa was virtually eaten down to the
VoL. 52, No. 4, OcToBEerR 1976 299
last abdominal segment at this time, and the other pupa (located about
one meter from the first one) was just in the initial stages of being eaten
by two different types of ants. I observed this pupa for two subsequent
days and eventually all that remained was the last two abdominal seg-
ments and cremaster. When first discovered the pupa was being eaten
near the head appendages by an ant, Crematogaster sp. (Fig. 3), but
two days later, when about half the pupa was gone, three individuals of
a much larger ant (Ectatomma sp.) were also present with the other
ants (Fig. 3). Subsequently I discovered three more eaten pupae in
the same area, giving a total of five pupae found as ant prey.
In the laboratory, three of the original group of eight individuals
being reared died as pupae as prey for cockroaches. These pupae were
kept in a small screen cage with pupae of other butterflies (Ithomiidae,
Brassolidae), but they were the first ones to be attacked by cockroaches
during the night.
DISCUSSION
In terms of phylogenetic considerations (Emsley, 1965), H. chari-
tonius is one of the advanced species of the genus, and the morphology
of its early stages illustrate close relatedness with other species such as
Hf. hecale and H. cydno, which occur in the same habitat in northeastern
Costa Rica. A good characteristic to separate later instar caterpillars of
H. charitonius from those of H. cydno or H. hecale is the relatively
shorter head scolus in the former species. Also, the pupa is more highly
sculptured in Hf. charitonius than in these other species. Recently, Brown
and Benson (1975) have argued for the operation of larval mimicry in
microsympatric species of Heliconius. Even though the caterpillars of
differently closely related species of Heliconius might be feeding on dif-
ferent host plants in the same habitat, it is held that larval mimicry can
act to reduce predation rates by vertebrates. Thus selection would favor
the occurrence of species with similar caterpillars in the same habitats.
Given divergence in host plants, to reduce competition for larval food,
several closely related advanced species might co-occur to reduce preda-
tion on caterpillars. Thus it is not unexpected to find H. charitonius
occurring with H. hecale, cydno, erato, and sara. At least one of these
species (H. hecale) feeds on Passiflora vitifolia here, and it indicates
divergence in oviposition and larval feeding from H. charitonius. In
this region of Costa Rica, there are likely no more than seven species of
Passifloraceae that could be exploited by Heliconius, so that each species
may have a different primary larval host plant (Gilbert, 1975).
The multiple oviposition of several eggs on the same vine (either by
300 THE Pan-PaciFIc ENTOMOLOGIST
the same or different female) suggests that (1) the larval host plant is
viewed by the species as an abundant resource, and (2) this species is
not food-limited in the caterpillar stage. Strict resource limitation would
be normally expected to provide selection pressure favoring a greater
scattering of the eggs. Gilbert (1975), however, points out that vines of
Passiflora generally occur at low densities. All Heliconius butterflies
are associated with Passifloraceae. Further field observations are needed
to see how egg densities of Heliconius correlate with vine size or bio-
mass. For example, H. charitonius may lay one or few eggs on smaller
vines of 7. lobata than on larger ones. Benson et al. (1975), report T.
lobata as the host plant of H. charitonius in southeastern Costa Rica
and Panama; my studies indicate that it is also the host plant in north-
eastern Costa Rica. Benson et al. (1975) also report that H. charitonius
either scatters several eggs on the meristem of 7’. lobata, or else lay a
single egg on the meristem. Despite the occurrence of other Passi-
floraceae in the habitat (and in the same vine patch), only 7. lobata is
locally used as the larval host plant; this points to specialization by H.
charitonius, something not unexpected (Gilbert, 1975). Lamas (1974)
makes some interesting comments regarding larval host plants of the
subspecies peruvianus of H. charitonius (Comstock and Brown, 1950;
Brown and Comstock, 1952): he maintains, that in addition to several
species of Passiflora, this subspecies, presumed near extinction in Lima,
Peru, may exploit other plants as larval host plants.
T. lobata is very likely the host plant for this butterfly in other life
zones (Holdridge, 1967) in Costa Rica since Standley (1937) mentions
that it also occurs in montane tropical wet forest regions. It is possible
that H. charitonius uses other Passifloraceae as larval host plants in
Costa Rica, but for montane rain forest and premontane wet forests on
the eastern slopes of the central Cordillera, the primary host plant is 7.
lobata. However, other passifloraceous host plants of H. charitonius are
known from Costa Rica (Benson e¢ al., 1975). Although T. lobata pos-
sesses large tendrils, it is interesting that H. charitonius does not lay
eges on them. Heliconius hecale and H. cydno are both known to lay
eggs singly on the tendrils of passifloraceous vines (Young, 1973; 1975).
There is divergence in the oviposition habits among advanced heli-
coniines. Brown (1973) reports that H. nattereri in Brazil oviposits on
Tetrastylis ovalis. Apparently this primitive passifloraceous genus is
not exploited as a larval host plant by many heliconians.
Larval host plant specialization (i.e., one primary host plant in a
given region), expected for most species of Heliconius (Gilbert, 1975),
couple with considerable mortality of eggs and caterpillars from preda-
VoL. 52, No. 4, OcroBEeR 1976 301
tion (Gilbert, 1975), probably keep the adult populations of individual
species at or near the carrying capacity of the environment. Predation
by ants on Heliconius pupae warrants further study in terms of its im-
pact on populations. The well developed and apparently functional extra-
floral nectaries of TJ. lobata, P. vitifolia and perhaps other
microsympatric Passifloraceae used as larval foods may serve to at-
tract pupa predators such as ants. Gilbert (1975) points out that these
glands attract predatory insects such as ants and wasps; Ehrlich and
Gilbert (1973) found that eggs of H. ethilla are eaten in large numbers
by ants in Trinidad. Thus the observed disappearances of eggs seen
here might be attributed to predation by ants. The fact that only a few
out of several vines had Heliconius eggs might be indicative of intense
egg predation. It is known that the female pupae of H. charitonius re-
lease a pheromone which attracts males (Edwards, 1881; Emsley, 1965) ;
it would be interesting to know if such a pheromone attracts potential
predators such as ants as well.
As adult longevity of H. charitonius is high (Cook, Thomason, and
Young, in prep.) and vagility generally low (Young and Thomason,
1975; Young and Carolan, 1976), there is further stress placed on
Heliconius for local species packing, promoting K-selection as the adap-
tive strategy. It is generally known that Heliconius butterflies are dis-
tasteful to vertebrate predators, since presumably cyagenic glycosides
and alkaloids from Passifloraceae are retained as “plant poisons” in the
insects (Gilbert, 1975). Certainly the vivid yellow and black wing color
pattern of H. charitonius suggests unpalatable properties. Although
K-selection might be the general adaptive strategy for these butterflies,
in regions of Central America such as northeastern lowland Costa Rica,
where a large spectrum of secondary habitats are available at the present
time, there is opportunity for species such as H. charitonius to be flex-
ible in the adaptive context.
The above considerations suggest that H. charitonius and other mem-
bers of the genus are not food-limited as caterpillars, owing to a variety
of factors primarily (1) availability of a large host plant biomass, (2)
specialization by different species on different passifloraceous species in
the same region, (3) diversity of oviposition habits by different species
(Gilbert, 1975), and (4) high levels of mortality of juvenile stages.
The large number of Heliconius species and other heliconiines that usu-
ally exist in the same region (Gilbert, 1975 mentions 10) warrants fur-
ther field study directed towards elucidating the mechanisms of co-
occurrence. At least for H. charitonius, from my preliminary studies,
(1) host plant specialization, (2) high selectivity of flower-type visita-
302 THE Pan-PaciFic ENTOMOLOGIST
tion, (3) apparent high mortality of early stages (yet to be confirmed).
(4) well developed adult population cohesiveness, and (5) micro-
sympatry with other heliconiines, with perhaps the existence of larva!
mimicry, suggests K-selection in secondary tropical habitats. Although
very preliminary and limited, the flower visitation observations sug-
gest repeated visits to the same flower patches by H. charitonius and
other species. Perhaps these butterflies are “traplining” for nectar and
pollen. Gilbert (1975) mentions that advanced species such as /7. chari-
tonius are expected to trapline daily for nectar and pollen sources. and
that they also trapline for egg laying sites. These properties contribute
to adult population cohesiveness.
ACKNOWLEDGMENTS
This research is a by-product of National Science Foundation Grant
GB-33060. I am grateful to Lenore Durkee (Grinnell College) and Luis
Poveda (Museo Nacional de Costa Rica) for assistance with the identi-
fication of plant species. The full cooperation of Dr. J. Robert Hunter
of Compania Agricola Myristica, S.A. (CAMSA) with logistic details
is greatly appreciated. Dr. Paul Kannowski identified the ants.
LITERATURE CITED
ALEXANDER, A. J. 196la. A study of the biology and behavior of the caterpillars
and emerging butterflies of the subfamily Heliconiinae in Trinidad.
West Indies. Part I. Some aspects of larval behavior. Zoologica 46:1—24.
ALEXANDER, A. J. 1961b. A study of the biology and behavior of the caterpillars
and emerging butterflies of the subfamily Heliconiinae in Trinidad.
West Indies. Part IT. Molting, and the behavior of pupae and emerging
adults. Zoologica 46:105—124.
Breese, W., Crane, J. AND FLeminc, H. 1960. A comparison of eggs, larvae and
pupae in fourteen species of heliconiine butterflies from Trinidad, Wesi
Indies. Zoologica 45:111-154.
Benson, W. W., Brown, K. S., Jr., anp Gitpert, L. E. 1975. Coevolution of plants
and herbivores: passion flower butterflies. Evolution 29:659-680.
Brown, F. M. anp Comstock, W. P. 1952. Some biometrics of Heliconius
charitonius (Linnaeus) (Lepidoptera, Nymphalidae). Amer. Mus. Novit.
1574:1-52.,
Brown, K. S., Jr. 1973. A portfolio of neotropical Lepidopterology. Organizacao
Tecnica de Taquigrafia e Gravacoes Editora, Rio de Janeiro, 28 pp.
Brown, K. S., Jr., AND MieLkE, O. H. H. 1972. The heliconians of Brazil ( Lepi-
doptera: Nymphalidae). Part II. Introduction and general comments.
with a supplementary revision of the tribe. Zoologica 57:1—40,
Brown, K. S., Jr. anp Benson, W. W. 1975. The heliconians of Brazil (Lepidop-
tera: Nymphalidae). Part VI. Aspects of the biology and ecology of
Heliconius demeter, with description of four new subspecies. Bull.
Allyn Museum 26:1-19,
Vou. 52, No. 4, OcrosBErR 1976 303
Comstock, W. P., anp Brown, F. M. 1950. Geographical variation and sub-
speciation in Heliconius charitonius (Lep.: Nymphalidae). Amer. Mus.
Novit. 1467:1-21.
Epwarps, W. H. 1881. On certain habits of Heliconia charitonia Linn., a species
of butterfly found in Florida. Papilio 1:209-215.
ExR.IcH, P. R. ann Giipert, L. E. 1973. Population structure and dynamics of
the tropical butterfly Heliconius ethilla. Biotropica 5:69-82.
Emstey, M. G. 1965. Speciation in Heliconius (Lep., Nymphalidae) : morphol-
ogy and geographic distribution. Zoologica 50:191-254.
GitserT, L. E. 1972. Pollen feeding and reproductive biology of Heliconius but-
terflies. Proc. Nat. Acad. Sci. USA 69:1403-1407.
GILBERT, L. E. 1975. Ecological consequences of a coevolved mutualism between
butterflies and plants. pp. 210-240, In Coevolution of Animals and
Plants, L. E. Gilbert ed., Univ. Texas Press. 246 pp.
Hovopripce, L. R. 1967. Life zone ecology. Tropical Science Center, San Jose,
Costa Rica. 35 pp.
Jones, F. M. 1930. The sleeping Heliconias of Florida. Nat. Hist. 30:635-644.
Lamas, G. 1974. Supuesta extincion de una mariposa en Lima, Peru (Lepi-
doptera, Rhopalocera). Revista Peruana de Entomol. 17:119-120.
Poutton, E. B. 1931. The gregarious sleeping habits of Heliconius charitonius
L. Proc. Roy. Entomol. Soc. London 6:4—10.
STANDLEY, W. 1937. The flora of Costa Rica. 210 pp.
Younc, A. M. 1973. Notes on the biology of the butterfly, Heliconius cydno
(Lepidoptera: Heliconiinae) in Costa Rica. Wasmann J. Biol. 31:
337-350.
Younc, A. M. 1975. Observations on the life cycle of Heliconius hecale zuleike
(Hewitson) in Costa Rica. Pan-Pacific Entomol. 51:76—85.
Younc, A. M. ann TuHomason, J. H. 1975. Notes on communal roosting of
Heliconius charitonius (Nymphalidae: Heliconiinae) in Costa Rica. J.
Lepid. Soc. 29:243-255.
Younc, A. M. ann Carotan, M. E. 1976. Daily instability of communal roost-
ing in the butterfly Heliconius charitonius in Costa Rica. J. Kansas
Entomol. Soc.: In press.
304 THE Pan-PaciFic ENTOMOLOGIST
Notes on the Biology of the Introduced Elaterid
Conoderus exsul (Sharp)
(Coleoptera: Elateridae)
M. W. STONE
131 Sir Damas Dr., Riverside, California 92507
The Sugarcane wireworm Conoderus exsul (Sharp), originally from
New Zealand and later Hawaii, was first discovered in 1937 at Alameda,
California; currently it is recorded from 14 counties in California. (Stone,
1975). In the Hawaiian Islands the larvae have been found feeding on
sugarcane shoots and in Alameda Co., Calif., on grass roots. Larvae of
this species made rapid growth when reared in salve cans on wheat.
This species could become an important pest of vegetable and field crops
with an increase in numbers.
Light trap collections in 1974-75 indicate that C. exsul adults have
become increasingly abundant in Riverside and Orange counties. This
information on seasonal abundance together with preliminary data on
the biology of this insect are presented.
CONODERUS EXSUL (SHARP)
Monocrepidius exsul Sharp 1877: 20.
Monocrepidius exsul; Brown 1880. Manual New Zealand Coleoptera I: 294. (copy
of Sharp’s original description).
Monocrepidius exsul; Sharp 1908. Fauna Hawaiiensis, vol. III, part V: 369 (records
it from Hawaii).
Monocrepidius exsul; Williams, 1931: 168. Reports larvae feeding on sugarcane
shoots in Hawaii.
Conoderus bicarinatus Van Dyke, 1932: 297. Type male, U.S.A. Arizona, Mt.
Washington, near Nogales, 6000’, July 8, 1919. (J. A. Kusche) No. 3107, CAS.
Monocrepidius exsul; Graves, 1938: 91. Larvae and adults collected at Alameda,
Cal. Det. by Van Dyke.
Conoderus arizonicus Van Dyke, 1939: 11. New name for C. bicarinatus Van Dyke
(nec Reitter, 1891).
Conoderus duplicatus Van Dyke, 1943: 44. Unnecessary new name for C. bicari-
natus Van Dyke (nec Reitter, 1891).
Conoderus exsul; Lane, 1954: 246. Report on distribution in Calif.
Conoderus exsul; Stone, 1975: 165. Widespread in central and southern Calif.
Conoderus bicarinatus Van Dyke appears to be a synonym of exsul.
Becker, E. C. (personal comm.) examined the type of exsul and it com-
pared favorably with California specimens. Adults collected throughout
California, Arizona, and also in Hawaii have been found to be similar
to the holotype of bicarinatus Van Dyke (No. 3107 CAS). The Van
Tue Pan-Pactric ENtomo.wocist 52: 304-310. OctToser 1976
Vout. 52, No. 4, OcToBER 1976 305
Dyke description of this species appears adequate. Additional notes and
measurements on both sexes follow.
