The
PAN-PACIFIC
ENTOMOLOGIST
Volume 72 January 1996 Number 1
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PAN-PACIFIC ENTOMOLOGIST
72(1): 1-4, (1996)
PHENOLOGY OF PANDORA MOTH
(LEPIDOPTERA: SATURNITDAE) ADULT EMERGENCE AND
EGG ECLOSION IN CENTRAL OREGON
DARRELL W. Ross
Department of Forest Science, Oregon State University,
Corvallis, Oregon 97331
Abstract.— Pandora moth, Coloradia pandora Blake, adult emergence and egg eclosion were
monitored at three sites in central Oregon. Accumulated degree-days above a threshold of 5° C
from the beginning of adult emergence to the end of adult emergence were between 386.0 + 7.8
and 459.3 + 10.8, and from the beginning of adult emergence to the end of egg eclosion were
between 648.7 + 17.9 and 685.3 + 21.1.
Key Words. —Insecta, Coloradia pandora, phenology, degree-days, adult emergence, egg eclosion
The pandora moth, Coloradia pandora Blake, is a defoliator of ponderosa,
lodgepole, and Jeffrey pines (Pinus ponderosa Dougl. ex Laws., Pinus contorta
Dougl. ex Loud., and Pinus jeffreyi Grev. & Balf., respectively) in the western
United States (Carolin & Knopf 1968, Furniss & Carolin 1977). Most pandora
moths have a two-year life cycle. Adults emerge, mate, and oviposit in mid-
summer. Eggs hatch in late-summer and the young larvae feed in groups on needles
near branch tips. The larvae overwinter on the branches and continue feeding the
following spring. Most defoliation occurs in the spring prior to host shoot elon-
gation. The mature larvae crawl down the bole in early-summer to pupate in the
soil and litter. The pupal stage typically lasts about 12 months, although a small
percentage of the population remains in the pupal stage for two to five years
(Carolin 1971).
The pandora moth is normally inconspicuous, but periodic outbreaks have
severely defoliated pines in parts of Oregon, Colorado, and Arizona (Patterson
1929, Wygant 1941, Schmid & Bennett 1988). Because defoliation occurs in
alternate years and the larvae do not feed on the current years needles, tree
mortality is minimal and usually associated with stressed trees (Wagner & Ma-
thiasen 1985, Bennett et al. 1987). However, defoliation can cause significant
growth loss and may predispose trees to bark beetle infestations (Patterson 1929,
Miller & Wagner 1989). The large number and size of caterpillars and moths and
the dramatic appearance of completely defoliated stands can be a considerable
nuisance to property owners and recreationists within infested areas. The collapse
of outbreak populations after three or four generations has usually been attributed
to a naturally occurring virus.
Knowledge of pandora moth phenology is potentially useful for research and
management applications. Published records of pandora moth phenology are
based upon calendar dates that can vary considerably among generations de-
pending upon weather conditions. This paper reports phenological observations
of pandora moth adult emergence and egg eclosion based upon heat accumulations
from the beginning of moth flight.
2 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
METHODS AND MATERIALS
Pandora moth phenology was monitored at three sites on the Fort Rock Ranger
District of the Deschutes National Forest about 20 km S of Bend, Oregon. The
three sites were within 2.5 km of one another. Plant associations for the study
sites were ponderosa pine/bitterbrush/fescue and ponderosa pine/bitterbrush-
manzanita/fescue (Volland 1985). Elevations were 1400-1440 m, aspects were S
to SW, and slopes were 0-—5%. Ponderosa pine was the only tree species present
on the plots. At the time of the study, tree age (mean + SE) was 62 + 2 years,
diameter at breast height was 18.4 + 2.0 cm, height was 11.7 + 1.9 m, and basal
area was 37.4 + 6.2 m?/ha.
These stands were moderately to heavily defoliated by the pandora moth in
1990 (USDA Forest Service 1990). Between 28 May and 5 Jun 1991, 25 cages
(34 x 47 cm) were staked to the ground at 5 x 5 m spacings at each site to
monitor adult emergence. The cages were inspected on 16, 17, 22, and 30 Jul; 4,
8, 13, 19, and 27 Aug; and 9 Sep 1990. The number and sex of moths present
on each visit were recorded. For each date, adult emergence was calculated as a
percentage of the total seasonal emergence.
Pandora moth eggs and larvae were sampled on 14 and 29 Aug and 10, 17,
and 24 Sep 1991. On each date, 10 randomly selected dominant or codominant
trees were sampled at each site. One 30-45 cm long branch tip was removed with
a pole pruner from the mid-crown on both the N and S sides of each tree. The
branches were taken to the laboratory and eggs and larvae present were counted.
Percent egg hatch was calculated as the number of larvae divided by the number
of larvae plus the number of unhatched eggs.
At the time that emergence cages were installed, an electronic data recorder
with two temperature sensors (Omnidata Datapod® model DP-212, Logan, Utah)
was placed at each site. The temperature sensors were placed in partially shaded
locations within the crowns of understory trees about 1—2 m above the ground.
The instrument scanned temperatures at 5 min intervals and recorded daily max-
imum and minimum temperatures. Degree-days were calculated from the tem-
perature data by the modified sine wave method (Allen 1976) beginning on 17
Jul when the first adults emerged in the plots. A lower threshold of 5° C was
arbitrarily chosen for degree-day calculations because developmental thresholds
for the pandora moth were unknown. However, temperatures below 5° C interrupt
incubation of pupae and eggs of closely related bombycid and saturnid moths
(Rivnay & Sobrio 1967, Wang 1989), and the lower developmental thresholds
for eggs of other North American forest defoliators are near 5° C (Wickman 1976).
RESULTS AND DISCUSSION
The first adult moths were observed in Bend, Oregon on 12 Jul (R. G. Mitchell,
personal communication) and in La Pine, Oregon (about 20 km S of the study
sites) on 16 Jul (unpublished data). Emergence holes in the soil surface and adult
moths were first observed on the plots on 17 Jul, although no adults were captured
in emergence traps until 4 Aug.
The last adult moths were collected from the emergence traps on 27 Aug when
the mean degree-day accumulation (+ SE) was 459.3 + 10.8 (Fig. 1). Those moths
could have emerged on or after 20 Aug, the day following the previous collection,
when the mean degree-day accumulation was 386.0 + 7.8 degree-days. The last
1996 ROSS: PANDORA MOTH PHENOLOGY 3
100
° Adultemergence /
= Egg eclosion /
80
60
40
Percent of total
20
0 100 200 300 400 500 600 700
Degree-days
Figure 1. Percent pandora moth adult emergence and egg eclosion as a function of degree-days
accumulated since the beginning of adult emergence in central Oregon, 1991. Bars indicate standard
errors.
male moths were collected on 19 Aug when the mean degree-day accumulation
was 373.0 + 7.8 degree-days. Those moths could have emerged on or after 14
Aug, when the mean degree-day accumulation was 309.0 + 6.1 degree-days.
Egg eclosion reached 96.2 + 3.8% by 24 Sep when the mean degree-day ac-
cumulation was 685.3 + 21.1 degree-days (Fig. 1). The last eggs could have
hatched on 18 Sep, the day after the previous sample date, when the mean degree-
day accumulation was 648.7 + 17.9 degree-days. The few eggs that remained
unhatched on the last sample date were likely infertile or unhealthy.
Degree-day accumulations can be used to estimate the end of adult emergence,
egg eclosion, or to time control activities. For example, a fall application of a
non-persistent insecticide will be most effective if it is applied soon after all eggs
have hatched (Ragenovich et al. 1986). This date could be determined by re-
peatedly sampling foliage for the presence of eggs and larvae. Alternatively, the
date could be estimated by accumulating degree-days from the date adult emer-
gence begins untila total of 649 is reached. These degree-day accumulations should
4 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
be used only in central Oregon until they are verified in other parts of the geo-
graphic range of the pandora moth.
ACKNOWLEDGMENT
I thank the staff of the Fort Rock Ranger District of the Deschutes National
Forest for providing maps and records, and permitting access to the study sites.
This research was supported, in part, by funds provided by the Research Council,
Oregon State University.
LITERATURE CITED
Allen, J.C. 1976. A modified sine wave method for calculating degree-days. Environ. Entomol., 5:
388-396.
Bennett, D. D., J. M. Schmid, S. A. Mata & C. B. Edminster. 1987. Growth impact of the North
Kaibab pandora moth outbreak. U.S. Dept. Agric, For. Serv. Res. Note, RM-474.
Carolin, V. M.,Jr. 1971. Extended diapause in Coloradia pandora Blake (Lepidoptera: Saturniidae).
Pan-Pac. Entomol., 47: 19-23.
Carolin, V. M., Jr. & J. A. E. Knopf. 1968. The pandora moth. U.S. Dept. Agric., For. Serv. For.
Pest Leaflet, 114.
Furniss, R. L. & V. M. Carolin. 1977. Western forest insects. U.S. Dept. Agric., For. Serv. Miscell.
Publ., 1339.
Miller, K.K. & M. R. Wagner. 1989. Effect of pandora moth (Lepidoptera: Saturniidae) defoliation
on growth of ponderosa pine in Arizona. J. Econ. Entomol., 82: 1682-1686.
Patterson, J. E. 1929. The pandora moth, a periodic pest of western pine forests. U.S. Dept. Agric.,
For. Serv. Tech. Bull., 137.
Ragenovich, I. R., J. M. Schmid, D. D. Bennett, J. W. Barry & C. E. Richmond. 1986. Field
evaluations of four insecticides against the pandora moth, 1982. Insecticide and Acaricide
Tests, 11: 426.
Rivnay, E. & G. Sobrio. 1967. The phenology and diapause of Saturnia pyri Schiff. in temperate
and subtropic climates. Z. Angew. Entomol., 59: 59-63.
Schmid, J. M. & D. D. Bennett. 1988. The North Kaibab pandora moth outbreak, 1978-1984. U.S.
Dept. Agric., For. Serv. Gen. Tech. Rep., RM-153.
USDA Forest Service. 1990. Aerial survey map of the Deschutes National Forest. U.S. Dept. Agric.,
For. Serv., Forest Pest Management, Region 6, Portland, OR.
Volland, L. A. 1985. Plant associations of the central Oregon pumice zone. U.S. Dept. Agric., For.
Serv., R6-ECOL-104-1985. PNW Region, Portland, OR.
Wagner, M. R. & R. L. Mathiasen. 1985. Dwarf mistletoe-pandora moth interaction and its con-
tribution to ponderosa pine mortality in Arizona. Great Basin Naturalist 45: 423-426.
Wang, S. 1989. Silkworm egg production: volume III. Food and Agriculture Organization of the
United Nations, FAO Agricultural Services Bulletin 73/3.
Wickman, B. E. 1976. Phenology of white fir and Douglas-fir tussock moth egg hatch and larval
development in California. Environ. Entomol., 5: 316-322.
Wygant, N. D. 1941. An infestation of the pandora moth, Coloradia pandora Blake, in lodgepole
pine in Colorado. J. Econ. Entomol., 34: 697-702.