MALE.—Length 8.5 to 12 mm. width 2.6 to 3.2 mm. Antenna slender and thread-
like extending 14 to 1 segment beyond apex of hind angles of pronotum. Segments
2 to 5 with following lengths in mm.; .17, .25, .46, .46. Male genitalia (Fig. 6)
FEMALE.— (Figs. 4 & 5). Length 11 to 13 mm.; width 3.8 to 4mm. More robust
and larger than male. Antenna shorter extending to apex of hind angle of pro-
notum.
BIOLOGICAL AND MorpHo.ocicaL Notes
Egg.—The egg is whitish, smooth, oval shaped and measures 0.43 by 0.53 mm.
Newly deposited eggs have a sticky coating which cause soil particles to adhere,
making the eggs difficult to locate in the soil. Adults obtained at black light and
confined indoors in 2 oz. soil-filled, salve cans laid eggs from June to early Septem-
ber. A total of 218 eggs were obtained from one large reared female.
Larva.—(Fig. 1). The newly hatched larvae are whitish, but become cream
colored after the second moult. When mature the head, thorax and terminal lobes
of the ninth abdominal segment are dark reddish. The caudal notch on the ninth
segment is nearly closed on immature larvae but gradually enlarges and is V shaped
on mature larvae (Fig. 2). Full grown larvae measure 15 to 19 mm. long by 2.2
to 2.8 mm. wide.
Pupa.— (Fig. 3). When first formed the pupa is opaque white, but later becomes
cream colored and the eyes become conspicuous as dark spots. Mouth parts, an-
tenna, wing pads and legs are all visible. A sharp spine is present near each ante-
rior margin of the pronotum and a similar spine is also present at each posterior
angle. Two stout shorter spines are located at the tip of the abdomen. Pupae vary
greatly in size: 8 to 14 mm. long by 2.5 to 3.4 mm. wide.
The duration of the larval period of Conoderus exsul was determined
by confining newly hatched larvae individually in 2 oz. salve tins con-
taining moist 30 mesh soil and wheat, which were replenished at 2 week
intervals. The larvae were confined indoors at temperatures of 15—25° C
(60 to 78° F). Of a group of 34 larvae which hatched during August
1974, 5 pupated in March 1975, 11 in April, 10 in May and 8 in June—
July 1975. The wide range in the duration of the larval period, from
8 to 12 months, is difficult to explain since the larvae, depending on
size, were fed equal amounts—2 to 5 kernels of wheat at each feeding.
The first adult was obtained on March 20, 1975 or 2.7 months before
adults were collected at light outdoors. The pupal period for 29 indi-
viduals ranged from 14 to 18 days, an average of 15.7 days. Adult
longevity varied depending upon temperature, moisture and food
availability. When confined in glass jars indoors with moistened cotton
and slices of carrot adult life ranges from 25 to 65 days, averaging 38
days.
In 1975 rearings of C. exsul were conducted to determine if the dura-
THE Pan-Pactric ENTOMOLOGIST
306
Vou. 52, No. 4, OcToBER 1976 307
¥
Fics. 5-6. Conoderus exsul Sharp. Fig. 5. Body outline A. Apices of elytra
with rounded notches; found only in C. scissus Schfr. B. Unusual in that the hind
angles of the prothorax have paired carina. Fig. 6. Male genitalia (Ventral view)
Bt Toe
tion of the larval period varied according to the time at which the egg
hatched. Larvae hatching on June 15 and from August 15 to September
15 were reared indoors in containers on wheat as previously described.
In the above two groups designated as early (June 15) and late (August
15 to September 15) hatch, 15.8% and 12% of the larvae matured the
same year. The larval period in these groups ranged from 54 to 103
days. The pupal period averaged 14 days. (Table 1). In the early
<
Fics. 1-4. Conoderus exsul Sharp. Fig. 1. Larva, dorsal and ventral view, x 6.
Fig. 2. Larva, ninth abdominal segment, dorsal view. Fig. 3. Pupa, ventral view,
< 8. Fig. 4. Adult, female, x 7.
308 THE Pan-PaciFic ENTOMOLOGIST
TABLE 1. Duration of larval stage of Conoderus exsul Sharp. River-
side, Cal. 1975-76.
Larvae completing development in
1975 1976
ise Pupa- Larval period Larval period
hatched Records _ tions Range Ave. Pupations Range Ave.
1975 (No.) (%) (Days) (Days) (%) (Days) (Days)
June 15 38 15.8 81-103 91 84.2 218-292 251
Aug. 15 to
Sept. 15 24 IZ 54-78 62 87.5 180-214 214
hatch overwintering group, pupations began January 22 and terminated
April 1, 1976. First and last adults were obtained on February 8 and
April 20.
In the late hatch overwintering larval group, the first pupation oc-
curred a month later on February 21 and the last April 23. The above
pupae transformed to adults March 12 and May Ill. There was not
much difference in the duration of the larval period in the early and late
hatch groups, as indicated by the averages, 251 and 214 days, respec-
tively. The same was true of the pupal period for both groups, which
ranged from 17 to 23 and averaged 19 days. (Table 2).
One would expect that the higher temperatures indoors would be
responsible for accelerating larval feeding and for their reaching ma-
turity abnormally early. Nevertheless, it is interesting to note that of 10
medium size larvae dug up outdoors in January and fed lima beans, all
pupated in February and March, as was the case with specimens in the
containers kept indoors.
Information obtained to date indicates that in the presence of ample
TABLE 2. Duration of the pupal period of Conoderus exsul. River-
side, Cal. 1975-76.
Larvae completing development in
1975 1976
Date cen 8 19, SA mente tt Fe Ta a ats Pee
hatched Range Ave. Range Ave.
1975 (Days) (Days) (Days) (Days)
June 15 12-16 14 18-23 19.4
Aug. 15 to
Sept. 15 14-15 14 17-20 18.6
VoL. 52, No. 4, OcToBER 1976 309
€ of total catch
June July Aug. Sept, Oct.
Fic. 7. Bimonthly catches of Conoderus exsul adults at black light. Riverside,
Calif. 1974-75.
food a small percentage of C. exsul larvae mature the same season,
whereas the majority complete their life cycle in the year following.
PHENOLOGY
At Riverside adults were collected nightly at a 15 Watt florescent
black light located adjacent to a bare field formerly planted to citrus.
J. Wilcox employed a similar trap set-up in an avocado grove located
4 miles east of Olive, CA. (Orange County), about 30 miles from River-
side.
In 1974 at Riverside, adults were first collected on June 14 and were
most numerous in the period July 31 to Aug. 31. Emergence terminated
on Oct. 17. Trapping at Olive was started later—on Aug. 2—and ter-
minated Oct. 22, yielding a total of 518 adults. At Riverside 286 adults
were collected over a much longer period. Peak emergence at Olive
ocurred in the period Aug. 31 to Sept. 15.
In 1975 at Riverside the first adult was collected at light on June 11.
The largest numbers of adults were obtained when outdoor temperatures
at 8 to 9 p.m. remained at 24 to 26° C (75 to 80° F), usually after day-
time temperatures reached maximums of 35 to 38° C (95 to 100° F).
No adults were collected when evening temperatures dropped to below 15°
C (60° F). Peak emergence in 1975 occurred in the period Aug. 1 to
31, the same period as in 1974. Adult activity ceased on Oct. 6. A total
of 211 adults were collected, with males and females in equal numbers.
Fig. 7 shows catches of adults bi-monthly at Riverside in 1974-75.
At Olive in 1975 (fig. 8) adults were first collected on June 26 and
terminated Nov. 2. The peak of collections occurred Sept. 1 to 15. A
total of 1238 adults were taken—an exceptional number for an intro-
310 THE Pan-Paciric ENTOMOLOGIST
30
nw
Oo
% of total catch
| a
June July Aug, Sept. Uct.
Fic. 8. Bimonthly catches of Conoderus exsul adults at black light. Olive,
Calif. 1974-75.
duced species, considering that the soil in this grove was of coarse
texture, not overly moist and generally unfavorable for wireworm sur-
vival. Of the total catch 48% were females.
ACKNOWLEDGMENTS
I am especially indebted to J. Wilcox, Olive, California, for making
nightly catches of adults during the two seasons of these studies and for
his valued suggestions in the preparation and reviewing of the manu-
script. My thanks to E. C. Becker, Biosystematics Research Institute,
Ottawa, for his interest and helpful suggestions. Thanks also to D. H.
Kavanaugh (Calif. Academy of Sciences) who provided the Type of
C. bicarinatus, and to Floyd G. Werner, Univ. of Arizona, who furnished
specimens and collection records of C. exsul in Arizona.
LITERATURE CITED
Graves, H. W. 1938. A Hawaiian Elaterid introduced into California. Pan-Pac.
Entomol., 14(2): 91.
Lane, M. C. 1954. Distribution of several introduced species of wireworms. U.S.
Plant Pest Control Branch, Coop. Insect Rept. 12: 246.
SHarp, D. 1877. Elateridae of New Zealand. Ann. Mag. Nat. Hist. 19(4): 20.
Stone, M. W. 1975. Distribution of 4 introduced Conoderus species in California.
(Elateridae). Coleopt. Bull. 29(3) : 163.
Van Dyke, E. C. 1932. Proc. Calif. Acad. Sci. (4th Ser.) 20(9): 297.
Van Dyke, E. C. 1939. A new name for bicarinatus V. D. Pan-Pac. Entomol. 15:
11.
Van Dyke, E. C. 1943. Conoderus bicarinatus VY. D. Pan-Pac. Entomol. 19: 44.
WitiiAMs, F. X. 1931. Handbook of the insects and other invertebrates of Ha-
waiian sugarcane fields. Hawaiian Sugar Planters Assoc. Honolulu, 168
pp.
VoL. 52, No. 4, OcTroBER 1976 oii
A New Species of Apatolestes from California
(Diptera: Tabanidae)
Wooprow W. MIpDLEKAUFF
Department of Entomology, University of California, Berkeley
AND
Rosert S. LANE
Vector and Waste Management Section, California State Department of Health,
Berkeley.
The pangoniine genus Apatolesies consists of a primitive group of
species confined to the western United States, northern Mexico, and
southwestern Canada. It is comprised of 10 previously described species
and | subspecies, 3 of which are found only in California. The addition
of rugosus n. sp. to this list brings the total number of species to 11, with
4. of them confined to the state.
The new species was discovered while the authors were studying the
tabanid fauna of California, and is described here to make the name
available.
This species is closely related to A. willistoni Brennan to which it will
run in existing keys due to the infuscation present in the costal cells at
the bifurcation of veins R4—R; and along the apical cross veins (Fig. 1).
A. rugosus can be distinguished from willistoni by its more robust body
and average larger size (12-13 mm); wrinkled (rugose) subcallus; ex-
tensive brown pollinose areas lateral to the ocelli; and hairless middle of
the frontoclypeus in the female (Fig. 2). A. willistoni is more slender
and averages smaller in body length (8-9 mm) ; the subcallus is smooth,
not rugose; the pollinosity next to the eyes on the vertex is gray and the
frontoclypeus is hairy. The male has the area of the small eye facets
blending into that of the large facets, whereas these areas are sharply
demarcated in willistoni.
Apatolestes rugosus, n. sp.
(Figs. 1-2)
Female holotype—Body length 12 mm. Ratio of basal width of frons to its
height and width at vertex is 1:2:1.5; vertex between ocelli and eyes with rusty-
brown pollinosity, the pollinose area acutely angled below median ocellus and
adjacent to eye; remainder of frons including basal callus shiny, jet black, with a
patch of long stramineous hairs above basal callus and adjacent to each eye; basal
callus markedly swollen, laterally attaining each eye; subcallus with gray to tan
pollinosity and a characteristic pair of wrinkles (Fig. 2); frontoclypeus sparsely
THe Pan-Paciric ENtomotocist 52: 311-313. Octoser 1976
312 THe Pan-PaciFiIc ENTOMOLOGIST
Fics. 1-2. Apatolestes rugosus, female. Fig. 1. Wing, showing venation and
areas of infuscation. Fig. 2. Frontal view of head showing rugose subcallus.
covered with long, straw-colored hairs; genae with numerous black hairs especially
near eye margins; apical palpal segment with a conspicuous dorsal groove, covered
with a few pale, but mostly black hairs; scape and pedicel with gray pollinosity;
the flagellum black.
Thorax and scutellum with gray pollinosity, the former with a submedian and
a broad black line laterally on each side and both with numerous, short, appressed,
pale golden-yellow hairs; pleurae with numerous, long, white and black intermixed
hairs. Femora and dorsal surface of tarsi black; tibiae and plantar surface of tarsi
dark brown.
Abdominal tergites black, each with a transverse band of gray pollinosity expand-
ing medially on tergites II and III to form small, indistinct triangles. Wing hya-
line except for infuscated costal cell, cross veins, and a spot at bifurcation (Fig. 1).
Male allotype——Body length, 10.5 mm. Except for sexual differences similar to
female holotype. The area of small eye facets brown to black blending impercepti-
bly into the brown area of large facets above; a few short black hairs at tip of
palpi.
VoL. 52, No. 4, OcToBer 1976 313
Holotype female, University of California Field Station, Hopland, Mendocino
Co., California, June 2, 1973, John R. Anderson, collector.
Allotype male, taken in copulo with the holotype female.
Paratypes.—47 2 2 and 20¢ 64, all topotypic, collected on various dates from
late May to late July, 1966, 1969, 1972, and 1973. In addition the following speci-
mens were seen but not placed in the paratype series: 19, Red Bluff, Tehama Co.,
VI-12-51 (California Department of Food and Agriculture) and 19, 20 mi. n.w.
Macdoel, Siskiyou Co., VII-14-67, L. L. Dunning, collector (U. C. Davis).
The holotype and allotype plus some additional paratypes will be deposited in the
entomological collection, California Academy of Sciences, San Francisco. Remain-
ing paratypes will be placed in the U.S. National Museum, Canadian National
Collection, Cornell University, and the California Insect Survey.
SCIENTIFIC NOTE
Liogorytes joergenseni (Bréthes), a cicada killer in Argentina (Spheci-
dae, Nyssoninae).—The only two wasp genera previously known to provision
cicadas are the well-known and widespread Sphecius, and the Australian Exeirus,
both genera in the sphecid tribe Gorytini. Now a third gorytin genus, Liogorytes,
is known to use cicadas.
A nesting colony of L. joergenseni (Bréthes) was discovered in a natural spring
zone at La Ciénaga, Catamarca, Argentina. The first sighting of this colony was
made 12 December, 1973, when Stange found a female dominated population (sex
ratio 30 females to 1 male) busily engaged in provisioning nests with a species
of Cicadidae (Chonosia sp.). This cicadid was present in tremendous numbers,
apparently having emerged from nymphs that had been feeding on the dominant
plant in the study zone, a species of Clump Grass (Sporolobus). Females were
observed carrying this medium size cicadid to its ground burrows which were about
one-half inch in diameter. Bohart and Stange revisited the site on 27 November,
1975, and found a male dominated population (38 males to 9 females). In contrast
to the first experience, only one adult cicadid was found and the population
activity was centered upon males searching for females. Based on the material
collected to date (78 specimens) the females are rather constant in size (about
22 mm long), whereas the males are highly variable (12 mm to 22 mm). When
the site was first visited in 1975, adults of LZ. joergenseni were quite abundant
(about 10 a.m.) but within an hour, after a marked increase in temperature, the
adults ceased flying. On a return to the site by Bohart on 16 December, 1975
no adults of either sex were seen and nesting was presumably complete.—R. M.
Bouart, Department of Entomology, University of California, Davis, 95616, and
LrioneL STANGE, Instituto Miguel Lillo, Tucuman, Argentina.
314 THe Pan-Paciric ENTOMOLOGIST
Bembicini of Baja California Sur: Notes on Nests, Prey, and
Distribution
(Hymenoptera: Sphecidae)
Howarp E. Evans
Department of Zoology and Entomology
Colorado State University
Fort Collins, 80523
During a trip by air and rental car to Baja California Sur in early
June 1975, my assistants and I collected 9 species of Bembicini and ob-
tained nesting data on two of these. We had hoped to study Bembix
magdalenae, described from Bahia Magdalena by C. L. Fox in 1926 and
not recovered since. Although we spent several days on the shores of
Magdalena and adjacent bays, we failed to find magdalenae but did
find nesting aggregations of the related and equally unstudied species
B. rugosa Parker. This proved to be a reasonably typical member of the
genus in nesting behavior despite its unusual structural features. Brief
notes are also presented on other Bembicini collected, as there has been
little systematic collecting of Sphecidae in Baja California Sur.
BEMBIX RUGOSA PARKER
This species was described from Arizona and to my knowledge has not
previously been taken in Baja California Sur. We found it to be not un-
common in two localities on Bahia Magdalena, on the Pacific side of the
peninsula. On June 9 we took several females on tall composites along
the roadside about 3 km east of San Carlos, and later the same day we
found several females nesting in a man-made excavation into an an-
chored sand dune, about 1 km from San Carlos but only 100 m from the
shores of the bay. The wasps nested in flat or slightly sloping soil
within the excavation, but not in the steep-sloping soil walls; we did
not see any on top of the dune or in adjacent, more active dunes. The
soil in the nesting site was a whitish sand of somewhat coarse texture,
with occasional streaks of more compact clay-sand. Empty, broken
Bembix cocoons of previous generations were abundant on the sand
surface and at depths to 20 cm. Evidently they were in the process of
being exposed by the winds from the bay; the active nest-cells we found
were deeper than 20 cm.
Several of the empty cocoons dug from the sand had fragments of
prey around them. These proved to be wings and body fragments of
muscoid and syrphid flies, as well as one bombyliid. Several old but
Tue Pan-Paciric Entomo.ocist 52: 314-320. Octosper 1976
VoL. 52, No. 4, OctosBer 1976 315
unbroken cocoons were measured and found to be 8-9 mm in maximum
diameter, 20-25 mm in length; the number of pores per cocoon varied
from 3 to 6 (x =4.7, N= 12). In one old cell containing an inviable
cocoon, at a depth of 20 cm, there was a female mutillid, Dasymutilla
gloriosa (Saussure). These mutillids were also seen walking over the
sand in some numbers. We have no firm evidence that this species is
a parasite of Bembix.
From 1000 to 1130 we watched one female B. rugosa make 110 trial
burrows, no more than 0.5 cm deep, before finally remaining at one
point and digging persistently for several hours. Each trial burrow
required only a few seconds of digging, and all were within an area of
about 3 square meters. When she finally persisted at one site, she
backed out periodically, spraying the sand a considerable distance and
producing a rather flat mound. She dug intermittently and had reached
a depth of only 25 cm by 1600 hours.
We marked several apparently active nests and excavated three of
these. One which was open most of the day was found to have a straight,
oblique burrow 105 cm long that reached a depth of 40 cm and then rose
several cm before terminating blindly. We assume this to have been an
incomplete nest that had been abandoned. A second nest was of similar
structure, the burrow passing obliquely downward for 35 cm, then
obliquely upward for 9 cm, then gradually downward for 3 cm before
reaching a cell 4 cm long at a depth of 22 cm. The cell was empty, so we
assume this nest had also been abandoned, or at least temporarily evac-
uated.
The third nest was of similar structure although with two lateral turns
near the bottom. The major part of the burrow was 70 cm long, forming
about a 45 degree angle with the surface that later steepened to about 65
degrees. The female was at the bottom of this burrow, at a depth of 35
cm. Beyond her the burrow passed obliquely upward for 6 cm, then
obliquely downward for 5 cm before reaching a horizontal cell measur-
ing 4 cm long by 1.5 cm high. The cell contained a single fly, Psilo-
cephala sp. nr. tergisa (Say) (Therevidae) lying on its back, with a
wasp egg 5 mm long attached erect to its side.
We also encountered B. rugosa at a locality about 60 km further south,
at a site about 2 km south of Puerto Chale, on Bahia de las Almejas,
which is an extension of Bahia Magdalena. An estimated 12 females and
a few males occupied the top of a low sandhill only 50 m from the man-
erove-fringed bay. The hill was covered with cacti and bushes; the
wasps nested in bare areas but were much attracted to one large bush
which was in bloom. Males flew about the flowers and open areas in
316 THE Pan-Paciric ENTOMOLOGIST
the morning hours of June 10, and several attempted matings were ob-
served. About 6 nests were marked in various stages of construction.
One female dug intermittently for the greater part of the day, then at
1640 made an initial closure of an apparently completed nest. The
mound at the nest entrance measured 25 cm long by 14 cm wide and 1.5
cm deep in the center. For 3 minutes the wasp passed over that half of
the mound closest to the entrance in irregular zigzag patterns, partially
leveling this half of the mound but leaving the other half intact. She
then made an elaborate closure, during which she made 18 radiating
lines, 9-14 cm long, emanating from the entrance. Between each line
she made a short flight, landing at or near the entrance and initiating
another line. The result was a small heap of sand covering the entrance
and flanked by radiating lines on both sides. This behavior also re-
quired about 3 minutes, after which the wasp dug through the entrance
and closed behind her at 1646 hours. Precisely the same behavior was
observed at the nest of a second individual that had been digging most
of the day.
We did not excavate either of these nests, but rather spent much time
digging out another nest which was being provisioned during the after-
noon. This nest was very deep and difficult to excavate because of the
dry, powdery condition of the sand and the fact that the burrow had two
sharp lateral curves. It began at about a 45 degree angle with the sur-
face, then steepened to about 60 degrees; the female was found in this
burrow about 1 m from the entrance. At 110 cm from the entrance the
burrow made a right angle turn, then went down another 10 cm, to a
vertical depth of 78 cm, before passing upward 3 cm and making
another right angle bend before passing obliquely downward 4 cm to a
cell. The cell measured 1.5 X 4 cm and contained 3 fresh flies and many
fly remains, as well as a nearly full-grown wasp larva. Two of the flies
were Asilidae (Ablautus flavipes Coquillett), one Mydidae (Pseudo-
nomoneura sp.).
We observed an apparent final closure of a fourth nest. This female
left a hole 4 cm long by 2 cm wide and 3 cm deep, to one side of which
there were about 20 radiating lines each measuring about 15 cm long.
We were unable to trace this burrow, which was tightly packed with
sand. We did attempt to excavate a fifth nest and were able to follow
the burrow for 1.3 m, to a vertical depth of 75 cm, but we failed to find
a cell.
BEMBIX SAYI CRESSON
This species has previously been reported from La Paz by C. L. Fox
(1923), and there is a pair from Cabo San Lucas in the California
VoL. 52, No. 4, OctoBer 1976 317
Academy of Sciences. Its nesting behavior has been described from
Florida, Kansas, Colorado, and New Mexico by Evans (1957, 1966).
We encountered the species at only one locality, 16 km west of La Paz,
on June 12. About 20 nests were scattered over the flat bottom of an
arroyo, about 0.3 km from the shore of Bahia Pichilingue. The nesting
area measured about 3 X 10 m, no nests being closer together than 0.5
m and most much more widely spaced than that. The area was sur-
rounded by tall cacti and bushes. We found no males at this site, and
all females showed much wear of the wings and mandibles.
Most nest entrances were open during the day, and a bombyliid fly,
Exoprosopa sima O.S., was observed flying from hole to hole, apparently
Ovipositing in them. Several Bembix females were provisioning during
the morning hours. One of them brought in 4 flies in a 30 minute period
from 1005 to 1035, in one instance requiring only 3 minutes to obtain
prey. In each case she plunged quickly into the open hole and emerged
within a few seconds. We dug out this nest in the afternoon, at which
time it was closed from the inside. It was surprisingly shallow consider-
ing the extreme heat and very dry soil in this locality. The burrow was
oblique, 46 cm long, reaching a cell at a vertical depth of 22 cm. The
female was about 35 cm from the entrance and there was a small closure
between her and the cell as well as one at the entrance. The cell was 4
cm long by 1.5 cm high and contained a half grown larva, 8 fresh flies,
and many fly remains.
The flies in this cell were identified as Diacrita costalis Gerstacker (4)
(Otitidae), Peleteria neotexensis Brooks (2) (Tachinidae), and Copesty-
lum isabellina (Williston) (2) (Syrphidae). We also found the remains
of flies around several old cocoons unearthed during our excavations. A
sample of these revealed the following: 14 Syrphidae of two species, 3
Sarcophagidae, 1 Callitroga (Calliphoridae), and 1 Diacrita costalis
(Otitidae).
We observed two instances in which females were engaged in building
large mounds of sand, presumably following final closure of the nest.
These females would land on top of the mound, then turn off to one side
for a distance of 7-10 cm kicking sand behind them, then make a brief
flight and land on top of the mound to repeat the performance over a
slightly different track. One female made 40 such lines radiating from
the mound over a period of 50 minutes, then started a new nest 40 cm
away. This “mound-building behavior” was reminiscent of that of B.
littoralis Turner, an Australian species which, however, performs the
behavior at the initial nest closure (Evans and Matthews, 1973). It is
also reminiscent of the still larger mound of Stictia lineata (Fabricus) in
318 THE Pan-PaciFic ENTOMOLOGIST
South America (Evans and Matthews, 1974). The function of these
large mounds is in no case understood.
Measurements of one of the B. sayi mounds showed it to be 25 cm
long by 17 cm wide and 2.5 cm deep in the center. In one case we were
able to locate an oblique, open burrow, 20 cm long, at one end of the
mound. This is presumably the “back burrow” described for U.S. popu-
lations. Although U.S. wasps make an elaborate final closure involving
radiating lines, then make a “back burrow,” leaving the mounds from
both burrows intact, there is no present evidence that they actually
build up these mounds (Evans, 1966, Fig. 157). This example of ap-
parent geographic variation in behavior seems worthy of further study.
BEMBIX OCCIDENTALIS W. J. Fox
We observed females of this species capturing flies from dead fish
lying on the beach a few km west of La Paz. We did not discover where
they were nesting. The species was not encountered within the nesting
areas of B. sayi or B. rugosa. It is, however, widely distributed in Baja
California Sur, and in fact part of the type series was from San José
del Cabo. There are specimens in the California Academy of Sciences
from Mulege and Coyote Cove (Bahia Concepcion), both on the Gulf
of California.
BICYRTES VARIEGATA (OLIVIER)
Two females of this species were seen digging on the beach, well below
the high water mark, 16 km west of La Paz. C. L. Fox (1923) recorded
variegata from La Paz and from Bahia Concepcion, both records also’
being from June.
MICROBEMBEX ARGYROPLEURA BOHART
We took 1 male of this species near San Carlos, and 4 males in dunes
near Puerto Chale, 60 km south of San Carlos. It has not previously
been reported from south of El Arco, Baja California Norte (Bohart,
1970). Quite possibly some of the Microbembex reported by C. L. Fox
(1923, 1926) from the La Paz area properly belong to this recently
described species. Alcock (1975) has studied its nesting behavior in
Arizona.
STENIOLIA DUPLICATA PROVANCHER
This is an abundant wasp in Baja California Sur. We collected 2
females and 3 males at Todos Santos, 1 female near La Paz, and 2 males
77 km northwest of La Paz. Gillaspy (1964) presented several addi-
Vou. 52, No. 4, OcToBER 1976 319
tional records. Evans and Gillaspy (1964) reported on several nests
from western Texas.
GLENOSTICTIA BITUBERCULATA (PARKER)
We took two males at flowers of Umbelliferae at Todos Santos. The
species has not previously been reported from Baja California, but I
have also seen a male from 1.5 mi. east of San Jorge, B.C. Sur, collected
July 25, 1971 (Real & Main) [Calif. Acad. Sci.]. Our specimens were
compared with the type in the U.S. National Museum.
GLENOSTICTIA BIFURCATA (C. L. Fox)
We also took 1 male of this little known wasp on Umbelliferae at
Todos Santos. The species was described from Baja California Norte
(Isla Angel de la Guarda and Bahia de Los Angeles). C. L. Fox (1923)
also described Stictiella directa from Bahia de los Angeles and recorded
it from Isla del Carmen and Isla Espiritu Santo, B.C. Sur. I have
studied part of his type series and regard directa as a synonym of bi-
furcata (new synonymy).
GLENOSTICTIA GILVA GILLASPY
We took a female and 3 males on flowers near La Paz, June 11-14.
This species was misidentified as exigua W. J. Fox by C. L. Fox (1923),
who recorded it from Isla Espiritu Santo and Todos Santos. Notes on
the nests and prey were presented by Evans (1966) and by Alcock
(1975).
ACKNOWLEDGMENTS
I am indebted to the following specialists for identifying the fly prey
and parasites: G. Steyskal, C. W. Sabrosky, L. Knutson, W. Wirth, and
F. C. Thompson. I am especially indebted to Darryl T. Gwynne
and William L. Rubink, who accompanied me on the trip to Baja Cali-
fornia Sur and assisted in collecting and making field observations.
This paper is part of a study of the comparative behavior of solitary
wasps, supported by the National Science Foundation, grant GB43790.
LITERATURE CITED
Aucock, J. 1975. The behavior of some bembicine wasps of southern Arizona
(Hymenoptera: Sphecidae, Microbembex, Glenostictia, Xerostictia) .
Southwest. Nat., 20: 337-342.
Bowart, R. M. 1970. New species, synonymy and lectotype designation in North
American Bembicini. Pan-Pac. Entomol., 46: 201-207.
320 THE Pan-Paciric ENTOMOLOGIST
Evans, H. E. 1957. Studies in the comparative ethology of digger wasps of the
genus Bembix. Comstock Publ. Assoc., Cornell Univ. Press, Ithaca, N.Y.
248 pp.
Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps.
Harvard Univ. Press, Cambridge, Mass. 526 pp.
Evans, H. E. anp J. E. Gittaspy. 1964. Observations on the ethology of digger
wasps of the genus Steniolia (Hymenoptera:Sphecidae:Bembicini).
Amer. Midl. Nat., 72: 257-280.
Evans, H. E. anp R. W. Matruews. 1973. Systematics and nesting behavior of
Australian Bembix sand wasps. Mem. Amer. Entomol. Inst., 20: 1-387.
Evans, H. E. anp R. W. Mattuews. 1974. Observations on the nesting behavior
of South American sand wasps (Hymenoptera). Biotropica, 6: 130-134.
Fox, C. L. 1923. Expedition of the California Academy of Sciences to the Gulf
of California in 1921. The Bembicini (digger wasps). Proc. Calif.
Acad. Sci., (4)12: 429-436.
Fox, C. L. 1926. Expedition to the Revillagigedo Islands, Mexico, in 1925. The
Bembicini (digger wasps). Proc. Calif. Acad. Sci., (4)15: 219-222.
Gittaspy, J. E. 1964. A revisionary study of the genus Steniolia (Hymenoptera:
Sphecidae:Bembicini). Trans. Amer. Entomol. Soc., 89: 1-117.
ZOOLOGICAL NOMENCLATURE
Required six months’ notice is given of the possible use of plenary powers by the
International Commission on Zoological Nomenclature in connection with the fol-
lowing names listed by case number.
ANNOUNCEMENT A. N. (S) 99
See Bull. Zool. Nom. 33 part 1, 26th June, 1976.
896. Tipula oleracea (Diptera: TIPULIDAE): revived proposals for stabilizing
names in species-group.
ANNOUNCEMENT A. N. (S) 100
See Bull. Zool. Nom. 33 part 2, 30 September 1976.
Z. N. (S.) 2140 ERIOCOCCIDAE Cockerell, 1899, proposed conservation of, and
Eriococcus Targioni-Tozzetti, 1868, proposed designation of type-
species for (Insecta: Homoptera).