PAN-PACIFIC ENTOMOLOGIST
72(1): 5-12, (1996)
DISCOVERY OF COLPOCLYPEUS FLORUS (WALKER)
(HYMENOPTERA: EULOPHIDAE) IN APPLE
ORCHARDS OF WASHINGTON
JAY F. BRUNNER
Tree Fruit Research and Extension Center, Washington State University,
1100 North Western Avenue, Wenatchee, Washington 98801
Abstract. —Routine collections of leafroller larvae in Washington apple orchards for the purposes
of assessing the kinds of parasites and levels of parasitism revealed the presence of an unknown
species. Specimens sent to the USDA-ARS Systematic Entomology Laboratory, Beltsville, were
identified as Colpoclypeus florus (Walker). This report represents an extension of its geographic
range, the first record in the U.S., as well as a new host record. Colpoclypeus florus is a gregarious
ectoparasitic eulophid which is widely distributed in Europe where it is the most common natural
enemy of tortricids in orchards and other crops. It was introduced into Canada and released in
small numbers in 1966 and 1967. No recovery was reported until 1992 when it was reared from
larvae of obliquebanded leafroller and eyespotted bud moth in Ontario. In that same year the
level of parasitism by C. florus in two unsprayed apple orchards in Washington was 66% and
75%. The parasite’s behavior and key characters that indicate its possible presence in an orchard
are discussed. Notes on the distribution in Washington and methods of rearing C. florus on
pandemis leafroller larvae are described, along with its potential as a biological control agent in
Washington.
Key Words.—Insecta, Colpoclypeus florus, Eulophidae, leafroller parasite, new host record
Two species of leafroller (Lepidoptera: Tortricidae), Pandemis pyrusana Kear-
fott, the pandemis leafroller (PLR), and Choristoneura rosaceana (Harris), the
obliquebanded leafroller (OBLR), are the most common pests of pome fruit in
Washington State (Beers et al. 1993; Brunner 1984, 1991). Both species overwinter
as a second or third instar larva and complete two generations per year (Brunner
& Beers 1990). Insecticides are the primary tactics used to control these pests
(Brunner 1988). Natural enemies have been reported attacking leafrollers in the
western United States. In Washington, however, the natural suppression of leaf-
rollers has generally not been sufficient to provide commercially acceptable con-
trol. This is probably due to the use of broad-spectrum insecticides used to control
codling moth, Cydia pomonella (L.), that are toxic natural enemies of leafrollers.
This is the first report of the parasite, Colpoclypeus florus (Walker), attacking
PLR in Washington apple orchards. It represents an extension of the geographic
distribution of C. florus, as well as a new host record.
MATERIALS AND METHODS
Third through fifth instar PLR and OBLR were collected from several apple
orchards near Wenatchee, Washington, and Milton-Freewater, Oregon, in the
spring of 1992 (Table 1). Potential leafroller feeding sites, one or more leaves or
flower parts held together with webbing, were collected, placed in plastic food
storage bags and returned to the laboratory. Feeding sites were examined and
healthy larvae were placed individually in one-ounce plastic portion cups (Prairie
Packaging, S-100), along with a cube of modified pinto bean diet (Shorey & Hale
1965). If parasite larvae or pupae were found, they were placed into a cup along
with the remains of the host and leaf. Cups were kept at a constant temperature
6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
(23° C) and a long photoperiod (16:8, light : dark) until larvae died or adult leaf-
rollers or parasites emerged.
In late July PLR larvae of a second summer generation were collected from
orchards in Wenatchee (Table 1). Some PLR larvae were attacked by an unknown
ectoparasite. The webbing of the rolled leaf shelter of larvae attacked by this
parasite was uncharacteristically tough. When such sites were found, the entire
leaf, or the portion containing the parasitized leafroller larva, was placed in a
small tight-sealing plastic petri dish (Falcon 1006—50 x 9 mm) which was then
placed in a food storage container. When an apparently healthy larva was found,
it was placed in a one-ounce plastic portion cup with a cube of the modified pinto
bean diet. All cups were held in the conditions described above and were examined
two times per week until a parasite or an adult leafroller emerged. Any parasites
emerging from leafroller larvae or pupae were placed in glass vials with alcohol
and labeled as to host-collection site and date. Adults of the ectoparasitic eulophid
not previously reared from PLR were sent to the USDA-ARS Systematic Ento-
mology Laboratory, Beltsville, for identification.
In late August, several unsprayed orchards were sampled for the density of
leafroller larvae. Twenty shoots, on 40 to 60 trees, were examined for presence
_ of larval feeding sites. Feeding sites were opened and a record was kept of those
which were parasitized by the eulophid.
Adults of the ectoparasite which emerged from field-collected PLR larvae were
counted and sexed. One male and one female were placed inside a one-ounce cup
along with a fourth instar PLR larva. Each cup contained a cube of bean diet as
a food source for the leafroller, and honey was streaked on the top or side of the
cup as a carbohydrate source for the parasites. Cups were placed inside a food
storage container and kept at constant temperature 23.9° C and long photoperiod
(16:8, light:dark). Cups were examined three times per week to determine the
status of leafroller and parasites. The number of leafrollers which emerged as
adults and the number of parasites emerging from each leafroller larva were
counted and sexed. The size of each life stage of C. florus was determined by
measuring the length and width of 30 individuals of each stage, randomly selected
from the parasite colony maintained in the laboratory, using a ocular micrometer
and dissecting microscope.
RESULTS AND DISCUSSION
The ectoparasitic eulophid reared from PLR larvae was identified as C. florus
(M. E. Schauff, Systematic Entomology laboratory, USDA, Beltsville). The dis-
covery of C. florus attacking larvae of PLR in Washington represents the first
record of the species in the United States, as wellas a new host record. Colpoclypeus
florus was transported to Ontario, Canada, from France and Italy and released
against the strawberry leafroller, Ancylis comptana fragariae (Walsh & Riley), in
1966 and the redbanded leafroller, Argyrotaenia velutinana (Walker), in 1967.
However, no record of its recovery was ever noted (Williamson 1966, Hikichi
1971). Hagley & Barber (1991) reported rearing C. florus from OBLR and Spi-
lonota ocellana (Dennis & Schiffermiieller), the eyespotted bud moth. This report
represented the first official recovery of C. florus in Canada.
Colpoclypeus florus is a gregarious ectoparasite of tortricids and is often the
most common and important natural control in European orchards and other
1996 BRUNNER: COLPOCLYPEUS FLORUS IN WASHINGTON 7
Table 1. Site descriptions where leafroller larvae were collected for determining parasitism levels.
Site Location Crop Management type
TFREC Wenatchee apple unsprayed
Birchmont Wenatchee apple soft pesticide/unsprayed
Marley Yakima apple conventional
WSU Pullman apple feral trees
Davis Milton-Freewater apple conventional/non-bearing
BF Milton-Freewater apple semi-abandoned
Baneck Milton-Freewater apple conventional
Wondra Milton-Freewater apple abandoned
crops (Evenhuis 1974a, Evenhuis & Vlug 1983, Gruys & Vaal 1984, Limon &
Blasco 1973, Monta & Gambaro 1973, Scaramozzino & Ugolino 1978, van Veen
and & van Wijk 1987, Vives 1980). Colpoclypeus florus parasitizes larvae of several
tortricid species in Dutch apple orchards and survives control programs based
on insect growth regulators (Gruys & Vaal 1984, de Reede et al. 1984, Helsen &
Blommers 1989). In Germany C. florus, along with Meteorus ictericus (Nees) and
Apanteles ater (Ratzburg), parasitized up to 80% of Adoxophyes orana (Fisher
von Réslerstamm) in apple orchards treated with the insect growth regulator
fenoxycarb (Harzer 1990). In Italian orchards, C. florus was responsible for 95%
parasitism of Argyrotaenia pulchellana Haworth (Monta & Gambaro 1973).
Colpoclypeus florus is not well synchronized with most tortricid species found
in European apple orchards. It is most common in mid- to late summer when it
parasitizes a high percentage of tortricid larvae (Gruys & Vaal 1984). However,
it is present in low levels in the spring and does not appear to overwinter on A.
orana, 1ts most common host in apple orchards (Evenhuis 1974b). European
orchard fruit tortricid pests overwinter as young larvae or eggs (Alford 1984),
unsuitable host stages for C. florus. Colpoclypeus florus probably uses overwinter
hosts that occur outside the orchard, such as Syndemis musculana (Hiibner) which
overwinters as full-grown larvae and may provide an important overwintering
reservoir for C. florus (Blommers et al. 1988). The most important leafroller pests
of pome fruit in Washington overwinter as eggs or young instar larvae (Beers et
al. 1993). As in Europe, a lack of suitable overwintering hosts may represent a
barrier to the survival and consistency of C. florus populations in the Pacific
Northwest.
How C. florus was transported to the western U.S. and why it was not detected
until 1992 are unknown. Colpoclypeus florus was probably established in British
Columbia by the mid-1970s, based on the inclusion of Colpoclypeus sp. in a list
of parasites reared from leafroller larvae as part of a research project under the
direction of Dr. H. F. Madsen (Jerry Vakenti, pers. comm.). Unfortunately this
information has not been published. Because the genus Colpoclypeus was not
known from North America until it was introduced into Canada (Williamson
1966, Hikichi 1971), it is likely that this unpublished report represents the first
indication of the presence of C. florus in western North America.
Leafroller Parasitism. —Low levels of leafroller parasitism, 0 to 13%, were found
in all orchards in spring (Table 2) and no C. florus were detected. Summer par-
asitism levels were higher, especially in two unsprayed apple blocks, TF-5 and
8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Table 2. Leafroller larvae collected from different sites in 1992 and the results of rearing to de-
termine levels of parasitism.
; : Number of parasites reared
Locations: Spring 1992 LLR/OBLR: ——————————————— eS .. 1% total.
Site ID # larvae adults C. florus Apanteles_ Tachinidae Other parasitism
PLR sites
WSU-TFREC-21 86 75 0 0 0 0 0.0
WSU-TFREC-5 100 74 0 0 0 1 1.3
WSU-TFREC-24 87 75 0 0 0 0 0.0
Birchmont-4F east 92 89 0 1 0 0 1.1
Birchmont-4F west 51 47 0 0 0 0 0.0
Birchmont-1A 73 64 0 1 0 0 1.5
Marley 63 48 0 0 0 4 Tish
OBLR sites
WSU-Pullman 100 72 0 0 0 0 0.0
Davis 100 84 0 0 0 0 0.0
BF 31 22 0 0 0 3 12.0
Baneck? 100 41 0 0 0 0 0.0
Wondra 37 17 0 0 0 2 10.5
. Number of parasites reared
Locations: Summer 1992 PLR a SS SSS A 8 ttl,
Sites ID # larvae adults C. florus Apanteles Tachinidae Other parasitism
PLR sites
WSU-TFREC-5 52 11 38 0 0 2 78.4
WSU-TFREC-24a 59 9 40 0 2 0 82.3
WSU-TFREC-24b 109 32 67 0 2 0 68.3
Birchmont-4F west 78 63 0 0 12 0 16.0
Birchmont-1D1 81 46 3 4 11 l 29:2
Birchmont-1D2 71 42 6 6 12 1 37.3
4 Low survival level of leafroller larvae probably due to use of Bt products in the orchard prior to
larval collections.