Comments should be sent in duplicate, citing case number, to the Secretary,
International Commission on Zoological Nomenclature, c/o British Museum (Nat-
ural History), Cromwell Road, London, SW7 5BD, England, if possible within 6
months of the date of publication of this notice. Those received early enough will
be published in the Bulletin of Zoological Nomenclature. R. V. MELVILLE, Secretary
to the International Commission on Zoological Nomenclature.
VoL. 52, No. 4, OcToBER 1976 321
A new Rheumatobates from Costa Rica
(Hemiptera: Gerridae)
Joun T. PoLtHEmust
3115 So. York, Englewood, Colorado 80110
AND
LANNA CHENG
Scripps Institution of Oceanography, P.O. Box 109, La Jolla, California 92037
The water strider described here was found in a mangrove swamp.
The name is published so that it will be available for a forthcoming
work on marine insects. For all measurements 60 units = 1 mm, except
where given in mm.
Rheumatobates ornatus, n. sp.
Length, apterous male, 2.5 mm; macropterous forms unknown. Color deep brown
to black; dorsum of thorax and abdomen mostly frosted. Posterolateral portion of
thorax, head except median longitudinal black stripe, dorsum of first genital seg-
ment, connexival margins, orange brown. Broad medial area of pronotum, pleura,
most of mesosternum, prosternum and venter of head yellowish to leucine. An-
tenna brown, lighter ventrally and on base of segment I. Fore femur basally, fore
trochanter, posterior trochanter, yellowish brown; remainder of legs brown to deep
brown. Rostrum brown.
Structural characteristics. Antenna, fore and middle legs of male modified.
Antennal formula LIV: male, 40:3:48:37; female, 23:4:27:26. Male antennal
segment I as in Fig. le, with tufts of stout hairs basally on segment II directed
caudally and ventrally, also long (7) stout curved spine directed ventrally at distal
three-fourths on segment III (not visible in figure). Head (between eyes) of male
long (30), broad (27); shorter (27) but equally broad (27) in female. Male
abdominal dorsum broadly, transversely depressed, tergite II lowest; venter of seg-
ment VII modified to form a raised clasper (Figure la). Connexiva of male almost
vertical, flatter in female. Male fore femur with a dorsal knob basally; middle
trochanter with knob directed dorso-posteriorly; posterior trochanter with dorsal
knob basally. Anterior tibia of male flattened, with two stiff leucine brushes
distally, one dorsal, one ventral; adjacent to dorsal leucine brush is another brush
of short stiff black bristles; tibia and tarsus modified as shown in Figure lc, d.
Middle femur of male highly modified, excavate above medially, ridged along
entire length of caudal margin dorsally, set with two rows of recurved hairs plus
several rows of stiff setae directed both anteriorly and posteriorly (Figure 1b) ;
excavation with a brush of long, stiff black setae which are recumbent or obliquely
posteriorly directed (not visible in figure). Acetabula of male excavate laterally,
forming an oblique, almost vertical sulcus bordered anteriorly by a tumescence
1 Contribution from the Department of Environmental, Population and Organismic Biology, University
of Colorado, Boulder 80302 and Martin Marietta Corp., Denver, CO 80201.
Tue Pan-Paciric ENTOMOLOGIST 52: 321-323. OcroBEeR 1976
322 THe Pan-Paciric ENTOMOLOGIST
\/@mm
Fic. 1. Rheumatobates ornatus n. sp., male. a. Seventh abdominal and genital
segments, ventral view. b. Middle femur and trochanter, dorsal view. c. Fore leg,
dorsal view. d. Tip of fore tarsus and claws. e. Antenna, dorsal view.
bearing a brush of stiff posteriorly directed black setae, which overlay the sulcus.
Female similar to male in coloration; length (3 mm). Appendages without
modification.
Material: Holotype (4), allotype (2), and paratypes, 316 6, 162 9, COSTA
RICA, Mata de Limon, 29 Dec. 1974, L. Cheng. Holotype, allotype and 2 paratypes
in California Academy of Sciences; paratypes in Polhemus Collection and Scripps
Oceanographic Institution.
Discussion: This species is the only known Rheumatobates with an
excavate and expanded middle femur, and it is not close to any described
species. R. crassifemur Esaki and its subspecies have the middle femur
swollen but not highly modified. The type locality is close to Puntare-
nas, and not far from Boca de Barranca; the latter is the type locality
of another recently described Rheumatobates associated with the marine
habitat (Polhemus, 1975).
Biological Notes: The mangrove swamp where R. ornatus was col-
lected is quite extensive, and connected to the sea only at high tide. The
insects were very abundant among the aerial roots of Rhizophora man-
gle, occurring in groups of 50-100. They came out in the open only
when chased from the roots. Both adults and nymphs were present, and
several mating pairs were captured. They were found with fish larvae
VoL. 52, No. 4, OcToBER 1976 323
that occurred in great numbers in certain areas. The habitat as well as
behavior of these skaters were very similar to those of Rheumatobates
aestuarius Polhemus found in Baja California (see Cheng and Lewin,
1971).
LITERATURE CITED
Cuenc, L. anp R. A. Lewin. 1971. An interesting marine insect, Rheumatobates
aestuarius (Heteroptera:Gerridae), from Baja California, Mexico. Pac.
Ins. 13(2): 333-341.
PotHemus, J. T. 1975. New estuarine and intertidal water striders from Mexico
and Costa Rica (Hemiptera:Gerridae, Mesoveliidae). Pan-Pac. Entomol.
51(3): 243-247.
SCIENTIFIC NOTE
Records of Rearing of Tachinids (Diptera) from Insects, Mostly
in Washington.’"—During the summers of 1965-7 numerous insects, largely
Lepidopterous larvae, were collected in the vicinity of Pullman and elsewhere in
Washington by Bert A. Freeman and reared at the laboratory in Pullman for de-
termining their tachinid parasites and parasite-host relationships. Because of the
illness of Mr. Freeman the project had to be terminated prematurely and was,
consequently, incomplete. Considerable unpublished information was obtained,
however, and this is presented here.
Various collecting methods were used, but in general larvae of different instars
and sometimes pupae and adults were collected under natural conditions and
brought into the laboratory where rearing was continued until the parasites
emerged and until adult hosts were obtained for purposes of identification or con-
firmation of their identities. Winter collecting was made under loose bark and in
leaf duff on the ground.
Some tachinid pupae do not seem very tolerant of artificial media or holding
conditions and will not emerge unless natural conditions are closely simulated.
For the most part, however, satisfactory results were obtained by exposing tachinid
pupae to temperatures of 3° C for two months and then bringing them gradually
out of the cold. For winter holding of tachinid and lepidopterous pupae sawdust
from kiln-dried lumber proved the best. It could be made to hold sufficient mois-
ture and, under eastern Washington conditions, very little fungous growth is en-
countered if the sawdust is changed every three or four weeks.
In the following annotated list common names are given only for insects other
than Lepidoptera; lack of such indicates a lepidopterous host. Unless otherwise
indicated the host is in the larval stage. Collections and rearing, unless otherwise
indicated, were done by Mr. Freeman. Some information based on rearing of in-
sects obtained other than in Washington is included. The order of listing is alpha-
betical by tachinid parasites. Dates are those of the collections of the hosts, not of
emergence of the parasites.
1 The work was supported by USDA grant 12-14-100-8005(33). Scientific paper number 4526, Wash-
ington State University College of Agriculture Research Center, Pullman, Washington. Work was
conducted under project 1852.
324, THE PaAn-PaciFic ENTOMOLOGIST
Athrycia cinerea (Coquillett), ex Prodenia praefica Grote, Prosser, WA., Sept.
3, 1965.
Bessa harveyi (Townsend), ex larch sawfly, Pristiphora erichsoniit (Hartig), Mt.
Spokane, WA., July 24, 1965, and Moscow Mt., near Moscow, IDA., Aug. 6, 1967.
Bigonicheta spinipennis (Meigen), ex adult European earwig, Forficula auricularia
Linnaeus, Pullman, WA., July 5, 1965, 42 parasites obtained from 209 hosts. Ex
Archips cerasivoranus (Fitch), Puyallup, WA., summer, 1965, reared by E. P.
Breakey.
Blondelia hyphantriae (Tothill), ex Celerio lineata (Fabricius), Pullman, WA.,
July 16, 1967.
Compsilura concinnata (Meigen), ex Malacosoma disstria Hiibner, Pullman, WA.,
May 24, 1965, 37 parasites from 347 hosts; Pullman, WA., May 25 and June 1,
1965; Puyallup, WA., June 14, 1965; South Prairie, WA., June 3, 1967. Ex
Malacosoma pluviale (Dyar), Pullman, WA., June 17, 1965, 6 parasites from 87
hosts. Ex Stilpnotia salicis (Linnaeus), Pullman, WA., July 2, 1965, 19 para-
sites from 69 hosts. Ex Vanessa cardui (Linnaeus), Colton, WA., June 18, 1967.
Ex Halisidota argentata Packard, Enumclaw, WA., June 24, 1967. Ex Pieris
rapae (Linnaeus), Puyallup, WA., reared by Louis Getzin.
Doryphorophaga doryphorae (Riley), ex Colorado Potato beetle, Leptinotarsa de-
cimlineata (Say).
Eleodiphaga pollinosa Walton, ex adult tenebrionid, Eleodes hispilabris Say, Pros-
ser, WA., June 10, 1967; Snake River, near Pullman, WA., March—April, 1967,
10 parasites from about 100 hosts. Ex adult tenebrionid, Eleodes hovaverrucula
Boddy, reared “in numbers.” Ex adult tenebrionid, Eleodes sp., Steptoe Canyon,
near Colton, WA.
Erynnia tortricis (Coquillett), ex Archips cerasivoranus (Fitch), Pullman, WA.,
June 20, 1965, and July 14, 1967. Ex Lespeyresia pominella (Linnaeus), dia-
pausing larvae, Yakima, WA., Feb., 1967, R. B. Hutt.
Eusisyropa blanda (Osten Sacken), ex Hyphantria cunea (Drury), Puyallup, WA.,
June 5, 1965; Orting, WA., Oct. 11, 1966. Ex Archips argyrospilus Walker,
Orting, WA., Aug. 6, 1967.
Exorista mella (Walker), ex Halisidota maculata (Harris), last larval instar, Step-
toe Butte, WA., Oct. 6, 1966. Ex Datana ministra (Drury), Kamiak Butte, near
Palouse, WA., Oct. 4, 1966. Ex Prodenia praefica Grote, Genessee, IDA., Aug.
23, 1965.
Euexorista futilis (Osten Sacken), ex Malacosoma pluviale (Dyar), on willow,
Pullman, WA., June 17, 1966.
Gymnoclytia occidentalis Townsend, ex pentatomid, Euschistus impictiventris Stal.
Hemisturmia tortricis (Coquillett), ex Archips argyrospilus (Walker), Orting,
WA., June 4, 1965, and Aug. 6, 1967.
Hyphantrophaga hyphantriae (Townsend), ex Hyphaniria cunea (Drury), Steptoe
Butte, near Palouse, WA., Sept. 27, 1965, and Sept. 29, 1966; emerged directly
from pupa of host.
Leschenaultia americana (Brauer and Bergenstamm), ex Malacosoma disstria
Hiibner, South Prairie, WA., June 14, 1967.
Lespesia sp., ex Papilio rutilis Lucas, Pullman, WA., Nov. 26, 1965 (?overwinter-
ing pupa).
Lespesia archippivora (Riley), ex Vanessa cardui (Linnaeus), Colton, WA., June
VoL. 52, No. 4, OcToBER 1976 325
18, 1967. Ex Prodenia praefica Grote, Prosser, WA., Sept. 3, 1965. Ex Colias
eurytheme Boisduval, Othello, WA., June 27, 1966.
Lixophaga variabilis (Coquillett), ex Laspeyresia pominella (Linnaeus), Wenat-
chee, WA., Aug. 16-20, 1967.
Mericia ampelus (Walker), ex Peridroma saucia (Hiibner), Orting, WA., July 30,
1966. Ex Hyphantria cunea (Drury), Steptoe Butte, near Palouse, WA., Aug.
16, 1965.
Microphthalma disjuncta (Wiedemann), ex scarabaeid, Polyphylla crinita LeConte,
Westport, WA., June 5, 1967, 1 parasite from 24 larvae.
Nemorilla pyste (Walker), ex Archips cerasivoranus (Fitch), collected as pupa,
Pullman, WA., June 20, 1965.
Phryxe pecosensis (Townsend), ex Heliothis phloxiphaga G. & R., Yakima, WA.,
June 12, 1966.
Phryxe vulgaris (Fallén), ex Heliothis phloxiphaga G. & R., Yakima, WA., July
12, 1966, 1 parasite from 21 larvae. Ex Pieris rapae (Linnaeus), Prosser, WA.,
May 19, 1965.
Tachinomyia sp., ex Halisidota maculata (Harris), on willow, Steptoe Butte, near
Steptoe, WA., summer, 1966.
Tachinomyia similis (Williston), ex Malacosoma dissiria Hiibner, South Prairie,
WA., June 14, 1967. Ex Stilpnotia salicis (Linnaeus), Pullman, WA., June 2-4,
1967.
Thelymyia mathesoni (Reinhard), ex Hemerocampa pseudotsugata McDunnough,
Kamiak Butte, WA., Aug. 15, 1966, collected as last larval instar, parasite
emerged from pupa.
Uramya halisidotae (Townsend), ex Halisidota argentata Packard, under Douglas
fir on ground after windstorm, Tacoma, WA., June 10, 1966; parasites emerged
from last larval instar just prior to pupation.
Winthemia (?rufopicta Bigot), ex Peridroma saucia (Hibner), Pullman, WA.,
June 17, 1966.
Winthemia quadripustulata (Fabricius), ex Haliothis zea (Boddie), Grandview,
WA., Aug. 28, 1967.
Zenillia (? blandita (Coquillett) ), ex Hyphantria cunea (Drury).
A hyperparasite, the chalcidoid Dibrachys cavus (Walker), was commonly ob-
served and reared from puparia of Bigonicheta spinipennis (Meigen) during the
winter months. Incidence of parasitism ranged from 22 to 35 percent, with three
to nine larvae occurring in each of the parasitized pupae. Pupae of the tachinid
host were common under the loose bark of cottonwoods and poplars during the
winter months in the Pullman area.
The above rearing records extend the geographical distribution recorded in the
Stone et al. (1965 USDA Agric. Handbook, 276) catalog as follows: Eleodiphaga
pollinosa Walton, Washington, previously recorded from New Mexico and Califor-
nia; Thelymyia mathesoni (Reinhard), Washington, previously recorded from New
York to Manitoba and Ohio.— Maurice T. James, Professor Emeritus of Entomol-
ogy, Washington State University, Pullman 99163.
326 THE PaAn-PaciFIC ENTOMOLOGIST
A New Species of Heteromurus from the Solomon Islands
(Collembola: Entomobryidae)
José A. Mart Mutt
University of Illinois, and Illinois Natural History Survey, Urbana, 61801
Through the courtesy of Dr. P. N. Lawrence and the British Museum
(Natural History) I have studied the Orchesellini collected by the Royal
Society Expedition of 1965 to the Solomon Islands. The tribe was
represented by three species: the one described herein and two new spe-
cies of Dicranocentrus to be described in an upcoming revision of that
genus.
The species described below is named after Dr. Lewis J. Stannard, Jr.,
a leading authority on the Thysanoptera, who has recently retired from
the Illinois Natural History Survey after 30 years of service.
Heteromurus stannardi, n. sp.
Length up to 1.9 mm. General appearance typical of the genus (Fig. 4), the
fourth abdominal segment not being over 1.6 times longer than the third. Body
background color yellow, very diffuse purple pigment is distributed over the head,
body, antennae, and leg segments up to the coxae. Head, body, and appendages
(with the exception of the last two antennal segments) covered with scales (Fig.
7).. Antennae five segmented, about half as long as the body. Ant. 5 annulated.
Two eyes present on each side of the head (Fig. 5). Trochanteral organ composed
of about 26 setae. Tibiotarsi covered with ciliated setae, devoid of smooth setae
with the exception of the opposite seta to the tenent hair, which is present on the
third pair of legs. Claw structure as in figure 6. Tenent:hair apically lanceolate,
about as long as the unguiculus. Dental spines absent. Mucro as in figure 3, lack-
ing a basal spine. Head and body macrochaetotaxy as in figures 1 and 2.
Material Examined: Solomon Islands, Guadalcanal, Popamanasiu, 1,320 m. el.,
November 1-4, 1965, mossy ridge forest litter, P. N. Lawrence, coll. Royal Society
Expedition, B. M. 1966-1. Holotype and 15 paratypes. Two paratypes are deposited
in the collection of the Illinois Natural History Survey, the rest of the material is
in the British Museum.
Diagnosis: By the persistent absence of the mucronal basal spine, the
species comes closest to Heteromurus nitens Yosii (1964), from the
Tonga Islands, and to the African species Heteromurus subduvius Barra
(1968). Heteromurus stannardi may be readily separated from these
species by the structure of the claws and body macrochaetotaxy, which
by the way, is different from that of any species for which the pattern is
known.
Tue Pan-Paciric ENtomo.ocist 52: 326-330. Octoser 1976
Vou. 52, No. 4, OcTroBer 1976 327
5
Fics. 1-3. Heteromurus stannardi. Fig. 1. Dorsal head macrochaetotaxy, each
dot represents one seta. Fig. 2. Dorsal body macrochaetotaxy. Fig. 3. Mucro.
>
Fics. 4-5. Heteromurus stannardi. Fig. 4. Photomicrograph of a paratype.
100. Fig. 5. Scanning electron micrograph (SEM) of the eyes. 3,000.
Fics. 6-7. Heteromurus stannardi. Fig. 6. SEM of the hind claw. 1,300.
Fig. 7. SEM of dorsal abdominal scales. 1,600.
328 THE Pan-PactFic ENTOMOLOGIST
VoL. 52, No. 4, OcToBER 1976
330 THE Pan-Paciric ENTOMOLOGIST
LITERATURE CITED
Barra, J. A. 1968. Contribution a l’étude du genre Heteromurus Wankel. 1860
(Collemboles). Biol. Gabonica 4: 105-117.
Yosu, R. 1964. Some Collembola of the Tonga Islands. Kontyi 32: 9-17.
RECENT LITERATURE
AGRICULTURAL I[NsEcT Pests OF THE Tropics & THemR Controt. Dennis S. Hill.
1975. 516 pp. Cambridge University Press, 32 East 57th Street, New York,
New York 10022. Price $34.50.
Factors AFFECTING DISPERSAL DISTANCES OF SMALL ORGANISMS. D. O. Wolfen-
barger. 1975. 230 pp. Exposition Press, Inc., 900 So. Oyster Bay Road,
Hicksville, New York 11801. Price $15.00.
GuIDE TO THE ADULT AND LARVAL PLUSIINAE OF CALIFORNIA (LEPIDOPTERA:
Noctuwar). Thomas D. Ejichlin. August 15, 1975. Occasional Papers in
Entomology, no. 21, 73 pp. State of California, Department of Food and Agri-
culture, Division of Plant Industry, 1220 N Street, Sacramento, California
95814. Individual copies sent free of charge upon request.
Insect PHystoLocy AND ANATomy. N. C. Pant and Swaraj Ghai (eds.). 1973.
276 pp. Division of Entomology, Indian Agricultural Research Institute, New
Delhi 110012, India. Rs 6.00.
Livine Insects [THE AUSTRALIAN NATURALIST LiprARy]. R. D. Hughes. 1975.
304 pp. Taplinger Publishing Company, 200 Park Avenue South, New York,
New York 10003. Price $14.95.
Mires InjuR1ous To Economic Puiants. Lee R. Jeppson, Hartford H. Keifer, and
Edward W. Baker. 1975. 638 pp. University of California Press, 2223 Fulton
Street, Berkeley, California 94720. Price $30.00.
THe Woritpd oF THE HoneyBee. Colin G. Butler. 1975. 226 pp. Taplinger
Publishing Company, 200 Park Avenue South, New York, New York 10003.
Price $14.95.
Tue ButTerFiies OF NortH America. William H. Howe (coordinating editor
and illustrator) and twenty contributors. 1975. 633 pp., 97 colored plates.
Doubleday & Company, Inc., Dept. ZP-510, Garden City, New York 11530.
Trade edition, price $39.95. Limited edition, limited to 200 copies, numbered
and signed by the artist, including a print of special painting suitable for
framing, price $150.00.
VoL. 52, No. 4, OcToBER 1976 331
Nesting Behavior of Encopognathus rufiventris
Timberlake
(Hymenoptera: Sphecidae)
R. M. Bowart anp B. VILLEGAS
Department of Entomology, University of California, Davis 95616
Encopognathus is a relatively primitive or unspecialized genus of
the tribe Crabronini. According to Bohart and Menke (1976), there
are 23 described species occurring in widespread temperate and tropical
regions of the world except Australia. The only biological reference
of which we are aware is a brief note by Arnold (1932) that E.
chirindensis (Arnold) provisioned with ants of several species in
Rhodesia. In light of our findings this may have been in error.
ENCOPOGNATHUS RUFIVENTRIS TIMBERLAKE
Nesting site—A small colony was observed in Arroyo Seco Camp
in the hilly Upper Sonoran Life Zone of Monterey County, California
at an elevation of 246 meters. From May 28—June 13, 1975 about 35
nests were located in a two square meter area of hard-packed sandy soil
alongside wheel tracks of a seldom used dirt road. Perhaps 20 females
were active at a time, some constructing burrows and others provision-
ing. Entrances were scattered but some were only a few cm apart. Daily
activity at the nest site extended from 10:15 AM to 5:00 PM, Pacific
Standard Time. Females were engaged mostly in nest construction
during the morning and provisioning during the afternoon. No males
were seen at the nest site but a few were collected on small yellow com-
posites several hundred feet away. Females exhibited an erratic hover-
ing flight for two to three seconds just before entering or just after
leaving nests.
Nest structure.—Ten nests were excavated and dissected, some at the
site and others after removal to the laboratory. The entrances were
about 3.5 mm in diameter, open at all times, and without a chimney
although sometimes under leaves. There was no accumulation of dirt
around the entrance since the female carried away excavated material
in her mandibles and front legs. In this process she flew backwards
from the nest opening for about 25 cm before dropping the load. Typi-
cal burrows are diagrammed in figs. 3 and 4. Each main tunnel was
relatively simple, somewhat undulate, and slanting into the ground for
9.5 to 12 cm. The tunnel ended in a whorl of three to five branches with
cells arranged in linear fashion (figs. 3, 4). Loose sandy soil was
Tue Pan-Paciric Entomotocist 52: 331-334. Octroper 1976
332 THE Pan-Paciric ENTOMOLOGIST
i
‘et
'
uy
‘ J
Crt
par.
yy
{li
|
(
Lea}
—L.
Fics. 1-4. Encopognathus rufiventris Timberlake. Fic. 1, mature larva, lateral
view. Fic. 2, facial view of mature larva with enlargements of labium and mandible.
Fics. 3-4, profiles of typical nests.
placed between the cells by the wasp as she worked back toward the
entrance. Each oval cell was about 3.5 mm wide and 6.5 mm long.
The greatest number of cells observed in a single nest was 16.
Provisioning.—Prey were small Miridae, all but a few of them adults.
Four species were utilized and these were apparently obtained from
nearby plants of Verbena lasiostachys and Lotus scoparius. Most fre-
VoL. 52, No. 4, Ocroper 1976 333
quent prey were Macrotylus lineolatus Uhler. Other mirids used were
Psallus seriatus (Reuter) and several species of Plagiognathus (deter-
minations by Jon Herring, U.S. Department of Agriculture, Division
of Insect Identification, etc.). The female held the paralyzed prey in
the middle legs as she returned to the nest. Near the opening the wasp
displayed a short hovering and somewhat sidewise motion before plung-
ing into the open nest. The time absent from the nest by 10 actively
provisioning females was 50 seconds to six minutes (average 136
seconds). The prey were packed tightly, head forward or the reverse,
and 12 to 18 per cell.
Immature stages —-No eggs were found. Larvae pushed the uneaten
bug remains to one end of the cell. At maturity a silk lined cocoon was
formed which was completely covered with brownish sand grains as
illustrated for Entomognathus by Miller and Kurczewski (1972:fig. 4).
The cocoons were about 3.5 mm wide and 6.0 mm long. Morphologi-
cal details of a mature larva are shown in figs. 1 and 2. Figure 1 was
drawn from a specimen preserved in alcohol. Figure 2 was drawn from
a head briefly treated in hot KOH and mounted in balsam on a slide.
Parasites —Three females of an undescribed species of Spintharosoma
(Chrysididae) were collected after they were observed repeatedly en-
tering the burrows of Encopognathus rufiventris at the nest site.
DIscUSSION
Four presumably “primitive” genera of Crabronini have the following
prey relationships: nymphal and adult mirid bugs used by Enco-
pognathus, Lygus and other adult mirids by Anacrabro, leafhoppers
by Entomocrabro, and chrysomelid beetles by Entomognathus. The use
of different orders of insects by related genera may seem unusual. How-
ever, Belomicrus, which has many points of morphological resemblance
with Encopognathus but is in a separate tribe Oxybelini, is known to
provision with beetles in one section of the genus and nymphal and adult
mirids in another section (Bohart and Menke, 1976). Among pub-
lished reports on biology and larval morphology, the greatest simi-
larities with Encopognathus rufiventris have been found in the study
of Entomognathus memorialis Banks by Miller and Kurczewski (1972).
Burrow formation, larval structure, and cocoon appearance are all
much the same as those described and figured by the above authors.
Many small sphecids do not make a formal closure of the nest on
completion and EF. rufiventris belongs to this sort. The entrance is prob-
ably filled in a short time by blowing dust or as a result of light rains.
334 THe Pan-Paciric ENTOMOLOGIST
E. rufiventris females have a very short foreleg pecten as would be ex-
pected in a wasp which omits nest closure.
LITERATURE CITED
ARNOLD, G. 1932. New species of the Ethiopian Sphegidae. Occas. Pap.
Rhodesian Mus. (1)1: 38.
Bowart, R. M. anp Menke, A. S. 1976. Sphecid wasps: A world generic re-
vision. University of California Press Berkeley. 695 pp.
Mitier, R. C. anp Kurczewski, F. E. 1972. A review of nesting behavior in the
genus Entomognathus, with notes on E. memorialis Banks. Psyche 79:
61-78.
RECENT LITERATURE
Les CARABIDAE DU QUEBEC ET DU LABRADOR. A. Larochelle, Department de Biologie
de College Gourget, Rigand. Bull. I. 255 pp. 1975. Available from author,
$15.00 (Canadian).
This publication is difficult to categorize. The content is nearly evenly split
between an annotated catalogue of the species of Carabidae and Cicindelidae
known to occur in Quebec, and an atlas of distribution maps. Minor segments are
devoted to providing instructions for use of the catalogue and atlas, lists of
collections consulted, lists of species doubtfully recorded or to be expected in
Quebec, and a bibliography. Printing is by the offset process, with unjustified
right margins. A notice near the beginning of the work states that only 400 copies
have been printed, possibly explaining the relatively expensive price. Primarily
of interest to specialists of Carabidae or of the fauna of eastern Canada.—Kditor.
THE STONEFLIES, OR PLECOPTERA, OF ILLINoIs. T. H. Frison. Bulletin of the
Illinois Natural History Survey, 20:281-471. (Reprinted by Entomological
Reprint Specialists, P.O. Box 77224, Dockweiler Station, Los Angeles, California
90007). $15.00.
THe MAYFLies oR EPHEMEROPTERA, OF ILLINOIS. B. D. Burks. Bulletin of the
Illinois Natural History Survey, 26:1-216. (Reprinted by Entomological
Reprint Specialists, P.O. Box 77224, Dockweiler Station, Los Angeles, California
90007). $15.00.
These works remain useful summaries of the systematics, distribution and biology
of the taxonomic groups which are included. Despite the geographic restriction
suggested by the titles, the effective area of coverage is much larger, in terms of
the proportion of North American species included. Both volumes are generously
illustrated, and include lucid discussions of life history and ecology of these
important aquatic insects.—Editor.
VoL. 52, No. 4, OcroBER 1976 330
Observations on the Nesting Behavior of
Ammophila _ nasalis
(Hymenoptera: Sphecidae)
DarryL T. Gwynne!
Department of Zoology and Entomology, Colorado State
University, Fort Collins 80523
During June 8-9, 1975 I made some observations on the nesting be-
havior of Ammophila nasalis Provancher near San Carlos, Baja Cali-
fornia Sur, Mexico. These will add to the information on the nests and
prey of A. nasalis given by Hicks (1935) (under the name Sphex cras-
pedotus; see Menke 1965).
The wasps were nesting in the open spaces between low halophytic
shrubs in a large flat area of firm sand. A single nest of A. nasalis was
excavated by Dr. A. S. Menke at a similar nesting site on the University
of California at Santa Barbara campus at Goleta (June 1959) (personal
communication). Hicks’ observations were made along a sandy river-
bank of the Los Angeles river near Burbank, California.
During mid afternoon on June 8 female A was observed digging a
nest. She made several successive flights backwards from the nest
entrance dropping a spray of sand 20-30 cm from the nest entrance
and 10-20 cm above the ground. Most Ammophila species deposit soil
from burrow excavation while in flight (Evans 1959).
At 1345 on June 9 female B was observed leaving her nest, which had
a small white shell as a temporary closure. She flew about 30 meters
and started to investigate some of the nearby shrubs. During the after-
noon she was observed to return to the nest five times, carrying prey in
her mandibles. Upon landing at the nest she first dropped the prey then
removed the shell from the burrow entrance (fig. 1). She then grasped
the prey and backed into the burrow, then replaced the shell when she
left. This sequence was observed during the first four times she re-
turned to the nest. Similar observations were reported by Hicks (1935).
The hunting times ranged from 7 to 12 min. (XK =11, N=4). When
she left on the fifth hunting flight the nest was carefully excavated so
that one side was open (fig. 2); all of the caterpillar prey were removed
except the one bearing the egg. A plastic stake was used to cover the
exposed side of the burrow and both the sand and the temporary shell
closure were replaced. The female returned in 75 min. and entered the
1 Part of a study of the comparative behavior of solitary wasps, supported by the National Science
Foundation grant GB 43790 to Dr. Howard E. Evans.
Tue Pan-Paciric Entomouocist 52: 335-338. OctToser 1976
336 THE Pan-PaciFic ENTOMOLOGIST
*
Vou. 52, No. 4, OcToBER 1976 o07
nest with prey. She then made a final closure by pushing the shell down
into the burrow, depositing pieces of debris into the burrow with her
mandibles and using alternate movements of her front legs to push sand
into the burrow. The total time for the final closure was three minutes.
In his study of A. campestris Jur. (= pubescens Curtis) Baerends
(1941) found that each female takes care of more than one nest at a
time. He showed that the wasp visited each hole in the morning before
hunting. If prey were removed from a nest by an experimenter, before
the morning inspection, the female would bring in more prey than
usual to replace the missing ones. If cell contents were decreased after
the morning inspection the female did not bring in more prey. Ap-
parently A. nasalis mass provisions one burrow at a time but like A.
pubescens does not respond to the number of prey in the cell during the
day. This is evidenced by the above manipulation of cell contents; only
two small caterpillars, not enough to sustain a larva, remained with the
egg when the wasp made the final closure.