TF-24, located at the Washington State University Tree Fruit Research and Ex-
tension Center, Wenatchee (WSU-TFREC). In these blocks, C. florus was re-
sponsible for 75% and 78% of the total leafroller parasitism (Table 2). Two col-
lections of PLR larvae were made in block TF-24 three days apart. The first
indication that a different parasite was present in the WSU-TFREC blocks was
the discovery of abnormally dense and much tougher webbing than is typically
formed by PLR larvae. The webbing made by parasitized leafroller larvae was
similar in density to spider webbing, though not of the same consistency.
An average of 13.5 C. florus adults was reared from parasitized PLR larvae
(Table 3). The sex ratio favored females, 71.8%, as found in Europe (Evenhuis
1974b, Gruys & Vaal 1984, van Veen & van Wijk 1987). Colpoclypeus florus was
also reared on PLR larvae originating from a laboratory colony reared on artificial
diet. The average number of C. florus progeny produced from laboratory-reared
PLR larvae and their sex ratio were no different from field collected, parasitized
PLR larvae (Table 3). Colpoclypeus florus is easy to rear on PLR or OBLR larvae
in the laboratory, and thus it may be a promising agent for mass rearing and
augmentative release for biological control of leafroller pests in Washington or-
chards.
1996 BRUNNER: COLPOCLYPEUS FLORUS IN WASHINGTON 9
Table 3. Data on C. florus reared from PLR larvae collected from two unsprayed apple orchards
in Wenatchee and produced in the laboratory using PLR larvae from a colony.
Colpoclypeus florus
No. of Avg. no. progeny
Year Source leafroller larvae % male % female (adults + SE)
1992 Field 85 28.2 71.8 13.5 + 1.02
1992/3 Lab (PLR) 53 27.0 73.0 Qe lect le 27:
Distribution. — Colpoclypeus florus was detected parasitizing PLR larvae at low
levels in two commercial orchards, Birchmont-1D1 and 1D2, located 5 km north
of the WSU-TFREC orchards. These orchards were treated with conventional
insecticides in the pre-bloom period but did not receive additional insecticide
treatments during the summer. Colpoclypeus florus was also found in an organic
orchard in East Wenatchee, 10 km east of the WSU-TFREC location, and in an
unsprayed apple orchard 16 km north of Wenatchee along the Columbia River.
The abundance of C. florus at these sites was very low; only one or two parasitized
PLR larvae were found, but the finds indicate a rather widespread distribution
of C. florus in north-central Washington.
Additionally, C. florus was observed by an agricultural consultant (Steve Harris,
pers. comm.) in an organic orchard near Brewster, Washington, 100 km north of
Wenatchee. Leaves containing pupae from which C. florus had already emerged
were submitted to the WSU-TFREC for identification. Although this report does
not confirm C. florus presence in the Brewster orchard, no other ectoparasitic
eulophids attack leafrollers in Washington. The OBLR population in the Brewster
organic orchard had been very high in the spring, and control measures were
planned against the summer generation. However, when searches of growing
shoots in July and August revealed high levels of parasitism the grower decided
not to apply control sprays.
Parasite Description.—The adult parasite has a shiny black head and thorax
(sometimes appearing greenish under reflected light) with a lighter-colored ab-
domen (Fig. 1A). The average length (head to tip of abdomen) of the female is
1.82 + 0.01 mm (mean + SEM). The ventral side of the abdomen 1s a light cream
to golden-brown color. There are two small dark spots in the center and about
one-third of the way in from the tip of the abdomen that denote the base of the
Ovipositor (Fig. 1C). The legs are golden brown. The male C. florus is slightly
smaller than the female, 1.46 + 0.02 mm, and has a darker and less ovate
abdomen. Eggs are translucent when first laid but later turn a creamy white. They
are elongate, slightly curved and average 0.32 + 0.003 mm in length and 0.08 +
0.001 mm in width (Fig. 1B). Neonate larvae migrate to the host larva and begin
feeding. Colpoclypeus florus larvae are a bright green when young, turning to a
pale green or cream color as they near maturity (Fig. 1D). Mature larvae average
1.90 + 0.04 mm in length and 0.67 + 0.01 mm in width. When mature, C. florus
larvae leave the host larva but remain within the host’s webbed shelter to pupate.
Pupae are naked and a light brown color which soon darkens. Female pupae are
slightly more ovate than males, averaging 1.94 + 0.03 mm in length and 0.78 +
0.01 mm in width (Fig. 1E). Male pupae average 1.60 + 0.02 mm in length and
0.62 + 0.01 mm in width.
Figure 1.
,
Life stages of Colpoclypeus florus (Walker); A—adult female, B—eggs, C—female abdomen ventral view, D—larvae feeding on leafroller, E—pupae.
Ol
LSIDOTOWOLNA OMIOVd-NVd AHL
(ZL IOA
1996 BRUNNER: COLPOCLYPEUS FLORUS IN WASHINGTON 11
Parasite Behavior.—My observations confirm that C. florus does not paralyze
its host (van Veen & van Wijk 1987). Males typically emerge first and mate with
females as they emerge. In my laboratory, adults usually remained within the
host’s webbed shelter for two to three days until all had emerged, and then they
all departed over a short period. I observed female C. florus make rapid thrusting
movements with the ovipositor in the direction of the head capsule of a leafroller
larva. However, I have never observed the ovipositor penetrate the caterpillar’s
head capsule. After the attack there was significant change in the behavior of the
leafroller. It stopped feeding, began spinning a dense webbed shelter, and often
voided fluids. The C. florus female remained with the leafroller larva within the
densely webbed shelter and deposited eggs.
The presence of C. florus in Washington provides another natural enemy to
combat leafrollers. It has been reared on both PLR and OBLR in my laboratory
and may be a parasite that could be mass reared for timely releases. The unusually
high levels of leafroller parasitism noted in unsprayed orchards suggests that C.
florus is well adapted to the eastern Washington climate. The susceptibility of C.
florus to conventional insecticides in Europe suggests that it may be most useful
as a biological control of leafrollers in fruit orchards using alternative controls for
key pests, such as insect growth regulators, pheromones or bacterial insecticides.
ACKNOWLEDGMENT
I thank M. E. Schauff, Systematic Entomology laboratory, USDA, Beltsville for
the rapid identification of specimens submitted and Drs. Bill Turner, Elizabeth
Beers and Rich Zack for critical reviews and suggestions which helped improve
the manuscript. I am especially grateful to the able assistance of Mike Doerr and
Lisa Mill for their technical skills in rearing leafrollers and parasites in my lab-
oratory. I acknowledge the invaluable contribution of Washington’s fruit growers
who partially funded this research through the Tree Fruit Research Commission.
LITERATURE CITED
Alford, D.C. 1984. A colour atlas of fruit pests their recognition, biology and control. Wolfe House
Ltd.
Beers, E. H., J. F. Brunner, M. Willett & G. Warner. 1993. Orchard pest management: a resource
book for the Pacific Northwest. Good Fruit Grower, Yakima, Washington.
Brunner, J. F. 1984. Pandemis leafroller, a pest of increasing concern to Washington apple growers.
Proc. Wash. State Hort. Assoc., 79: 119-125.
Brunner, J. F. 1988. Apple pandemis in Washington apple orchards. Down to Earth, 44: 5-9.
Brunner, J. F. 1991. Leafroller pests of fruit crops in Washington state. pp. 185-197. Jn Williams,
K. (ed.). New directions in tree fruit pest management. Good Fruit Grower, Yakima, Wash-
ington.
Brunner, J. F. & E. H. Beers. 1990. Apple pandemis and obliquebanded leafroller. Cooperative
Extension, Washington State University EB1582.
Blommers, L., H. Helsen & F. Vaal. 1988. The autumn leafroller: phenology, damage and parasites
in a Dutch apple orchard. Netherlands J. of Plant Path., 94: 95-103.
de Reede, R. H., R. F. Groendijk & A. K.H. Wit. 1984. Field tests with the insect growth regulators,
epofenonane and fenoxycarb, in apple orchards against leafrollers and side-effects on some
leafroller parasites. Entomol. Exp. Appl., 35: 275-281.
Evenhuis, H. H. 1974a. The enemies of apple tortricids. Les organismes auxiliaires en verger de
pommiers. OILB/SROP, Brochuer, 3: 29-30.
12 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Evenhuis, H. H. 1974b. Colpoclypeus florus (Hymenoptera, Eulophidae), an important potential
parasite of Adoxophyes orana (Lepidoptera, Tortricidae) in apple orchards. Mededelinger van
de Faculteit van de Landbouwwetenschappen, Rijksuniversiteit Gent, 39: 769-775.
Evenhuis, H. H. & H. J. Vlug. 1983. The hymenopterous parasites of leaf-feeding apple tortricids
(Lepidoptera, Tortricidae) in The Netherlands. Tijdschr. Entomol., 126: 109-135.
Gruys, P. & F. Vaal. 1984. Colpoclypeus florus, a eulophid parasite of tortricids in orchards: rearing,
biology and use in biological control. Entomol. Exp. Appl., 36: 31-35.
Hagley, E. A. C. & D. R. Barber. 1991. Foliage-feeding lepidoptera and their parasites recovered
from unmanaged apple orchards in southern Ontario. Proc. Entomol. Soc. Ontario, 122: 1-7.
Harzer, W. 1990. Preliminary findings on the control of the fruit peel tortricid with Insegar. Obstbau
(Bonn), 15: 146-150.
Helsen, H. & L. Blommers. 1989. On the natural control of the summer fruit tortrix in a mildly
sprayed apple orchard. Mededelingen van de Faculteit Landbouwwetenschappen, Rijksuni-
versiteit Gent., 54: 905-909.
Hikichi, A. 1971. Argyrotaenia velutinana (Walker), red-banded leaf roller (Lepidoptera: Tortricidae).
In Kelleher, J. S. (ed.). Biological control programmes against insects and weeds in Canada
1959-1968. Commonwealth Institute of Biol. Control. Tech. Comm. No. 4, 10-11.
Limon, de la Oliva F. & P. J. Blasco. 1973. Measures to use against pests of Citrus in the northern
part of the Levante region, with a view to the establishment of a programme of integrated
control. Boletin Informativo de Plagas, 109: 69-86.
Monta, L. P. dalla & P. I. Gambaro. 1973. Observations on the biology and ethology of Colpoclypeus
florus Wik. (Hymenoptera, Chalcidoidea, Eulophidae) in the laboratory and in the open field.
Redia, 54: 243-259.
Scaramozzino, P. L. & A. Ugolino. 1978. Pandemis heparana (Den. & Schiff.), a tortricid injurious
to peach in Piedmont. Informatore Fitopatologico, 29: 3-6.