The shape of the burrow was typical of other Ammophila species as
given by Evans (1959) (fig. 2). The depth of the vertical burrows was
5.0 cm (female A) and 4.0 cm (female B). The single nest which was
excavated by Dr. Menke was 5.0 cm in depth but the three examined by
Hicks were deeper (8.2—8.6 cm).
Each burrow at the Baja California site ended in a horizontal cell
approximately 2.5 by 1 cm. The nest of female A, which was not com-
pleted when excavated, had a lump of sand as a temporary closure and
contained 10 geometrid caterpillars (identified by D. M. Weisman).
The egg was positioned transversely on the dorsal side of abdominal
segments 3 and 4 of the caterpillars. The nest of female B had a total
of 12 geometrid prey. Hicks (1935) also observed A. nasalis to use
geometrid prey but the single uncompleted nest at Goleta, California
contained a lycaenid caterpillar.
ACKNOWLEDGMENTS
I would like to thank Dr. Howard E. Evans for commenting on the
manuscript and Dr. A. S. Menke for additional information about A.
nasalis.
<
Fics. 1-2. Nesting of Ammophila nasalis. Fig. 1. Female B removing temporary
closure (shell) from burrow entrance. Fig. 2. Nest of female B showing the tem-
porary closure in the upper part of the burrow and geometrid caterpillars in the
cell.
338 THE PAn-PAciFIC ENTOMOLOGIST
LITERATURE CITED
Barrenps, G. P. 1941. Fortpflanzungsverhalten und Orientierung der Grabwespe
Ammophila campestris Jur. Tijdschr. Entomol., 84: 68-275.
Evans, H. E. 1959. Observations on the nesting behavior of digger wasps of the
genus Ammophila. Amer. Midl. Nat., 62: 449-473.
Hicks, C. H. 1935. Notes on rare western sphecid wasps. Pan-Pac. Entomol.,
11: 97-102.
Menke, A. S. 1965. The identity of some Ammophila observed by C. H. Hicks,
H. E. Evans and others in connection with biological studies (Hymenop-
tera, Sphecidae). Entomol. News, 76: 257-261.
SCIENTIFIC NOTE
Larvae of Coelus ciliatus parasitized by Reinhardiana petiolata (Coleop-
tera: Tenebrionidae; Diptera: Tachinidae). In May and June 1975 two
adults of Reinhardiana petiolata (Townsend) emerged from larvae of Coelus
ciliatus Eschscholtz collected from coastal sand dunes in California. Adults of
this fly are common over much of western North America (Stone, eé al., 1965,
Catalog of the Diptera of America North of Mexico, U.S. Government Printing
Office), but hosts of the immature stages were previously unknown for the tribe
Melisoneurini, in which Reinhardiana is classified (P. H. Arnaud, personal com-
munication).
Flies were reared from two collections of Coleus larvae. One adult female
emerged on V-22-1975 from a dead parasitized larva collected at Dune Lakes, 3 mi.
S Oceano, San Luis Obispo County, California, IV-29-1975. One adult male
emerged about VI-18-1975 from a larva collected at Point Reyes National Sea-
shore (South Beach), Marin County, California, IV-19-1975. This larva appeared
healthy when collected, showing no evident distress until the fly maggot pupated
between May 21-28, after killing the host. These dates indicate a pupal period
of 23-30 days.
The Reinhardiana larvae inhabit the haemocoele of the long, slender host
without causing a noticeable bulge until pupation. The fly puparia are about
twice the diameter of the beetle larva, whose body wall becomes stretched around
the parasite, giving the appearance of a snake containing a recent large meal.
Both puparia were lodged in abdominal segments 4-8, at about the midregion of
the host’s body. One puparium had the anterior end directed posteriorly in the
host, the other anteriorly. Emergence was through the dried body wall near the
apex of the puparium.
The broad ecological and geographic occurrence of Reinhardiana and the limited
Pacific coastal sand dune distribution of Coelus ciliatus suggest a broad host
range for this fly, possibly including other families of ground dwelling beetles.
The research was supported by National Science Foundation Grant BMS 74-
17924. P. H. Arnaud, Jr., kindly determined the adults of Reinhardiana, and assis-
tance with the fieldwork was provided by L. Fahrquar—Joun T. Doyen, Division
of Entomology and Parasitology, University of California, Berkeley 94720 and
C. N. Stospopcurkorr, Department of Biology, Northern Arizona University,
Flagstaff 86001.
Vout. 52, No. 4, OcTroBER 1976 309
A new species of Fuchsina Fall with
notes on some California Lathridiidae
(Coleoptera)
FRED G. ANDREWS
Insect Taxonomy Laboratory
California Department of Food and Agriculture
Sacramento, 95814
In his revision of North American Lathridiidae, Fall (1899) de-
scribed Fuchsina occulta from the coastal region of California. He
characterized this monotypic genus as “. . . the most extraordinary of
North American Corticariini” because of the 10-segmented antennae,
lack of eyes and differences in elytral and scutellar sculpture. He de-
scribed Revelieria californica in the same paper.
The Lathridiidae are known to feed on fungi in both larval and
adult stages (Hinton, 1941; Hammad, 1953). Some species are ar-
boreal while others are associated with moldy stored products, but most
are found in plant litter. Investigations of the litter fauna have largely
been in areas where there is a rich humus buildup. Along the Pacific
Coast this includes the Coastal Ranges, Transverse Ranges and west
side of the Sierra Nevada. The east side of the Sierra Nevada and the
basin ranges immediately to the east are in rain-shadow and plant litter
and humus are greatly reduced. Historically there has been little in-
vestigation of the soil fauna in this area. Recent sampling of the dry
sparse litter in this area using Berlese funnels has shown the Lath-
ridiidae to be abundant and widespread. Several new species have been
discovered, one of which will be treated in this paper.
This interesting but perplexing new desert inhabitant is a second
species in the genus Fuchsina. Inclusion of this species in Fuchsina
necessitates a slight redefinition of the genus. The following diagnosis
will suffice to separate Fuchsina from all other Corticariini.
FucuHsIna FALL
Small, reddish brown, parallel sided. Antennae 10- or ll-segmented with 3-
segmented club. Eyes absent or reduced to a single facet. Prosternal fovea
present. Scutellum transverse, without ridges or carinae, obtusely pointed poste-
riorly. Female with 5 abdominal sternites, male with a sixth present, shortened.
Fuchsina arida, new species
(Fig. 7)
Length, 1.2-1.4 mm. Width, 0.4-0.5 mm. Body elongate oval, convex, setiferous.
Cuticle shining, reddish brown. Head, thorax, scutellum and abdominal sternites
Tue Pan-Paciric ENtomo.uocist 52: 339-347. Ocrosper 1976
340 THe Pan-Paciric ENTOMOLOGIST
Fies. 1-6. Fuchsina spp.: 1) arida, head, single eye facet; 2) occulta, head,
eyeless; 3) occulta, elytral setation; 4) arida, elytral setation; 5) arida, scutellum;
6) occulta, precoxal fovea.
with reticulate microsculpture. Head transverse; tempora parallel-sided; frons
anterior to eyes sharply tapering to a blunt point; clypeus trapezoidal, expanded
anteriorly. Labrum trapezoidal, somewhat surrounding anterior clypeal margin.
Antennae 1l-segmented; Ist globose, 2nd ovoid, 3rd—8th of equal width, shortening
Vou. 52, No. 4, Ocroser 1976 o41
9) Revelieria californica Fall.
?
WHO OT O TNT OS
EO SOHO
8) Fuchsina occulta Fall
°
7
Fics. 7-9. Habitus: 7) Fuchsina arida new species
342 THE Pan-PaciFIc ENTOMOLOGIST
apically until transverse; club segments expanding, 9th oval, 10th transverse, 11th
with width one and one-half times length. Eye a single facet (Fig. 1). Pronotum
quadrate (width to length ratio 23:20); margin subdenticulate; punctation uni-
form, punctures separated by twice their diameter, fovea anterio-lateral to procoxae
(Fig. 6). Coxae conical, intercoxal process 14 diameter of coxa, closed behind.
Scutellum wider than long, ellipsoidal, microreticulate (Fig. 5). Elytra gently
inflated, smooth. Elytral setae in 14 or 15 confused rows, (Fig. 4) each emerging
from bilobed plaque. No elytral punctures. Tarsi with last 2 segments of equal
length. Aedeagus symmetrical.
Holotype. (Calif. Acad. Sci. No. 12394) Westgard Pass, Inyo Co., Calif., XI-
1972, berlese pinyon duff. D. Giuliani. Paratypes 2 same data, 12 same locality
except XII-72 and from juniper duff, 6 same locality except XII-72 and pinyon
duff.
Additional specimens. CALIFORNIA. Inyo Co.: Coyote Cr., 7 mi. SW Bishop,
7500’, IX-1972, D. Giuliani, pinyon duff [CDFA] (2); Fall Cr., 5200’, IX-1972,
D. Giuliani, golden oak duff [CDFA] (13); Grandview Camp, White Mts., 28-
VIII-1965, O. Clarke, Juniperus occidentalis duff, [CDFA] (17); Onion Valley
nr. Kearsarge Pass, 8-VIII-1964, O. Clarke, mixed leaf litter [CDFA] (1);
Waucoba Spring, 30-IV-1975, F. G. Andrews, A. R. Hardy, Pinus monophylla litter
[CDFA] (4); White Mts., 28-VII-1965, O. Clarke, Populus trichocarpa duff
[CDFA] (1). Modoc Co.: Benton Meadow, 12-VII-1974, soil [CISC] (11); Fan-
dango Pass, 4-VII-1974, soil [CISC] (2); Long Valley, 1-VII-1974 and 18-VII-
1974, soil [CICS] (2); Shields Creek, 26-VI-1974 and 5-VII-1974, soil and litter
[CISC] (4); Snell Spring, 26-VI-1974, 1-VII-1974, 2-VII-1974, soil [CISC] (11);
So. Deep Creek, 28-VI-1974 and 16-VII-1974, soil [CISC] (16). Riverside Co.:
Whitewater Canyon, 15-I]-1959, I. M. Newell, creosote bush duff [CDFA] (11).
San Bernardino Co.: 1 mi. NE Arrowbear, 6400’, 14-ITI-1964, E. L. Sleeper, pine-
oak duff [CDFA] (30); Joshua Tree Nat’]1 Monument, Fried Liver Wash, 4-II-
1967, Pinyon Wells, 14-V-1966 and 4-ITI-1967, Pleasant Valley, 7-I-1967, .7 mi. S
Squaw Tank, 11-XII-1965, E. L. Sleeper and S. L. Jenkins [LBSC] (5); New York
Mts., Bathtub Springs, 25-IV-1971, J. D. Pinto, leaf litter [CDFA] (1). San Diego
Co.: Borrego, Palm Canyon, 25-IV-1955, R. O. Schuster [CISC] (2). NEVADA.
Mineral Co.: 2 mi. W Mina, 6700’, IIJ-1973, D. Giuliani, pinyon duff [CDFA]
(3); Montgomery Pass, XII-1972, D. Giuliani, pinyon duff [CDFA] (4). ORE-
GON. Klamath Co.: Bly Mt., 19-III-1959, J. Schuh, yellow pine duff [JSCC] (1).
REMARKS: Fuchsina arida differs from F. occulta in having eleven-
segmented antennae; it resembles occulta in having (1) elytra without
striae or punctures, (2) large number of rows of setae (12 in occulta,
15: in arida) in somewhat confused lines, (3) scutellum without carina
or fovea, exposed portion subtriangular, (4) eye reduced (absent in
occulta, a single facet in arida) and (5) abdominal sternites numbering
5 in females, 6 in males. The naming of a new genus for arida based
upon the difference in antennal segment number (Fuchsina occulta is
the only known North American Corticariini to have other than an 11-
segmented antenna), or placing it in another existing Corticariini genus
Vou. 52, No. 4, OctoBEer 1976 343
TABLE 1. Morphological characters of the various genera of Corti-
cariini in North America.
Genus: Fuchsina Corticaria Corticarina) Melanophthalma Cortilena
No, of segments
in antennae &
antennal club
10-11/3 11/3 11/3 11/3 11/2
Posternal
fovea
Abdominal Q
present present absent absent absent
5
sternites CO 5
Elytral 12-14 con- 8 rows 8 rows ' 8 rows 8 rows
setation fused rows, punctate punctate punctate punctate
impunctate striae striae striae striae
Coxal lines
first abdominal | absent absent
absent present present
sternite
Relative length
tarsal segments
2&3
1>2 1>2
Shape of triangular truncate truncate truncate truncate
scutellum without carinate carinate carinate carinate
carina or or ridged or ridged or ridged or ridged
ridges
First tarsal smooth smooth smooth spinal smooth
segment projection
of male
a} genitalia symmetric symmetric asymmetric symmetric symmetric
was thought unwarranted because of overall similarity to occulta and
lack of similarity to other genera.
Table 1 depicts those characters historically used (Belon, 1897; Fall,
1899; Von Peez, 1967; Dejoz, 1970) to separate the various Corticariini
genera. It can be seen that Fuchsina as constituted by both occulta and
arida shows little overall similarity to any other North American genus
with the possible exception of Corticaria where a few characters are
shared.
FUCHSINA OCCULTA FALL
(Fig. 8)
In Mr. Fall’s original description of F. occulta he listed Los Angeles,
Los Gatos and Mill Valley as localities of collection and sifting litter at
the base of redwoods as the only host data. A number of new distribu-
tional records are known (Fig. 9). In addition to the coast range collec-
344, THE Pan-Paciric ENTOMOLOGIST
Fic. 10. Known geographical distribution of Fuchsina arida and F. occulta.
Dotted line depicts mountain crests with: desert climate to east.
VoL. 52, No. 4, OcToBER 1976 345
Fic. 11. Known geographical distribution of Revelieria californica Fall.
tion, it can be seen that it is also found on the west side of the Sierra
Nevada range and in the southern part of the Klamath Range. Host
data show records from Engelmann oak, Black oak, several undeter-
mined oak species, Pinus ponderosa, Heteromeles arbutifolia, Arcto-
staphylos and from the nest of the wood rat Neotoma fuscipes.
The pattern of distribution for F. arida and F. occulta show allopatry
with all known records of occulta being from the west side of the Sierra
Nevada and transverse ranges, while all records for arida are from the
east side of the Sierra Nevada and transverse ranges (Fig. 10). In San
Bernardino and Riverside Counties the two species are found within
8-10 miles of one another, but always on opposite sides of the moun-
tains.
346 THE Pan-Paciric ENTOMOLOGIST
REVELIERIA CALIFORNICA FALL
(Fig. 9)
This easily recognizable and distinct species was also described by
Fall in 1899. He included it in Revelieria along with R. genei of
Southern Europe and Northern Africa. It was then known from two
Basidiomycetes (Pleurotus and an unidentified polypore) and at Los
Gatos and Ojai, California, both coastal locations.
Subsequent collection, mainly by berlese sampling, has shown it to
be widespread in California, occurring in the lower elevations of the
Coast Ranges, Transverse Ranges, Klamath Mountains, southern end
of the Cascades and on both the east and west sides of the Sierra
Nevada (Fig. 11). It has not yet been found outside of California, but it
seems likely to be found in the Basin Ranges of Nevada and in Southern
Oregon. I have collected it in association with Myxomycete fructifica-
tions on several occasions. It was found in El Dorado Co. on the under
surface of Fuligo septica in association with Odontosphindus clavicornis
Casey (Sphindidae), and has been taken three times at Yuba Pass,
Sierra Co. on Arcyria versicolor Phill. On two occasions adult speci-
mens were collected. The third collection was 3 X 4 inch patch of
sporangia infested with Lathridiid larvae. Rearing yielded several
hundred Enicmus cordatus Belon adults and only three Revelieria cali-
fornica adults. Revelieria larvae were not isolated and studied, but
dissection of the adult gut yielded typical Arcyria versicolor spores. It
has not been recollected in association with higher fungi, but has been
berlesed from the litter of Heteromeles arbutifolia, Juniperus sp., Pinus
monophylla, Pinus sabiana and Quercus sp. It has not been found to be
locally abundant; rarely does a one-half square yard sample contain
more than a single specimen and has never contained more than three
specimens.
ACKNOWLEDGMENTS
I am indebted to the following individuals for help in obtaining
litter samples: R. F. Wilkey, formerly of the California Department of
Agriculture; A. J. Gilbert, T. R. Haig, A. R. Hardy, R. Hobza, E. L.
Paddock and R. E. Somerby of the California Department of Food and
Agriculture; J. T. Doyen and J. D. Pinto of the University of Cali-
fornia, Berkeley and Riverside respectively; and to D. Giuliani of Lone
Pine, California.
The following institutions and individuals made specimens available
for study: H. B. Leech, California Academy of Sciences; J. Chemsak,
VoL. 52, No. 4, OctoBER 1976 347
California Insect Survey, University of California, Berkeley; Saul
Frommer, University of California, Riverside; E. L. Sleeper, Long
Beach State University and J. Schuh, Klamath Falls, Oregon.
Ms. E. Parker prepared the habitus illustrations and the photomicro-
graphs were produced on the auto-scan at the University of California,
Davis, with the assistance of Mr. R. O. Schuster.
LITERATURE CITED
Beton, M. J. 1897. Essai. de classification générale des Lathridiidae avec le
catalogue systématique et alphabétique de toutes les espéces du globe.
Rey. Entomol. Franc. 1897: 105-221.
Dasoz, R. 1970. Etude des coléoptéres Lathridiidae de |’Afrique intertropicale.
Ann. Mus. R. Afr. Centr. Sér. in-8. Zool. 184: 1-49.
Fatt, H. C. 1899. Revision of the Lathridiidae of boreal America. Amer. Ent.
Soc., Trans. 26: 101-190.
Hammapb, S. M. 1953. The immature stages of Metophthalmus serripennis
Broun. R. Ent. Soc. Lond., Proc. (A) 28: 133-138.
Hinton, H. E. 1941. The Lathridiidae of economic importance. Bull. Ent. Res.
32(3): 191-247.
Prez, A. von. 1967. Familie: Lathridiidae, p. 168-182. In Freude, Harde, and
Lohse, eds., Die Kafer Mitteleuropas, Band 7, Clavicornia. Goeke and
Evers, Germany.
SCIENTIFIC NOTE
Distribution and ecology of Leistus ferruginosus Mannerheim (Coleop-
tera: Carabidae) .—Leistus ferruginosus is the only widespread American Leistus.
Its range as given by Erwin includes portions of Alaska, British Columbia, Wash-
ington, Oregon, and possibly California. There is one record from western Alberta.
Most published records are coastal; the following inland localities may be isolated,
relict populations: Jasper, Alberta; Mt. Revelstoke, British Columbia; and Mt.
Bonaparte, Okanagon Co., Washington. The following new record extends the
known range to western Montana: MONTANA: Ravalli Co., 5 miles SW Florence
3700’, June 27, 1967, Loren Russell collector. The single specimen is in the Uni-
versity of Washington collection. This locality is approximately 350 miles south-
east of both Mt. Revelstoke and Mt. Bonaparte.
All of the Oregon localities given by Erwin (1970, Pan-Pac. Entomol., 46:
111-119) are located west of the Willamette Valley, with five of the six strictly
coastal. The following record establishes the presence of L. ferruginosus in the
Oregon Cascade Range: OREGON: Marion Co., Silver Creek Falls State Park,
June 26, 1972, Loren Russell collector.
Erwin (loc cit) saw specimens collected between May and August, with one
teneral specimen taken in June suggesting adult hibernation. I have found this
species active between April 4 (Seattle, Wash.) and September 20 (Whatcom
348 THE Pan-PactFic ENTOMOLOGIST
Co., Wash.). A teneral specimen was taken April 22 (Seattle), so that larval
hibernation may also occur in this species.
Lindroth (1961. Opusc. Entomol., Suppl. 20: 1-200) has described the habitat
of L. ferruginosus as “on moderately moist, half shaded ground usually near
running water.” J have found it in such situations though usually away from
water, but also in the following habitats: at the edge of a snowfield (Whatcom
Co., Wash., Hannegan Pass 3000’, Sept. 30); under large rubble constantly
drenched by spray of a waterfall (Marion Co., Ore., Silver Creek Falls) ; and on
the seabeach near the mouth of a small tidal creek in association with Dyschirius
obesus LeC. and Bembidion tigrinum LeC. (Clatsop Co., Ore.). At the first two
localities cited the Leistus were associated with staphylinids of the genus Phlaeop-
terus. Although the maritime occurrence may have been untypical, it appears
that L. ferruginosus can survive in a variety of microthermal habitats.
An unusual behavioral characteristic which seems to have escaped notice is
the pronounced tendency of L. ferruginosus to ascend moist, shaded vegetation.
I have swept or beaten this species from salmon berry (Rubus spectabilis Pursh)
thickets in Washington and Oregon. Eleven specimens were taken in 20 minutes
sweeping at sunset (Coos Co., Ore.), including one hand-collected approximately
1 meter above the ground. Scansorial behavior is somewhat surprising in the
terrestrial Nebriini, though similar behavior has been noted in the cychrine
Scaphinotus angusticollis Mnh. in coastal forests (Van Dyke, 1944. Entomol. Amer.,
24: 1-9).
Unless stated otherwise, all records are based on material in the author’s col-
lection.—LorEN RusseELi, 828 NW 27th Street, Corvallis, Oregon 97330.
Ecological Notes on Chagas’ Zoonosis in New Mexico.—While recording
the biogeographical distribution of Chagas’ zoonosis and triatome insect vectors
in New Mexico, Wood & Wood (1961, Amer. J. Trop. Med. & Hyg., 10: 155-65)
reported a collection from Chaco Canyon National Monument of 11 Triatoma
protracta protracta (Uhler), which were renamed Triatoma protracta navajoensis
Ryckman (Ryckman, 1962, Univ. Calif. Publ. Entomol., 27: 114-15). Deducting
those specimens named 7’. p. navajoensis, Table 1 of Wood & Wood (1961, loc. cit.)
should now read 445 T. p. protracta collected, 436 examined and 16 or 3.8% infected
instead of 456-442-18-4%. Corresponding data for the 7. p. navajoensis in 1961
should be 11, 6, and 2 or 33% infected. The Navajo conenose bug has been
known for its annoyance to man in Chaco Canyon since 1952 when 1 6 and 1 @
naturally-infected with Trypanosoma cruzi Chagas were received from L. P.
Arnberger (Wood, 1953, Bull. So. Calif. Acad. Sci., 52: 57-60; 56: 51, 99).
The first T. p. navajoensis reported infected with 7. cruzi in Utah was from
a cabin in Wayne County (Ryckman, 1962, loc. cit.).
During the summers of 1962 and 1963, we investigated conenose bug annoyance
to humans at Chaco Canyon, which yielded additional data. Cooperative collecting
by all personnel at the Monument produced 18 T. p. navajoensis in 1962 and 9
in 1963. Single specimens received in 1961 and 1964 raised the total for the
Chaco Canyon personnel living areas to 40 (15 ¢, 22 9, 3 5th instar nymphs)
with 34 (13 ¢, 18 @, 3 5th instar nymphs) examined including 8 (2 ¢,6 9)
or 23.5% infected with Trypanosoma cruzi. Four T. p. navajoensis were taken
from house trailers and 24 from houses. These included infected bugs: 1 ¢,1 9
in bedrooms and 1 @ in bathroom, and uninfected bugs: 1 6, 49, 1 5th instar
VoL. 52, No. 4, OcToBer 1976 349
nymph in bedroom; 2 @ in kitchen and 2 @ in living room. Nine bugs were
captured outside homes. Infected were 1 ¢, 2 @ near a “blacklight” (loaned
by the Los Angeles County Museum of Natural History) and white muslin “bug
sheet” and 1 @ in an empty Sherman live trap set near the canyon wall. Un-
infected were 2 $,1 2 on front porches, 1 ¢ near the blacklight and 1 5th instar
nymph in a live trap. One infected 2 was captured inside the visitor center. No
Triatoma annoyance had been noted in recent years in Chaco Canyon according
to Archaeologist J. E. Mount.
From 1957 through 1960, Superintendent R. Taylor Hoskins and Naturalist
Paul F. Spangle forwarded triatomes to us from Carlsbad Caverns National Park.
Table 1 of Wood & Wood (1961, loc. cit.) included only the 1957 and 1958 speci-
mens of Triatoma gerstaeckeri (Stal). Additional bugs received in 1959 brought
the totals to 47 collected, 43 examined with 5 or 11.6% infected with Trypanosoma
cruzt. Sites of capture, where specified, for T. gerstaeckeri were as follows: in-
side homes (25): infected: 1 ¢@ in living room; 1 ¢ in bathroom; uninfected:
1 92 in living room; 1 ¢,1 92 in bedroom; 1 ¢,1 2 in bathroom; and 1 92 among
stored carpets. Two uninfected males were from the visitor center. Twenty bugs
were captured outside houses: infected: 2 @ at the cave entrance and 1 @ col-
lected by Dr. Gerald Harwood when it flew into the open windows of a parked
car at night; uninfected: 9 ¢, 1 2 on porches, 6 ¢ from outside screens, and
1 $ behind the visitor center. An infected T. gerstaeckeri in a car in the parking
lot suggests a possible dispersal method for this insect vector. High summer
temperatures could be lethal for most specimens in cars, but many cars with air
conditioning could harbor and transport live Triatoma hundreds of miles from
Carlsbad Caverns.
Most Triatoma protracta woodi Usinger were found in Neotoma dens (Table 1,
Wood & Wood, 1961, loc. cit.) ; however, 1 ¢ and 2 2 were collected inside homes
at Carlsbad Caverns. One ¢ was taken on 24.VII.1957 from under a damp cloth
under the kitchen sink and 2 2 on 6.VI.1960 were from beds. One ¢ was found
outside a home on 6.VIII.1958. All were negative for trypanosomes.
Recent correspondence from Environmental Specialist Philip F. Van Cleve at
Carlsbad Caverns reports “no annoyance” from Triatoma since 1968, although a
few specimens were found in areas of the visitor center and residences during the
summer seasons. The reduced incidence of Triatoma seems to be associated with
a reduction of the local small mammal population through improved waste-
disposal techniques.
In southwestern United States, humans may be protected from Chagas’ disease,
because of infrequent contacts with infected Triatoma due to better housing with
screening of doors and windows, exposure to smaller contaminative droplets with
fewer infective parasites per unit volume of feces (Wood, 1960, Exptl. Parasitol.,
10: 356-65), and the rapid evaporation of infected bug feces from the surface
of the skin in hot weather. Shorter feeding times for the insect vectors, delayed
defecation (Wood, 1951, J. Econ. Ent., 44: 52-54) and rapid dispersal from the
host after feeding probably reduce the potential for bite wound contamination.
Human infection in the United States has so far been linked by Woody et al
(1961, J. Pediat., 58: 744) to association with T. gerstaeckeri (28 mm), a larger
species than 7. p. navajoensis (20-23 mm). Triatoma p. navajoensis is the largest
of the widespread North American “protracta complex” of conenose bugs (Ryck-
man, 1962, Univ. Calif. Publ. Entomol., 27: 93-240). The blood meal size of
T. p. navajoensis has been determined by Wood (1976, Ann. Entomol. Soc. Amer.,
350 THE Pan-PaciFic ENTOMOLOGIST
In Press). Subspecies protracta and woodi take much less blood than navajoensis
and, therefore, presumably become infected less often from rodent parasitaemias.
The relatively smaller size of these insect vectors and consequent smaller amount
of ingested blood may assume great importance for the infrequent contaminative
transmission to man of this zoonosis of rodents in the United States.
The yet smaller size of Paratriatoma hirsuta Barber (12 mm) may explain,
in part, why it has not been found naturally-infected despite our examination of
486 specimens. P. hirsuta is readily infected experimentally with Chagas’ trypano-
some by feeding on an infected Peromyscus (Wood, 1941, Pan-Pac. Entomol., 17:
117). One negative ¢ P. hirsuta and 1 positive 5th instar T. rubida wuhlert
nymph were found in the same wood rat den at the Alvarado Mine, near Congress,
Yavapai County, Arizona on 30.V.1940. Twelve P. hirsuta nymphs taken in wood
rat dens in the same location on 2] & 22.XII.1940 were also negative. This was
at a time of winter temperature stress for the rats, a condition which should have
stimulated higher parasitaemias, as noted in Griffith Park, Los Angeles (Wood &
Wood, 1967, Pacific Insects, 9: 545). One negative @ P. hirsuta was collected
16.V.1956 from a bed in a home in Phoenix, Maricopa County, Arizona, where 1
positive 9 T. rubida uhleri was picked up from the cement patio on 12.VI.1970
(Wood, 1975, Pan-Pac. Entomol., 51: 167-8). One negative ¢ P. hirsuta col-
lected 9.VIII.1970 from the service room of a residence in Springdale, Washington
County, Utah, was associated with 2 positive 9 T. p. protracta collected 8 &
23.VII.1970 (Wood, 1973, Ibid, 49: 183-4). Probably heavy parasitaemias of
reservoir mammals are necessary for Paratriatoma to become infected due to its
much smaller blood meals——SHERWIN F. Woop anp Fare D. Woop, 614 PW.
Shenandoah St., Thousand Oaks, CA 91360.