Shorey, H. & R. L. Hale. 1965. Mass-rearing of the larvae of nine noctuid species on a simple
artificial medium. J. Econ. Entomol., 58: 522-524.
van Veen, J.C.& M.L. E.van Wijk. 1987. Parasitization strategy in the non-paralyzingectoparasitoid
Col poclypeus florus (Hym., Eulophidae) towards its common summer host Adoxophyes orana
(Lep., Tortricidae). 1. Host finding, acceptance and utilization. J. Appl. Entomol., 104: 402-
417.
Vives, J. M. 1980. An important pest of Spanish carnations, the South African carnation miner,
Epichoristodes acerbella Walk. Agricultura, Spain, 49 (580): 688-691.
Williamson, G. D. 1966. Summary of parasite and predator liberations in Canada and of insect
shipments from Canada in 1966. Canada Dept. of Agric. Report, Research Institute—Belleville,
Ontario.
PAN-PACIFIC ENTOMOLOGIST
72(1): 13-16, (1996)
APHIDIUS UZBEKISTANICUS
(HYMENOPTERA: APHIDITDAE) ESTABLISHED IN IDAHO
SUSAN E. HALBERT,!? JAMES B. JOHNSON,? PEGGy L. GRAVES,!
PAUL M. MARSH* AND DEBORAH NELSON?
1Aberdeen Research and Extension Center, University of Idaho,
Post Office Box AA, Aberdeen, Idaho, 83210
3Department of Plant, Soil and Entomological Sciences, University of Idaho,
Moscow, Idaho 83844-2339
4Post Office Box 384, North Newton, Kansas 67117;
>United States Department of Agriculture,
Animal and Plant Health Inspection Service, Plant Protection and Quarantine,
Niles Biological Control Laboratory, 2534 South Eleventh Street,
Niles, Michigan 49120
Abstract.—Exotic parasites of Diuraphis noxia (Mordvilko) (Homoptera: Aphididae) were re-
leased in Canyon Co. Idaho in 1988. A survey in 1994 indicated that Aphidius uzbekistanicus
Luzhetzki had become established; however, the parasites were found attacking two other cereal
aphids, Sitobion avenae (Fabricius) and Schizaphis graminum (Rondani), but not D. noxia. The
establishment was achieved by releasing wasps into small, shaded cages for 2-5 days.
Key Words.—Insecta, Sitobion avenae, Diuraphis noxia, Schizaphis graminum, Aphididae, bi-
ological control, Aphidius uzbekistanicus, Aphidiidae
Diuraphis noxia (Mordvilko) (Russian wheat aphid) is a recently introduced
pest that has caused considerable damage to small grains in the western United
States (Webster & Amosson 1994). It is under good natural control in its native
range (SEH, unpublished observation), suggesting that classical biological control
could minimize the impact of D. noxia, as well as reducing the need for pesticides.
A national effort was initiated under the auspices of the Western Regional Co-
ordinating Committee— 66 (WRCC— 66) to procure and establish exotic natural
enemies of D. noxia. This paper documents the release and establishment in Idaho
of one of the natural enemies (Aphidius uzbekistanicus Luzhetzki) obtained through
WRCC— 66.
MATERIALS AND METHODS
Origin of Released Natural Enemies. —The strain of A. uzbekistanicus released
in Parma (Canyon County) Idaho came from Turkey. It was collected by Tadeusz
Poprawski and Francis Gruber in the spring of 1988. The culture went through
quarantine at the European Parasite Laboratory near Paris, France (USDA-ARS)
and at Texas A&M University, College Station, TX. In August, 1988, a shipment
of this culture was received for propagation and release in Idaho.
Propagation of A. uzbekistanicus in Idaho.—The culture of A. uzbekistanicus
was maintained on D. noxia, which in turn, were raised on spring wheat, cv.
"IDO-232,” courtesy of the Idaho wheat breeding program. Wheat plants were
2 Present Address: Division of Plant Industry, Florida Department of Agriculture and Consumer
Services, Post Office Box 147100, Gainesville, Florida 32614-7100.
14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
seeded at a rate of about eight per 15 cm diameter pot. They were at least 6 wk
old (but not heading) when infested with D. noxia. Cultures were kept in 41 x
46 cm cages that had slanted glass tops installed about 40 cm above the floors.
Cultures were maintained by transferring 50-100 parasites into a cage with fresh
aphids rather than by adding aphid-infested plants to existing cultures. Parasites
were harvested in groups of about 100 for release.
Release Procedure. —Parasites were released throughout the late summer and
autumn of 1988 in cages about 1 m long, 2 m wide and 4 m high covered with
white chiffon material. A-frame tents of cotton percale cloth were used to shade
the cages in hot weather. Releases were made in the late afternoon or evening
into naturally occurring D. noxia infestations in mixed perennial grasses inter-
seeded with winter wheat and into volunteer wheat in the fall. Release locations
were 1-5 km north of Parma, Idaho. Cages and tents remained in place 2-5 d,
until they were needed for more releases.
Recovery Surveys.—An attempt was made to recover A. uzbekistanicus in the
same fields where it had been released by collecting mummified aphids and al-
lowing the parasites to emerge in a 95% humidity chamber; however, we were
unable to recover any A. uzbekistanicus in 1988. Limited collections in 1989-
1991 also failed to find any A. uzbekistanicus.
A more extensive survey was conducted at the University of Idaho Parma
Research & Extension Center during the 1994 growing season. Spring wheat, cv.
’Penawawa’ was planted on 15 Mar, 12 Apr and 10 May 1994. The last planting
was planted extremely late for the Parma area in order to ensure high populations
of aphids. Each planting occupied about '4 of the entire 0.4 ha. field.
Sampling began in the first two plantings on 11 May, and samples were taken
in all plantings every two weeks thereafter until the wheat matured past the point
where it would support aphid populations on 18 July. Fall samples were taken in
volunteer wheat at the Parma Research & Extension Center on 15 October. Mum-
mies were collected directly from the field and isolated by aphid species in 4
dram shell vials with cotton stoppers. Randomly selected tillers (n = 150-600)
(Feng et al. 1992) were inspected biweekly from each planting date until wheat
matured. Additionally, living aphids were collected each sampling period and
established singly on wheat leaf sections in 50 mm petri dishes. On each sampling
period, 100 D. noxia and 50 of each other species present in sufficient quantities
were collected (Feng et al. 1992).
RESULTS AND DISCUSSION
Parasites Reared from Mummies on Spring Wheat.—A total of 327 mummies
were collected from spring wheat during the sampling season. Rearing 25 D. noxia
mummies yielded 14 Diaeretiella rapae (M’Intosh) and 10 hyperparasites (Table
1). The most abundant aphid species for which mummies were collected was
Sitobion avenae (Fabr.). The 167 S. avenae mummies yielded 35 A. uzbekistanicus,
15 Aphidius ervi Haliday, four Aphidius sp., one D. rapae and 112 hyperparasites.
Samples of 12 Schizaphis graminum (Rondani) and 13 Metopolophium dirhodum
(Walker) yielded two and zero A. uzbekistanicus, respectively.
Parasites Reared from Living Aphids Established in Petri Dishes.—Thirty D.
noxia mummies were reared from which parasites eventually emerged (Table 2).
An additional 55 parasites were obtained from S. avenae. Diaeretiella rapae was
Table 1. Parasites collected from cereal aphid mummies on spring wheat, Parma, ID, 1994.
Primary parasites Hyperparasites
~ Aphidius~SAphidius’=<‘zssSA@*zihidius”=~=<CO*é‘éCé~Scereticiilas=—=—=—“‘éa ‘“OéCOC*é*é‘(C;C™SW Alloxysta Dendrocerus
Aphid host uzbekistanicus ervi spp. rapae Unknown spp. Pteromalidae spp. Total
Diuraphis noxia 0 0 1 14 0 2 8 0 25
Sitobion avenae 35 15 4 1 0 0 109 3 167
Metopolophium dirhodum 0 4 1 0 0 0 8 0 13
Schizaphis graminum 2 2 3 1 1 0 3 0 12
Unknown host? 21 53 9 24 0 1 , 0 110
Total 58 74 18 40 1 3 130 eS 327
9661
a Parasites emerged during field collection and transport to the laboratory, thus they could not be isolated by host.
OHVdI NI SQOINVISTYAIZA SAICIHdY “IV Lo LYUAGIVH
ST
16 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Table 2. Parasites that emerged from live aphids collected on spring wheat, Parma, ID, 1994.
Primary parasites Hyperparasites
Aphidius Aphidius Aphidius Diaeretiella Pteromalidae
Aphid host uzbekistanicus ervi spp. rapae spp. Total
Diuraphis noxia 0 2 0 27 1 30
Sitobion avenae 11 18 5 1 20 55
by far the most common parasite of D. noxia, while Aphidius spp. were more
common parasites of S. avenae. Hyperparasites were more common in S. avenae
mummies than in D. noxia mummies.
Eleven A. uzbekistanicus were recovered from S. avenae reared in petri dishes.
Aphidius uzbekistanicus accounted for 21 and 20% of total parasitism of S. avenae
in field collections and rearing of live specimens, respectively. The percentage
contributed by A. ervi was higher among live collected S. avenae than among
parasites reared from field collected S. avenae mummies (33% and 9% respec-
tively). The reverse was true for hyperparasites (36% for live collected; 65% for
mummies), suggesting that hyperparasites may prefer A. ervi mummies over 4A.
uzbekistanicus mummies; however, we could not associate specific hyperparasites
with A. uzbekistanicus because all Aphidius mummies are similar.
Our results are similar to the extensive pre-release baseline surveys of Feng et
al. (1992) in that D. rapae was more commonly found in D. noxia mummies than
in mummies of other aphid species. Hyperparasites were also much more common
in S. avenae mummies in both surveys. The collections of A. uzbekistanicus
represent a recovery of parasites released in 1988; however, so far, none have
been found parasitizing D. noxia.
ACKNOWLEDGMENT
We thank June Thomas, Debra Stansell and Irene Shackelford for field and
laboratory assistance. We are grateful to J. P. McCaffrey, E. J. Bechinski, and two
anonymous reviewers for their helpful comments. The work was made possible
through USDA-APHIS cooperative agreement Number 94-8 100-0254 (CA) and
support from the Idaho Wheat Commission. This is Idaho Agricultural Experi-
ment Station Scientific Paper No. 95723.
LITERATURE CITED
Feng, M.-g., J. B. Johnson & S. E. Halbert. 1992. Parasitoids (Hymenoptera: Aphidiidae and Aphel-
inidae) and their effect on aphid (Homoptera: Aphididae) populations in Irrigated Grain in
Southwestern Idaho. Environ. Entomol., 21: 1433-1440.
Webster, J. A. & S. Amosson. 1994. Economic impact of the Russian wheat aphid in the western United
States: 1992-1993. Great Plains Agricultural Council Publication No. 152. Stillwater, OK.