MAILING DATES FOR VOLUME 52
No. 1—January _ mailed 22 April 1976
INOe sage: a at oes mailed 23 July 1976
No. 8--s)aly 5 ee mailed 8 September 1976
No. 4 October mailed 24 March 1977
Vou. 52, No. 4, OcToBER 1976
dol
INDEX TO VOLUME 52
Acanthocinus leechi 203
Adaina ambrosiae 251
Aegialites
californicus 82
fuchsi 82
subopacus 82
Alexander, California Crane Flies 244
Allen and Edmunds, A New Mayfly 133
Alloxysta nothofagi 286
Ammophila
campestris 337
nasalis 335
Andrews, A New Alloxysta 286
Andrews, Lathridiidae, esp. Fuchsina
309
Anomala
diabla 220
eximia 221
hardyorum 221
imperialae 222
kanei 223
sabinae 224
suavis 225
Apatolestes
actites 212
rugosus 311
Apiomerus crassipes 178
Aprostocetus diplosidis 80
Armstrong, Distribution of Stelidota 83
Arnaud, A Parasite of Arctidae 90
Arnaud, Charles H. Martin 90
Arnaud, Sarcophaga intercept 89
Ault, Behavior of Belomicrus 29
Barr and Westcott, Notes on Chryso-
bothris 138
Belomicrus columbianus 29
Bembex
occidentalis 318
rugosa 314
sayi 316
Bicyrtes variegata 318
Bionomics
Adaina ambrosiae 251
Aegialites 82
Ammophila nasalis 335
Anthophora urbana urbana 120
Apatolestes actites 212
Apiomerus crassipes 178
Apoidea 177
Aprostocetus diplosidis 80
Belomicrus columbianus 29
Bembix spp. 314
Chalybion californicum 229
Chrysobothris spp. 138
Coelus ciliatus 338
Conoderus exsul 304
Cyrtopogon inversus 288
Encopognathus pectinatus 331
Fuchsina spp. 339
Heliconus charitonius 291
Hoplitis 110
Hordnia circellata 33
Leistus ferruginosus 347
Panurginus atriceps 161
occidentalis 159
Pleocoma hirticollis vandykei 95
Proteriades shoshone 73
Reinhardiana petiolata 388
Revelieria californica 346
Siphloplecton basale 265
Sphaeridium spp. 14
Synnoma lynosyrana 1
Tabanidae 84
Tachinidae 323
Templemania 91
Tenebrio molitor 84
Veromessor pergandei 63
Bohart and Stange, Liogorytes in Argen-
tina 313
Bohart and Villegas, Behavior of Enco-
pognathus 331
Book Notices and Reviews 62, 158, 233
Bouseman, Biology of Apiomerus 178
Brephos infans oregonesis 83
Caccoplectus
improvisus 156
probus 156
Callophrys (Mitoura)
turkingtoni 60
Cattle droppings 13
Chagas’ disease 348
Chalybion californicum 229
Chandler, New Caccoplectus 154
Chandler, on Attracting Anthicidae 179
Charles H. Martin 90
Chemsak and _ Linsley,
Triacetelus 216
Chemsak and Linsley, Studies of Lep-
turinae 38
Cheng, A New Hermatobates 209
Chontalia 44
cyanicolor 44
Choreutis apocynoglossa 256
Choriolaus 38
derhami 40
gracilis 41
Chrysobothris
axillaris 139
azurea 139
barri 140
breviloba 140
biramosa calida 140
chrysoela 140
cupressicona 141
deleta 144
edwardsii 145
fiskei 145
ignicollis 145
The
Genus
302
inaequalis 146
laricis 146
lata 147
neotexana 147
peninsularis 148
piuta 148
prasina 148
pusilla 149
quadrilineata 149
rossi 150
rotundicollis 150
sexfasciata 151
subcylindrica 151
texana 15]
ulkei 152
woodgatei 152
Clement and Rust, Biology of Hoplitis
110
Clifford, Life Cycle of Siphloplectron
265
Coelus ciliatus 338
Coleoptera
Anthicidae 179
Bruchidae 50
Buprestidae 138, 272
Carabidae 347
Cerambycidae 37, 38, 137, 203, 216,
227
Elateridae 304
Hydrophilidae 14
Lathridiidae 339
Malachiidae 92
Meloidae 179
Nitidulidae 83
Pselaphidae 97, 154
Pyrochroidae 213
Salpingidae 82
Scarabaeidae 14, 95, 220
Staphylinidae 14, 56
Tenebrionidae 84, 338
Collembola 326
Conoderus exsul 304
Compsilura concinnata 90
Computerized Discriminant Functions
Contarinia sorghicola 80
Coville, Spider Wasp Behavior 229
Curicta
howardi 208
pronotata 207
Cyrtopogon inversus 288
Daly, Leucopsis Reared from Xylocopa
271
Deformities—Collops 92
Dennis & Lavigne, Oviposition of Cyrto-
pogon 288
Diedrops 126
aenigma 129
hitchcocki 129
Diptera 18
Asilidae 288
THE Pan-PaciFic ENTOMOLOGIST
Drosophilidae 167
Ephydridae 126, 242
Muscidae 234
Sarcophagidae 89
Tabanidae 84, 311
Tachinidae 90, 323, 338
Tephritidae 90
Tipulidae 244
Doyen, Book Reviews 158, 233
Doyen and Slobodchickoff, Parasitiza-
tion of Coleus 338
Drosophila cuauhtemoci 167
Editorial Notice 285
Encopognathus rufiventris 331
Ephemeroptera 133, 265
Ephydra usingeri 242
Epirrhoe plebeculata 83
Euplecturga 107
fideli 108
impressicollis 107
norstelcha 109
Evans, Bembicini of Baja California 314
Evodinus lanhami 227
Fannia
conspicua 238
thelaziae 235
Felix et al., Mexican Drosophila 167
Fuchsina
arida 339
occulta 343
Garrison, Variation in Libellula 181
Giuliani, Habitat of Aegialites 82
Giulianium 56
campbelli 56
Glenostictia
bifurcata 319
bituberculata 319
gilva 319
Goeden and Ricker, Ragweed Plume
Moth 251
Gregarines 84
Grigarick and Schuster,
Oropodes 97
Gwynne, Nesting Behavior of Ammoph-
ila 335
Halisidota 90
Heliconus charitonius 291
Hemiptera
Gerridae 209, 321
Nepidae 204
Notonectidae 172
Reduviidae 178, 264, 340
Heppner, A New Choreutis 256
Hermatobates singaporensis 209
Hesperoconopa anthracina 245
Heteromurus stannardi 326
Hippomelas 272
caelata 282
carolinensis 273
castaneus 274
compacta 275
Revision of
Vou. 52, No. 4, OcTOBER 1976
cuneata 283
dianae 274
fulgida 275
imperialis 284
insularis 275
juniperinus 275
obliterata 275
pacifica 276
planicauda 279
planicosta 276
sphenica 280
Homoptera
Cicadellidae 33
Hoplitis
hypocrita 111
fulgida 114
sambuci 116
Hordnia circellata 33
Hydriomena nubilofasciata 83
Hymenoptera 19
Andrenidae 159
Anthophoridae 120
Apoidea 177, 271
Chalcidoidea 271
Cynipidae 286
Eulophidae 80
Formicidae 63
Megachilidae 73, 110
Sphecidae 29, 229, 313, 314, 331, 335
Hyrtanella 133
christineae 135
Ischalia chinensis 213
James, Records of Tachinids 323
Johnson, A New Callophrys 60
Johnson, Classification of Kytorhinus 50
Kaplan, Deformities of Collops 92
Kytorhinus prolixus 50
Langston, Geometridae 83
Lee et al., A California Coastal Tabanid
212
Leistus ferruginosus 347
Lepidoptera 233, 262
Arctiidae 90
Geometridae 83
Glyphipterygidae 256
Lycaenidae 50
Nymphalidae 291
Pterophoridae 251
Tortricidae, 91
Leptochoriolaus 47
opacus 47
Lepturges infilatus 137
Leucopsis klugii 271
Lewis, A New Evodinus 227
Lewis, Female Acanthocinus 203
Libellula luctuosa 181
Linsley, Defensive Behavior of Bees 177
Linsley, Galapagos Moths 262
Linsley and Ross, Plant Associations of
Hippomelas 272
Liogorytes joergenseni 313
393
Literature Notices 12, 22, 59, 153, 166,
228, 243, 250, 290, 330, 334
Lycidocerus 44
sanguineus 45
Mannophorus forreri 37
Mari Mutt, A New Heteromurus 326
Marqua, Cerambycid Distribution 37,
137
Mathis and Wirth, New Neotropical
Ephydridae 126
Mayer and Johansen, Biology of An-
thophora 120
Megachoriolaus 41
chemsaki 41
Merritt, Book Notice 62
Merritt, Insects of Cattle Droppings 13
Metretopus 265
Microbembix argyropleura 318
Middlekauff and Lane, A New Apato-
lestes 311
Molophilus spiniapicalis 247
Moore, A New Staphylinid 56
Mordellistenomimus 43
nanus 43
Nebria desolata 91
Nemognathomimus 42
pallidulus 42
Nothofagus 286
Notonecta 172
Odonata
Libellulidae 181
Ormosia
denningi 247
loretta 248
Oropodes 97
arcaps 103
dybasi 101
nuclere 105
orbiceps 99
rumseyensis 100
Page and Fisher, Pleocoma 95
Panurginus
atriceps 161
occidentalis 159
Parker, A new Megachilidae 73
Philip, Galapagos Tabanidae 84
Pleocoma hirticollis vandykei 95
Polhemus, North American Nepidae 204
Polhemus and Cheng, A New Rheu-
matobates 321
Potts, New Anomala 220
Powell, Oviposition of Templemania 91
Powell, Biology Synnoma 1
Proceedings 83
Proteriades shoshone 73
Pseudophilotes 89
Purcell, Host Preference of Hordnia 33
Ranatra
montezuma 204
spatulata 206
texana 206
304
Reinhardiana petiolata 338
Revelieria californica 346
Rhabdomastix brevicellula 244
Rheumatobates ornatus 321
Rissing and Wheeler, Foraging of Vero-
messor 63
Russell, Biology Leistus ferruginosus
347
Sarcophaga tibialis 89
Scientific Notes 37, 80, 82, 83, 84, 89,
90, 91, 92, 95, 137, 177, 178, 179,
203, 212, 262, 264, 271, 313, 323,
338, 347, 348
Scott and Shepard, Computerized Iden-
tifications 23
Siphloplectron 265
Stelidota geminata 83
Steniolia duplicata 318
Stone, Biology of Conoderus 304
Summers, Scientific Note 80
THe Pan-Paciric ENTOMOLOGIST
Synnoma lynosyrana 1
Triacetelus 216
emarginata 218
sericatus 218
viridipennis 217
Triatoma protracta navajoensis 264
Turner, on Fannia Flies 234
Verromessor pergandei 63
Voigt and Garcia, Notonecta nymphs
172
Wirth, A New Brine Fly 242
Wood, Cannibalism in Triatoma 264
Wood and Wood, Chagas’ Zoonosis in
Mexico 348
Xylocopa brasilianorum 271
Young, A New Chinese Pyrochroid 213
Young, Biology of Heliconus 291
Zhovreboff, Gregarines and Tenebrio 84
Zoological Nomenclature 28, 180, 241,
320
Published by the
Pacific Coast Entomological Society
in cooperation with
The California Academy of Sciences
VOLUME FIFTY-TWO
1976
EDITORIAL BOARD
J. T. DOYEN, Editor
J. A. CHEMSAK, Assistant Editor
E. G. LINSLEY
HUGH B. LEECH
E. S. ROSS
H. V. DALY
P. H. ARNAUD, JR., Treasurer
PUBLICATION COMMITTEE
1976 1977 1978
J. G. Edwards J. A. Powell, Chairman T. D. Eichlin
C. B. Philip R. W. Thorp J. D. Pinto
San Francisco, California
1976
i
CONTENTS FOR VOLUME 52
Alexander, C. P. .
Additions to the Crane flies of California 244,
Allen, R. K. and Edmunds, G. F., Jr.
Hyrtanella: A New Genus of Ephemerellidae from Malaysia _______ 133
Andrews, F. G.
A new species of Alloxysta, aphid hyperparasite, from South
PEPTIC eed ps. pun Eee Rt er Ve elses gis hour eke 286
Andrews, F. G.
A new species of Fuchsina with notes on some California
ARPS) s abst ihakel, (co. RERUN REE SLMS Seeing eA eenc ny ee Me 339
Ault, S. K.
Nesting behavior of Belomicrus columbianus _....----- 29
Barr, W. F. and Westcott, R. L.
Notes on Chrysobothris, and new species from California _________ 138
Bohart, R. M. and Stange, L.
Liogorytes joergensent, cicada killer in Argentina _.---_=___ 313
Bohart, R. M. and Villegas, B.
Nesting behavior of Encopognathus rufiventris 331
Book Notices and Recent Literature _.. 12, 22, 59, 62, 153, 166, 228,
233, 243, 250
Boi aewiew Ue ar 28 ee ee, EOE Rh oe 158
Chandler, D. S.
New species of Caccoplectus 154,
Chemsak, J. A. and Linsley, E. G.
Mexican and Central American Lepturine Cerambycidae related
to Euryptera and Choriolaus _... 38
Chemsak, J. A. and Linsley, E. G.
The genus T'riacetelus Bates _.....---- 216
Cheng, L.
A new species of Hermatobates 209
Clement, S. L. and Rust, R. W.
Nesting biology of three species of Hoplitis _.....-----___- 110
iii
Clifford, H. F.
Life cycle of Siphloplecton basale
Coville, R. E.
Predatory behavior of the Spider Wasp, Chalybion californicum 229
Daly, H. V.
Leucopsis klugii (Chalcidoidea) reared from Xylocopa brasili-
anorum (Apoidea) in Costa Rica yar
Dennis, D. S. and Lavigne, R. J.
Oviposition of Cyrtopogon inversus __........-----------------------------------—- 288
Doyen, J. T. and Slobodchikoff, C. N.
Larvae of Coelus ciliatus parasitized by Reinhardiana petiolata _. 338
radi fora TMI fs i bas Ne, SY MSc Ee a Rae 125, 264, 285
Evans, H. E.
Notes on Bembicini of Baja California Sur 314
Felix, R., et al.
Population genetics of Mexican Drosophila. II. A new species
0 PEE GUS CHT MOT OND oct cot og a reek ee ee ET 167
Garrison, R. W.
Multivariate analysis of geographic variation in Libellula
luctuosa 181
Goeden, R. D. and Ricker, D. W.
Life history of Ragweed Plume Moth in Southern California
Grigarick, A. A. and Schuster, R. O.
Revision of the genus Oropodes ____......-.--.----------------------- 07
Gwynne, D. T.
Nesting behavior of Ammophila nasalis 305
Heppner, J. B.
A new Choreutis from the San Francisco Bay Area _...... 256
James, M. T.
Rearing records of Tachinids 323
Johnson, C. D.
Redescription and phylogenetic affinities of Kvytorhinus
PRONE). com, Deve ea) o assay Lee) Be Sond dD eras 2 50
Johnson, K.
A new species of Callophrys from Mexico 60
1v
Lewis, A. E.
A new species of Evodinus from Wyoming 221
Linsley, E. G. and Ross, E. S.
Plant associations among adult Hippomelas _........_-______- 212
Mari Mutt, J. A.
A new species of Heteromurus from the Solomon Islands ________ 326
Mathis, W. N. and Wirth, W. W.
A new Neotropical Shore Fly Genus with two new species _____. 126
Mayer, D. F. and Johansen, C. A.
Biological observations on Anthophora urbana urbana _____-- 120
Merritt, R. W.
Food habits of insects inhabiting cattle droppings in north
eemtral “Galitormia: 08 sh ee eee Se 13
Middlekauff, W. W. and Lane, R. S.
A new species of Apatolestes from California __..__---______ 311
Moore, I.
- Giulianium campbelli, a new genus and species of Marine Beetle
ea AEE, eae Bree eee ee et oe a 56
Parker, G. D.
A new Proteriades reared from trap stems _.....-- 10
Polhemus, J. T.
Notes on North American Nepidae ___....---- 204:
Polhemus, J. T. and Cheng, L.
A new Rheumatobates from Costa Rica __._. B21
Potts, R. W. L.
New species of North American Anomala __...- 220
Powell, J. A.
Host plant preference, mating and egg development in
Synnoma lynosyrana ___.---------- nn 1
Proceedings, Pacific Coast Entomological Society __.._______»_»__ 83
Purcell, A. H.
Seasonal changes in host plant preference of blue-green Sharp-
shooter Hordnia circellata 33
Rrecoist itera yee ©, 2. te Se OE i cel 290, 330, 334
Rissing, S. W. and Wheeler, J.
Foraging responses of Veromessor pergandei to changes in seed
PEER TOC) 6600) 1% i oo Seen Ee eee MCs ae OO. Nees alee ie 63
Russell, L.
Distribution and ecology of Leistus ferruginosus ___-.....--.---- 347
Rust, R. W.
Notes on biology of North American Panurginus —_.-- 159
Scott, J. A. and Shepard, J. H.
Simple and computerized discriminant functions for difficult
identifications: A rapid nonparametric method 25
Stone, M. W.
Biological notes on Conoderus exsul 304,
Turner, W. J.
Fannia thelaziae, new species of eye-frequenting fly from
California and description of F. conspicua female 234,
Voigt, W. G. and Garcia, R.
Keys to Notonecta nymphs of West Coast United States 172
Wirth, W. W.
A new species of Brine Fly from California rice fields 242
Wood, S. F. and Wood, F. D.
Ecological notes on Chagas’ Zoonosis in New Mexico __.......- 348
Young, A. M.
Biology of Heliconius charitonius in Costa Rica 291
Young, D. K.
A. new species of /schalia from Southeastern China 213
Zoological Nomenclature 28, 180, 241, 320
ee
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