PAN-PACIFIC ENTOMOLOGIST
72(1): 17-26, (1996)
GEOGRAPHIC VARIATION IN ARBOREAL SPIDER
(ARANEAE) COMMUNITIES ON DOUGLAS-FIR IN
WESTERN OREGON
J. HALAS,’ D. W. Ross, R. R. MASON,?
T. R. TORGERSEN,” AND A. R. MOLDENKE!
Department of Forest Science, Oregon State University, Corvallis, Oregon 97331;
1Department of Entomology, Oregon State University, Corvallis, Oregon 97331;
*Pacific Northwest Research Station, United States Department of Agriculture,
Forest Service, La Grande, Oregon 97850
Abstract. —Relative abundance and community structure of arboreal spiders were estimated in
sapling stands of Douglas-fir, Psewdotsuga menziesii (Mirbel) Franco. Samples were collected on
two dates between June and August 1992 in three geographic provinces in western Oregon. A
total of 7693 spiders of 15 families, 46 genera and at least 62 species were collected from lower
crown branches. On both dates, mean spider density was significantly different among the prov-
inces (P < 0.0001 and P < 0.01). Overall web-spinning spiders dominated in both collections,
comprising 63 and 58% of the total spider fauna. The principal web-spinners were the Linyphiidae
and Erigonidae, followed by the Araneidae, Theridiidae and Tetragnathidae. The spectrum of
hunting families was dominated by the Salticidae and Philodromidae. Both families combined
represented 79 and 78% of spiders in this guild for the two sampling dates. On both dates, the
proportions of web-spinning to hunting spiders were significantly different among the provinces
(P < 0.001 and P < 0.0001). Web-spinners were dominant in coastal areas, whereas hunters
were a major group further inland. On both collecting dates, all geographic provinces were similar
in terms of their species composition. A slightly greater similarity was detected between the two
Coast Range provinces.
Key Words.— Arachnida, Douglas-fir, Pseudotsuga menziesii, spider community structure, geo-
graphic variation, western Oregon
The abundance, polyphagous habit, and widespread distribution of spiders
make these predators well suited as potential natural controls of insect populations
(Riechert 1974, Riechert & Lockley 1984). In forest ecosystems, spiders have been
reported preying on various life stages of important forest insect pests including
the Nantucket pine moth, Rhyacionia frustrana (Comstock), Douglas-fir tussock
moth, Orgyia pseudotsugata (McDunnough), and eastern spruce budworm, Chor-
istoneura fumiferana (Clemens) (Turnbull 1956, Loughton et al. 1963, Eikenbary
& Fox 1968, Dahlsten et al. 1977, Jennings & Houseweart 1978, Fichter 1984,
Mason & Paul 1988, Jennings & Houseweart 1989).
Spiders in Pacific Northwest forests are poorly studied despite their abundance
and potential roles in regulating insect pest populations. Turnbull (1956) studied
spider predation on C. fumiferana in British Columbia, Canada. Moldenke et al.
(1987) provide a list of species, key, and estimates of relative abundance for
arboreal spiders in Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco, and true
fir, Abies spp., forests of the Pacific Northwest. Fichter (1984) documented the
densities of arboreal spiders associated with early instar Douglas-fir tussock moth
in Abies concolor (Gordon & Glendinning) Hildebrand, in central California.
Mason (1992) studied spider populations on Douglas-fir and true firs in eastern
Oregon and Washington. He found a similar familial structure of spider com-
18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
munities across the region, with the spectrum largely dominated by two families
of hunting spiders. We hypothesized that spider abundance and community struc-
ture in the more mesic forests of western Oregon would differ from that in the
dry eastern portions of the state. This study quantifies spider abundance and
community structure on Douglas-fir canopies in three distinct geographic regions
in western Oregon.
MATERIALS AND METHODS
Geographic Provinces and Study Sites. —The study was conducted in three geo-
graphic provinces along an east-west transect in western Oregon. The provinces
were the western Coast Range, the eastern Coast Range, and the western Cascade
Range. Sites in the western Coast Range were located on the Siuslaw National
Forest about 10 km east of Waldport, in Lincoln County, Oregon. Sites in the
eastern Coast Range were located on the McDonald and Paul M. Dunn State
Forests about 10 km northwest of Corvallis, in Benton County, Oregon. Sites in
the western Cascade Range were located on the H. J. Andrews Experimental Forest
within the Willamette National Forest about 15 km northeast of Blue River, in
Lane and Linn Counties, Oregon. Mean elevations for sites within the three
provinces were 228, 270, and 845 m, respectively. All three provinces are char-
acterized by wet winters, warm dry summers and mild temperatures throughout
the year. The annual precipitation values for the provinces moving from west to
east are 228, 178 and 230 cm, respectively, with the majority of precipitation
occurring between November and March. Mean annual temperatures for the
provinces moving from west to east are 10.4, 9.8 and 7.9° C, respectively (Taylor
& Bartlett 1993, 1994, 1995).
Study sites were mostly pure sapling stands of Douglas-fir (< 15-years-old) with
occasional sparse components of western hemlock, Tsuga heterophylla (Rafin-
esque) Sargent, and western redcedar, Thuja plicata D. Don. The ground vege-
tation included dense patches of bracken fern, Pteridium aquilinum (Linnaeus)
Kuhn in Decken, salmonberry, Rubus spectabilis Pursh, Gaultheria, Gaultheria
shallon Pursh, fireweed, Epilobium angustifolium Linnaeus, and Pacific rhodo-
dendron, Rhododendron macrophyllum D. Don ex G. Don.
Sampling Methods.—Sampling procedures used in this study were similar to
those of Mason (1992). A total of eight sites in each of the geographic provinces
were sampled on two dates between Jun and Aug 1992 approximately one month
apart. The timing of sampling was selected to coincide with the period of maxi-
mum density and diversity of spider populations (Uetz 1979, Hatley & MacMahon
1980, Dodbel et al. 1990). On each collecting date, all provinces were sampled
within a two-week period. A total of fifty sapling Douglas-fir trees were randomly
selected for sampling at each of the sites. On each tree, 45-cm-long tips of three
branches selected from the lower third of the canopy were sampled by beating
over a hand-held drop cloth (Paul 1979). Sampling of the lower canopy was
assumed to provide a good estimate of the structure of the spider community
throughout the whole canopy. It has been shown that the relative abundance and
density of spiders in Douglas-fir remains fairly constant in relation to the height
of the canopy (Fichter 1984, Voegtlin 1982, Moldenke, unpublished data).
Spiders dislodged onto the drop cloth were collected using a portable vacuum
(Paul & Mason 1985) and preserved in 75% ethanol. Ten trees were sampled
1996 HALAJ ET AL.: ARBORAL SPIDERS ON DOUGLAS-FIR 19
before emptying the vacuum, resulting in five 30-branch samples from each site.
In the laboratory, spiders were sorted and identified. The identifications of spec-
imens were based mostly on taxonomic keys in Roth (1985) and Moldenke et al.
(1987). The densities of spiders were calculated as numbers of individuals per m7?
of branch area (Mason 1992).
Data Analyses. —Mean arthropod densities and ratios of web-spinning to hunt-
ing spider densities were compared among the three provinces by a one-way
analysis of variance (ANOVA). Differences in spider densities and diversity be-
tween the two sampling dates were tested using a paired f-test. In order to satisfy
the assumption of equal variance in ANOVA, mean arthropod densities and ratios
of spider guilds were transformed to their natural logarithms (In) before analyses.
Back-transformed values and 95% CI are reported here. Means were compared
and separated by Fisher’s protected least significant difference (LSD) test (Steel
& Torrie 1980). Diversity of spider populations was determined individually for
each of the sites using the Shannon diversity index (H’) (Pielou 1975). The pa-
rameter of evenness (J’) of this index which measures the equitability of abundance
of individual taxa in the sample was calculated. J’ ranges between O and 1 with
the value of | characteristic of hypothetical communities with an equal abundance
of all taxa. Similarity of spider populations among the provinces was determined
using the Sorensen similarity index (C,) (Southwood 1992).
Differences in spider diversity and evenness among the provinces were tested
by ANOVA and means were compared and separated by Fisher’s protected LSD.
All statistical analyses were performed using SAS computer programs (SAS In-
stitute 1985).
RESULTS AND DISCUSSION
Absolute Densities of Spiders.—Statistical analyses revealed significant differ-
ences in mean spider densities (total) among the three provinces on both sampling
dates (F = 28.59; df = 2, 21; P < 0.0001; and F = 5.02; df = 2, 21; P < 0.01).
In June, mean spider densities were significantly higher in the eastern Coast Range
compared with the other two regions which had similar densities (Fig. 1). In late
July—-early August, mean spider densities in both Coast Range regions were not
significantly different and spider densities in the eastern Coast Range were sig-
nificantly higher than those in the western Cascade Range (Fig. 1). Mean spider
densities ranged from 4.6 to 16.7 individuals per m? of branch area similar to
those found by Mason (1992) on true fir and Douglas-fir in eastern Oregon and
by Jennings & Dimond (1988) on balsam fir, Abies balsamea (Linnaeus) Miller,
and spruces, Picea spp., in east-central Maine. Some factors that may influence
spider abundance include tree density, type and abundance of associated vege-
tation, abundance of potential prey organisms, patterns of air movement, and
climate. Significant differences in spider densities among the provinces may have
been caused by natural variation in the structure of branches and foliage (Gun-
narsson 1988, 1990). In addition, we observed an increased abundance of her-
baceous and shrub vegetation at the Coast Range sites in comparison with the
Cascade Range sites. Further, the ground cover communities in the Coast Range
sites were dominated by dense, structurally more complex, patches of salmonberry
and Gaultheria, whereas the sites in the Cascade Range were mostly covered with
patches of architecturally simpler bracken fern and rhododendron.
20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
25
[ _]June
VN uly—August
20
4G
RNOW
ROH
OY
Y
j
U
SN
Number per m2Z branch area
N
W-—Coast E—Coast W—Cascade
Ranges Range Range
Figure 1. Mean spider densities (+ 95% CI) per m? of Douglas-fir branch area in three geographic
provinces in western Oregon. Bars with the same letters are not significantly different (P = 0.05).
There was a significant increase in spider densities between the first and second
sampling dates (t = 7.53; df = 23; P < 0.0001) (Fig. 1). Although the small number
of collecting dates in our study limits temporal analysis of trends in the spider
populations, we suspect that the increase in spider densities can be attributed to
the hatch of eggs laid in the early summer by individuals overwintering as im-
matures (Loughton et al. 1963). This assumption is supported by an increased
number of immature individuals (over 70% of total spiders) in the later samples.
Spider Diversity and Guild Composition. — A total of 7693 spiders of 15 families,
46 genera, and at least 62 species were collected from the foliage of Douglas-fir
in western Oregon. Our data are comparable to those of Turnbull (1956) who
collected 75 species of spiders representing 15 families from Douglas-fir foliage
in British Columbia, Canada. Mason (1992) collected spiders from 11 families
on Douglas-fir and true fir branches in interior Pacific Northwest forests. We
recorded about twice the number of genera and species collected on balsam fir
and spruces from east-central Maine (Jennings & Dimond 1988).
Analyses of variance indicated significant differences in spider diversity (H’)
among the provinces in June reflecting a higher diversity in the western Coast
Range and western Cascade Range provinces compared with the eastern Coast
Range province (F' = 9.32; df= 2,21; P < 0.01)(Table 1). No significant differences
were detected in late July-early August (F = 0.20; df = 2,21; P > 0.80). There
were significant differences in the evenness component of diversity (J') among the
provinces in June (F = 9.86; df = 2,21; P < 0.01), but not in late July—early
August (F = 0.06; df = 2,21, P > 0.90) (Table 1).
1996 HALAJ ET AL.: ARBORAL SPIDERS ON DOUGLAS-FIR 21
Table 1. Mean values (SEM) of the Shannon diversity index (H’) and evenness (J’) for spider
samples collected in western Oregon, 1992.
June Late July—-early August
Province n H’ J H’ J’
W—Coast Range 8 2.43 (0.02)a 0.86 (0.0l)a 2.28 (0.08)a 0.74 (0.02)a*
E—Coast Range 8 2.19 (0.10)b 0.71 (0.03)b 2.41 (0.04)a 0.77 (0.01)a
W—Cascade Range 8 2.32 (0.05)ab 0.83 (0.01)a 2.25 (0.06)a 0.75 (0.02)a*
4 Means within a column followed by the same letter are not significantly different (P = 0.05).
* Denotes a significant difference between two collecting dates (P = 0.05).
Significant differences in the evenness (J’) and diversity (H’) among the prov-
inces in June reflected a build up in the relative abundance of the Erigonidae
detected in the eastern Coast Range. In addition, a significant drop in the evenness
between June and late July—-early August in the western Coast and Cascade Ranges
(t = 5.87; df = 7; P < 0.001 and t = 3.58; df = 7; P = 0.009, respectively) can
be attributed to an increase in the relative abundance of the Linyphiidae and
Salticidae, respectively, in samples from these provinces. All three families were
dominated by only a few species comprising over 95% of individuals (see below).
Overall, relatively low values of evenness in our samples indicate an unequal
abundance of species reflecting the presence of a few dominant species. These
results are generally consistent with those of Mason (1992) who compared spider
community structure at the family level among geographic regions of the interior
Pacific Northwest. He found that a few families were dominant in each region,
but individuals of most families were present in low numbers. Similarly, Turnbull
(1966) recorded only a few spider species represented by a large number of in-
dividuals and the majority of species represented by a few individuals 1n a pasture
community in Ontario, Canada.
On both collecting dates, all three provinces were similar in terms of their
species composition. A slightly greater similarity was detected between both Coast
Range provinces in June and late July-early August (0.70 and 0.80, respectively)
(Table 2). In addition, relatively low values of C, between the two collecting dates
in each of the provinces (0.67, 0.75 and 0.69) indicate substantial temporal changes
in the species composition of spider populations (Table 2).
Spiders were divided into two functional guilds based on their foraging strat-
egies; web-spinners and hunters. Relative abundance of these two functional cat-
egories of spiders is often used to describe spider communities (Loughton et al.
1963, Fichter 1984, Jennings & Dimond 1988, Mason 1992). On both sampling
dates, the ratios of web-spinning to hunting spiders differed significantly among
the provinces (F = 13.23; df = 2,21; P < 0.001 and F = 23.24; df = 2,21; P <
0.0001, respectively). In general, the abundance of web-spinning spiders relative
to hunting spiders declined with distance from the coast (Fig. 2). On both sampling
dates, web-spinners were 1.4 to 2.3 times more abundant than hunters in both
regions of the Coast Range. In contrast, hunters were more abundant than web-
spinners in the western Cascade Range. Mason (1992) and Turnbull (1956) both
reported a higher abundance of hunting spiders relative to web-spinners on Doug-
las-fir in interior Pacific Northwest forests.
Table 2. Values of the Sorensen similarity index (C,) for spider samples collected in western Oregon, 1992.
June Late July—-early August
Province ~ W—Coast Range +~=E—CoastRange | W—CascadeRange | W—CoastRange | E—CoastRange | W—Cascade Range
June
W— Coast Range 1.00
E— Coast Range 0.70 1.00
W— Cascade Range 0.63 0.63 1.00
July—August
W —Coast Range 0.67 0.74 0.63 1.00
E—Coast Range 0.63 0.75 0.65 0.80 1.00
W—Cascade Range 0.55 0.69 0.69 0.74 0.78 1.00
CC
LSIDOTOWOLNG OIIOVd-NVd AHL
(DZL TOA
1996 HALAJ ET AL.: ARBORAL SPIDERS ON DOUGLAS-FIR 23
[| June
Wii, Suly- August
Ratio
W-Coast E—Coast W-—Cascade
Range Range Range
Figure 2. Average ratios (+ 95% CI) of web-spinning to hunting spiders in three geographic
provinces in western Oregon. Bars with the same letters are not significantly different (P = 0.05).
Spider Families and Dominant Species. Among the web-spinners, the Liny-
phiidae, Erigonidae and Araneidae combined were the principal families in all
three provinces comprising over 74% of the spiders in this guild (Fig. 3). These
were followed in abundance by the Theridiidae and Tetragnathidae. The Dictyn-
idae and Uloboridae comprised <1% of all spiders collected in this guild. Com-
pared with similar data from the interior Pacific Northwest (Mason 1992) the
Linyphiidae and Erigonidae were much more abundant and the Dictynidae were
much less abundant in our samples.
The Linyphiidae in all three provinces were dominated by Pityohyphantes rub-
rofasciatus Keyserling and Pityohyphantes costatus (Hentz); less abundant but
fairly common was Gnathantes ferosa Chamberlin & Ivie. Among the erigonids,
Ceraticelus atriceps (O. P.-Cambridge) and Ceraticelus vesperus Chamberlin &
Ivie were the most common species. In addition, Spirembolus mundus Chamberlin
& Ivie and Erigone denticulata Chamberlin & Ivie were frequently collected in
the western Coast Range. The theridiids were dominated by Dipoena nigra (Emer-
ton), Theridion simile C. L. Koch, Theridion neomexicanum Banks, Theridion
sexpunctatum Emberton, and Theridion differens Emerton. Araniella displicata
(Hentz) and Cyclosa conica (Pallas) were the most common araneid species.
The guild of hunting spiders was similar to that described by Turnbull (1956)
and Mason (1992). The Salticidae and Philodromidae were dominant hunters
composing over 75% of the spiders in this guild (Fig. 3). The salticids alone
represented about 50% of hunting spiders (Fig. 3). Metaphidippus aeneolus Curtis,
a species common throughout coniferous forests of the Pacific Northwest (Mol-
24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
100% 100% AT B
80 80 FS - Ly
70 70 ca oi ate
60 60 He ot
50 50 ae
40 40 ES - st
30 30
20 20
10 10
0 0
W-Coast E—Coast W—Cascade W-Coast E—Coast W—Cascade
Range Range Range Range Range Range
Figure 3. Relative abundance of individual spider families in the guild of web-spinners (A) and
hunters (B) in combined samples from both collection dates in three geographic provinces. Family
abbreviations are: ER, Erigonidae; SL, Salticidae;, LI, Linyphiidae; AR, Araneidae; PH, Philodromidae;
TR, Theridiidae; TE, Tetragnathidae; CL, Clubionidae; AN, Anyphaenidae; TO, Thomisidae; OX,
Oxyopidae; DI, Dictynidae; GN, Gnaphosidae; UL, Uloboridae; LY, Lycosidae.
denke et al. 1987), was the most common jumping spider in our samples. The
Philodromidae was dominated by Philodromus rufus pacificus Banks and Phil-
odromus spectabilis Keyserling. In contrast to the previous studies in interior
forests, our samples from western Oregon contained higher numbers of nocturnal
hunters like the Clubionidae and Anyphaenidae. This was particularly true in the
western Coast Range where clubionids alone comprised about 20% of the hunting
spiders (Fig. 3). Both families include hunting spiders that forage at night, mostly
on grasses and deciduous shrubs but which stray to tree canopies. The higher
abundance of these spiders in western Oregon may reflect more diverse habitats
with a rich layer of ground vegetation. The Oxyopidae, classified together with
salticids as agile hunters, were primarily limited to the Cascade Range, almost
completely lacking in samples from the Coast Range (Fig. 3).
The abundance of web-spinners in the Coast Range may be attributed to several
factors. For sit-and-wait predators, such as web-spinning spiders, the location of
a suitable site is critical to their success (Turnbull 1973, Riechert and Gillespie
1986). Robinson (1981) demonstrated experimentally that web-spinning spiders
prefer more complex substrates that provide a greater number of points of at-
tachment for web construction. Similarly, Chew (1961) postulated that a low
density of shrubs providing less opportunities for web construction was among
the factors responsible for under-representation of web-spinning spiders in a desert
community. The abundance of a more diverse vegetation at the coast sites would
add structural complexity to young conifer plantations possibly improving the
habitat for web-spinning spiders. Further, Scheidler (1990) observed in Germany
that about 70% of spiders inhabiting herbaceous vegetation in forest clearings
were web-spinning spiders dominated by linyphiids. This might explain the dom-
inance of this group in our samples.
1996 HALAJ ET AL.: ARBORAL SPIDERS ON DOUGLAS-FIR 25
In addition, the more consistent air movement near the coast may facilitate
dispersal of these spiders and thus their movement from the herbaceous and shrub
layers to tree canopies. Because of the sit-and-wait foraging strategy of these
spiders, numbers of prey coming into contact with the web is a limiting factor.
The regular air movements in coastal areas may increase the prey supply and,
consequently, affect the suitability of the site.
In summary, our data show that spider communities of western Oregon are
similar in terms of their species composition but significant differences exist in
the absolute density and relative abundance of individual species and families.
There is an increase in web-spinning spiders relative to hunters in forests closer
to the coast. Climatically, all three provinces are very similar, with slightly lower
annual temperatures in the western Cascades reflecting the higher elevation of the
study sites in this province.
Our assessment of the habitat quality indicates that both coast provinces were
characterized by denser and structurally more complex patches of ground vege-
tation which may be partly responsible for the observed differences. In addition,
microclimate conditions at the site or local abundance of prey organisms may be
equally important in determining the structure of spider communities. More re-
search is needed to describe the composition and temporal changes of spider
communities in this region and to experimentally assess the relative importance
of the habitat structure, prey availability and climatic factors in the biology of
these predators.
ACKNOWLEDGMENT
We thank D. T. Jennings (University of Maine, Orono) for identification of
spider taxa collected in June 1992 and John D. Lattin and David A. Perry (Oregon
State University, Corvallis) for their constructive comments on an early draft of
the manuscript. This research was supported, in part, by funds provided by the
United States Department of Agriculture, Forest Service, Pacific Northwest Re-
search Station.
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PAN-PACIFIC ENTOMOLOGIST
72(1): 27-30, (1996)
HYMENOPTERA REARED FROM
PLAGIOTROCHUS SUBERI
(HYMENOPTERA: CYNIPIDAE) GALLS IN CALIFORNIA
ROBERT L. ZUPARKO
Laboratory of Biological Control, University of California, Berkeley,
1050 San Pablo Avenue, Albany, California 94706
Abstract.—Nine species of Hymenoptera were reared from stem galls induced by Plagiotrochus
suberi Weld on Quercus suber L. in Albany, California. The most numerous species reared was
P. suberi, but Euderus crawfordi Peck was the most common parasitoid reared. Other species
reared were: Aprostocetus pattersonae (Fullaway), A. sp. nr. verrucarii (Balduf), Acaenacis taciti
(Girault), Sycophila wiltzae (Balduf), Sycophila sp. possibly foliatae (Ashmead), Brasema sp. and
Ceraphron sp. The rearings of all parasitoids (except E. crawfordi and S. wiltzae) constitute new
host records. All described parasitoid species are native to North America and represent new
associations with the introduced P. suberi.
Key Words. —Insecta, Plagiotrochus suberi, Quercus suber, Euderus crawfordi, new associations,
parasitoids
Plagiotrochus suberi Weld (Hymenoptera: Cynipidae) was first described by
Weld (1926) from specimens collected in California. It is restricted to cork oak
(Quercus suber L.) and it produces intrinsic stem galls on the twigs, occasionally
resulting in economic damage (Essig 1943). Weld (1926) suggested the cynipid
was a native Nearctic species that shifted hosts when cork oaks were introduced
from Europe, but Kinsey (1935) disagreed and considered it to be the agamic
form of some unspecified European species. The latter view was confirmed when
P. suberi was found in Switzerland in 1957 and in Portugal in 1963 (Bailey &
Stange 1966). It has since been reported from Argentina (Diaz 1972).
The biology of P. suberi was reported by Bailey & Stange (1966) based on studies
in Davis, California. The only parasitoid they found associated with P. suberi was
Euderus crawfordi Peck (Hymenoptera: Eulophidae).
The presence of P. suberi on cork oaks at our experiment station provided an
opportunity to study the guild of Hymenoptera associated with the galls. This
paper reports the species of Hymenoptera reared, and their relative abundance
and chronology of emergence.
MATERIALS AND METHODS
Two Q. suber trees located at the University of California, Berkeley’s Gill Tract
in Albany, California were sampled from 1992-1994. All samples were randomly
chosen from the distal ends of branches accessible from the ground or a 3 m
stepladder. Twigs with swellings or emergence holes were broken into pieces about
7.5 cm long and placed in shell vials stoppered with cotton. These twigs ranged
from 1.5—8 mm in width, and emergence holes were noted in twigs that were 2-
8 mm wide.
In 1992 about 100 twigs were collected on 5 May; these were held inside at
room temperature until 9 May 1993. The trees were sampled eight times (every
1-2 weeks) between 24 Feb and 14 May 1993. Approximately 10-20 twigs were
28 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Table 1. Number of adult Hymenoptera emerging from Quercus suber twigs, collected in Albany,
California, 1992-1994.
Date of emergence
"27Mar- 1May- 8May—- ~—s«215 May-—-22 May-__29 May
Year Species 30 Apr 7 May 14 May 21 May 28 May 16 Jul Total
19922 P. suberi —4 _ +++ +++ + 0 ca. 150
E. crawfordi — _ 33 9 5 2 49
Other species — — 0 0 3 10 13
1993> P. suberi 0 if: 24 53 29 by 115
E. crawfordi 8 1 3 3 3 3 21
Other species 9 0 0 0 1 1 11
1994¢ P. suberi 0 1 10 10 49 96 166
E. crawfordi 59 13 15 5 5 15 112
Other species 29 1 7 0 3 8 48
4From 100 twigs, collected on 5 May, 1992.
> From ca. 120 twigs, collected 24 Feb-14 May, 1993.
° From 1809 twigs, collected 27 Jan—26 May, 1994.
4 — = not sampled; +++ = many; + = few.
collected each time. In 1994, 10 twigs were collected once per week from 27 Jan
to 26 May (18 total samples). The 1993 and 1994 samples were held outside,
shaded from direct sunlight, until 30 July of each year. The number, sex and dates
of emergence of adults were recorded for all species (except in 1992, when numbers
of P. suberi were only estimated).
RESULTS
In all years P. suberi was the most common species reared, its 400+ adults
accounting for about 63% of all emergents (Table 1). The chronology of P. suberi
emergence was similar each year, peaking in May. No cynipids emerged from
samples collected in February or March. The earliest emergence in any season
was on 2 May 1994, and the latest on 16 Jun 1994. Only females were found.
The exact number ofadults emerging from each twig was not recorded, but samples
collected in April or May produced an approximate mean number of 1.5 adults
per twig in 1992, 1.53 in 1993, and 2.08 in 1994. The highest rate was 8.2 adults
per twig, from a sample collected in mid-May 1994.
Euderus craw fordi was the most common parasitoid reared, representing 23%
of all emergents in 1992, 14% in 1993 and 34% in 1994. In 1992, emergence by
this species peaked in the first half of May. In 1993, low numbers emerged steadily
throughout the spring with no distinct peak, and in 1994, emergence peaked in
mid-April. This was the only species reared from twigs collected during all 26
sampling periods in 1993 and 1994. One female emerged from a twig over 5
months after collection (27 Jan to 6 Jun 1994), and one male emerged over 4
months later (3 Feb to 14 Jun 1994).
Nine Aprostocetus pattersonae (Fullaway) (Hymenoptera: Eulophidae) emerged
in June 1992 and two in July 1994. This species emerged only from twigs collected
in May. Eight A. sp. nr. verrucarii (Balduf) were reared in April 1993 (twigs
collected from March-April) and 31 from April-May 1994 (collected January-—
April).
1996 ZUPARKO: PLAGIOTROCHUS SUBERI PARASILOIDS 29
Three females and one male Acaenacis taciti (Girault) (Hymenoptera: Pter-
omalidae) were reared in May 1992, and a single male in June 1993 (collected in
May). A male Brasema sp. (Hymenoptera: Eupelmidae) emerged in May 1993
(collected in May), and two females and nine males in May-June 1994 (collected
from January—May). In 1994, two female and one male Sycophila wiltzae (Balduf),
and a female Sycophila sp. possibly foliatae (Ashmead) (Hymenoptera: Eurytom-
idae) were reared in May (all collected in May). A single Ceraphron sp. female
(Hymenoptera: Ceraphronidae) emerged in April 1993.
DISCUSSION
The emergence pattern of P. suberi reported here agrees with that reported by
Bailey & Stange (1966). The absence of cynipid emergence from the samples
collected in February and March is probably due to larval mortality brought about
by the removal of the twigs from the trees. This species pupates in April, after
which any change in its substrate would have a decreased impact.
Bailey & Stange (1966) calculated that the average number of twig wasp exit
holes over 7 years ranged from 0.4 to 3.0 per linear cm of twig. From twigs
collected in April or May, I reared approximately 0.2 to 0.3 P. suberi adults per
cm. Two factors help explain this discrepancy. First, Bailey & Stange may have
overestimated P. suwberi numbers if they did not distinguish exit holes made by
E. crawfordi. Second, my methods underestimate P. suberi, because additional
adults might have emerged prior to collection, or died without emerging due to
trauma caused by the breaking of the twig. The density of exit holes I observed
in this study appeared similar to that pictured in Figure 4 of Bailey & Stange
(1966).
Albany may also have a higher parasitoid population than Davis. Bailey &
Stange (1966) noted that E. crawfordi reached “high numbers” in one tree, but
appeared to be of little importance in limiting the cynipid’s populations. In the
present study, the ratio of emergent E. craw fordi to P. suberi reached 1:3 in 1992,
and 2:3 in 1994.
Euderus crawfordi is known only from California and Arizona, and its only
other recorded host is Dryocosmus coxii (Bassett) (Hymenoptera: Cynipidae)
(Yoshimoto 1971). Dryocosmus coxii 1s closely related to P. suberi and produces
stem galls on Quercus emoryi Torrey and Q. hypoleuca Engelmann in Arizona
and New Mexico (Burks 1979). The parasitoid may have moved over onto P.
suberi when the latter was introduced through the planting of Q. suber in the
southwestern United States. Although I made no direct observations of parasit-
ization, the emergence pattern and relatively high numbers of FE. craw fordi support
its characterization as a primary parasitoid.
Both Aprostocetus species belong to the subgenus Quercastichus LaSalle, which
is known only from cynipid galls on oaks in North America (LaSalle 1994).
Sycophila wiltzae was reared from undetermined galls on Q. suber in San Jose in
1918 (Balduf 1932). The cynipids issuing from this material (= Hopkins U.S. No.
15608a) were eventually determined to be P. suberi (Weld 1926). Sycophila wiltzae
has also been reared from Andricus wiltzae Fullaway galls on Q. /obata Nee (Balduf
1932). The second Sycophila species resembles S. foliatae, which has been reared
from four genera of gall-forming cynipids on oaks in the eastern United States
(Burks 1979).
30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
The rearings reported here represent new host associations for both Aprostocetus
species, A. taciti, Sycophila sp., Braesma sp. and Ceraphron sp.
Plagiotrochus suberi is uncommon in Europe: although its sole known host is
native there, the cynipid was unknown there until 39 years after its description
in the New World (Bailey & Stange 1966). In contrast, the cynipid is common
throughout California (Essig 1943). This is a relatively recent introduction, with
the first collection made in 1918 (Weld 1926). In 1966, Bailey & Stange noted
that Q. suber in Davis had appeared to be pest-free since about 1930, but “in
recent years’” had begun to deteriorate due to the cynipid’s activity. They con-
cluded that E. crawfordi does not provide control of P. suberi.
Although the endemic natural enemy complex appears to be better-developed
in Albany than in Davis, the cynipid is still very numerous here. There are also
no records of natural enemies of P. suberi from Europe. This does not necessarily
indicate an absence of such organisms, but more likely is a reflection of the relative
scarceness of the cynipid itself.
ACKNOWLEDGMENT
Determinations were made by Kathy Schick (Plagiotrochus) and Dr. Steve
Heydon (Acaenacis), University of California, Davis, Dr. Michael Schauff (Eu-
derus) and Dr. Eric Grissell (Sycophila), Systematic Entomology Laboratory, Ag-
ricultural Research Service, U.S. Department of Agriculture, Dr. John LaSalle
(Aprostocetus), International Institute of Entomology (London), and Dr. Gary
Gibson (Braesma), Agriculture Canada (Ottawa). I am also grateful to Dr. Leo
Caltagirone and Dr. Ken Hagen of U.C. Berkeley for their comments on and
insights into this study, and to the two anonymous reviewers for their helpful
comments improving the ms.
LITERATURE CITED
Bailey, S. F. & L. A. Stange. 1966. The twig wasp of cork oak—its biology and control. J. Econ.
Entomol., 59: 663-668.
Balduf, W. V. 1932. Revision of the chalcid flies of the tribe Decatomini (Eurytomidae) in America
north of Mexico. Proc. U.S. Nat. Mus., 79: 1-95.
Burks, B. D. 1979. Families Eurytomidae and Cynipidae. pp. 835-860; 1060-1107. Jn Krombein,
K. V., P. D. Hurd Jr., D. R. Smith & B. D. Burks (eds.). 1979. Catalog of Hymenoptera in
America north of Mexico, vol. 1. Smithsonian Institution Press, Washington, D.C.
Diaz, N. B. 1972. Una nueva plaga del alcornoque en la Republica Argentina. Rev. Soc. Entomol.
Argentina, 34: 85-88.
Essig, E.O. 1943. The cork oak cynipid in California. J. Econ. Entomol., 36: 123-24.
Kinsey, A. C. 1935. The economic importance of the Cynipidae. J. Econ. Entomol., 28: 86-91.
LaSalle, J. 1994. North American genera of Tetrastichinae (Hymenoptera: Eulophidae). J. Nat. Hist.,
28: 109-236.
Weld, L. H. 1926. Field notes on gall-inhabiting cynipid wasps with descriptions of new species.
Proc. U.S. Nat. Mus., 68: 1-131.
Yoshimoto, C. M. 1971. Revision of the genus Euderus of America north of Mexico (Hymenoptera:
Eulophidae). Can. Entomol. 103: 541-578.
PAN-PACIFIC ENTOMOLOGIST
72(1): 31-36, (1996)
SUITABILITY OF A NON-HOST PALO VERDE FOR
DEVELOPMENT OF STATOR LIMBATUS (HORN)
(COLEOPTERA; BRUCHIDAE) LARVAE
CHARLES W. Fox, AMy D. HARBIN, AND TIMOTHY A. MOUSSEAU
Department of Biological Sciences, University of South Carolina,
Columbia, South Carolina 29208
Abstract.—In the southwestern United States, there are five species of palo verdes (Cercidium
and Parkinsonia sp.; Fabaceae). Stator limbatus (Horn), a seed beetle, has been reared from four
of these: Cercidium floridum (Benth.), C. microphyllum (Torr.) Rose & Johnst., Parkinsonia
aculeata Linnaeus, and P. macra (Johnst.). However, despite extensive collections in North and
Central America, S. /imbatus has never been reared from P. texana (A. Gray) S. Watson. We
tested the suitability of P. texana as a host for a Texas population of S. /imbatus. Survivorship
of S. limbatus on P. texana was high relative to the other palo verde species (except for C.
microphyllum, on which survivorship was also high). Development time and body weight of
emerging adults on P. texanawere each approximately intermediate between those on C. floridum
and P. aculeata (on which beetles developed slowly and emerged small) and C. microphyllum
(on which beetles developed rapidly and became large adults). These data indicate that P. texana
is a suitable host for S. Jimbatus and that, relative to other species of palo verde (except C.
micro phyllum), P. texana is a high-quality host. However, our study examines only the suitability
of these palo verdes as hosts for S. /imbatus in a controlled laboratory experiment. We discuss
other hypotheses that may explain why P. texana is not used by S. Jimbatus in nature.
Key Words.—Cercidium, diet breadth, Fabaceae, host range, Parkinsonia
Most herbivorous insects feed on few of the plant taxa available to them (Fox
& Morrow 1981). It is often observed, however, that herbivores develop very
well, or even better, on plants they will not oviposit or feed upon than on plants
they regularly use (Dethier 1954, Waldbauer 1962). Stator limbatus (Horn) (Co-
leoptera: Bruchidae) is a generalist seed beetle that uses >50 host plants in its
large geographic range (from northern South America to the southwestern United
States; Johnson & Kingsolver 1976, Johnson et al. 1989). In the southwestern
United States, there are five species of palo verdes (Isley 1975), and S. limbatus
has been reared from four of these: Cercidium floridum (Benth.), C. microphyllum
(Torr.) Rose & Johnst. Parkinsonia aculeata Linnaeus, and P. macra (Johnst.)
(Johnson & Kingsolver 1976, Nilsson & Johnson 1993). However, despite exten-
sive collections in North and Central America, S. /imbatus has never been collected
on or reared from Parkinsonia texana (A. Gray) S. Watson (Nilsson & Johnson
1993, C. W. Fox, unpublished observation), although other insects, such as Mi-
mosestes amicus (Horn) (Coleoptera: Bruchidae) have been reared from both P.
texana and other palo verde species (Nilsson & Johnson 1993). Here, we test the
suitability of P. texana as a host for a Texas population of S. limbatus. We find
that P. texana is indeed a suitable host for S. /imbatus. We speculate on numerous
hypotheses to explain the failure of S. /imbatus to use P. texana as a host plant,
andalso speculate whether, despite an inability to collect S. /imbatus on this plant,
P. texana is actually a host for this beetle.
32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
MATERIALS AND METHODS
Stator limbatus for these experiments were collected from >50 Acacia greggii
A. Gray (Fabaceae: Mimosoideae) plants along a ~ 30 km stretch of Hwy 90, just
south of Van Horn, Texas. Beetles were collected by picking mature pods from
A. greggii. These pods were transferred to the lab, and seeds containing beetles
were separated from uninfested seeds. We estimate that the laboratory population
was initiated with > 200 field collected individuals. Before initiating the laboratory
experiments, beetles were reared in the lab for one generation at 27° C, 24 h light,
on A. greggil.
Virgin female S. /imbatus, collected from isolated seeds <24 h after emergence,
were weighed and paired with a virgin male (also <24 h post-emergence). Each
pair was confined in a 30 mm plastic dish with 12 seeds of one of the following
species; C. floridum, C. microphyllum, P. aculeata, P. texana, or A. greggii. Al-
though A. greggii is not a palo verde (and is even in a different subfamily of the
Fabaceae), it was included here so that these data can be compared with previous
and future projects (e.g. Siemens & Johnson 1990; Siemens et al. 1991, 1992; Fox
et al. 1994, 1995).
Dishes were checked at 24 h intervals, and seeds bearing eggs were transferred
to clean dishes, until a female had laid an egg on each of >10 seeds. All eggs were
reared to adult at densities of one beetle per seed (additional eggs were scraped
from each seed), 27° C, constant light. Development time, body weight, and
survivorship were recorded for all offspring. Development time was estimated as
the time between egg-laying and adult emergence, and thus includes embryonic,
larval, and pupal development time. Emerging adults were weighed individually
on an electronic balance within 24 h of adult emergence.
RESULTS AND DISCUSSION
As in other experiments examining S. /imbatus life history (Siemens & Johnson
1990; Siemens et al. 1991, 1992; Fox etal. 1994, 1995), survivorship on A. greggii
was very high relative to survivorship on the palo verde species (Table 1). Sur-
vivorship on P. texana, which has not been documented as a host for this beetle,
was also high compared to the other palo verdes (except for C. microphyllum, on
which survivorship was also high); beetles reared on C. floridum and P. aculeata
had much lower survivorship than beetles reared on P. texana. This is a surprising
result because C. floridum is heavily attacked by S. /imbatus in southern California
and Arizona (Mitchell 1977, Siemens & Johnson 1990), and females regularly lay
eggs on P. aculeata in California (where it has escaped from cultivation), Arizona
(where it is likely native in Yuma Co. and escaped from cultivation elsewhere)
and in Texas (where it is native throughout most of the state; Isley 1975), but S.
limbatus has never been reared from P. texana.
As previously demonstrated for C. floridum (Siemens et al. 1993; Fox et al.
1994, 1995), most of the mortality on each palo verde species occurred as larvae
penetrated the seed coat (except for mortality on C. microphyllum; Table 1). For
example, although egg-to-adult survivorship on P. aculeata was only 4.6 + 6.9%,
survivorship of larvae that successfully penetrated theseed coat was 77.3 + 34.4%.
Extractions from the seed coat applied to other host species suggest that mortality
on C. floridum is due largely to allelochemicals in the seed coat (Siemens et al.
1992). The high mortality of larvae entering P. aculeata and P. texana, relative
1996 FOX ET AL.: STATOR LIMBATUS HOST PLANTS 33
Table 1. Survivorship of Stator limbatus reared on five host plants. Males and females are lumped
because dead larvae could not be sexed. Values sharing the same letter (within columns) are not
statistically different (Mann-Whitney U-tests).
Host species n
Survivorship
Egg—adult Entering seed Within seed
Acacia greggii 28 0.93 + 0.14a 0.99 + 0.03a 0.94 + 0.14a
Cercidium floridum 31 0.25 + 0.21 0.35 + 0.23 0.67 + 0.31b
Cercidium microphyllum 29 0.89 + 0.18a 0.99 + 0.03a 0.89 + 0.18a
Parkinsonia aculeata 31 0.05 + 0.07 0.06 + 0.09 0.77 + 0.34ab
Parkinsonia texana 32 0.70 + 0.28 0.75 + 0.28 0.93 + O.lla
to mortality within these seeds and mortality entering other seed species, suggests
that P. aculeata and P. texana have similar seed defense mechanisms to those of
C. floridum, although those of P. texana are clearly less effective at preventing
damage to seeds than the defenses of C. floridum and P. aculeata.
The observed patterns of development time and body weight (Table 2) of
emerging beetles closely resembled those for survivorship: C. floridum and P.
aculeata were generally poor hosts for S. limbatus, while A. greggii and C. mi-
crophyllum were generally good hosts for S. limbatus. Development time and
body weight on P. texana were each approximately intermediate between those
on the poor and good hosts.
Our data thus indicate that P. texana is a suitable host for S. limbatus, and
that, relative to other species of palo verde, including C. floridum, which is heavily
attacked by S. /imbatus in Arizona, P. texana is a high-quality host. Nonetheless,
there is no evidence that this host is used by S. /imbatus in nature (Johnson &
Kingsolver 1976, Nilsson & Johnson 1993). However, our study examines only
the suitability of these palo verdes as hosts for S. /imbatus in a controlled labo-
ratory experiment. It is often observed that many herbivores develop very well
in the laboratory on plants that they will not use in nature (Dethier 1954; Wald-
bauer 1962; Wiklund 1974, 1975, 1982; Hsiao 1982; Thompson 1988), suggesting
that the selective environment influencing diet breadth evolution includes nu-
Table 2. Development time and body weight of Stator limbatus reared on five host plants. The
low sample sizes on Parkinsonia aculeata reflect very high mortality on this host, and thus few emerging
adults. Values sharing the same letter, with sexes, are not statistically different from each other (Mann-
Whitney U-tests).
Host species Sex n Development time (days) Body weight (mg)
Acacia greggii f 28 28.5 + 0.3a 3.13 + 0.04a
m pag 28.8 + 0.3a 3.45 + 0.07a
Cercidium floridum f 18 36.3 + 0.7b 2.74 + 0.08b
m 17 36.9 + 0.6b 3.03 + 0.14b
Cercidium microphyllum i px) 29.8 + 0.3c 3.64 + 0.05c
m 29 29.8 + 0.3c 3.80 + 0.06c
Parkinsonia aculeata f 7 36.0 + 1.3b 2.74 + 0.20a,b,d
m 3 36.7 + 2.2b 3.00 + 0.32a,b,d
Parkinsonia texana f 30 31.0 + 0.3d 3.00 + 0.06d
m 2H 31.1 + 0.3d 3.09 + 0.07b,d
34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Table 3. The mean size of seeds of Acacia greggii and the four palo verde species used in this
experiment (+ 1 SE).
Host species n Seed weight (mg)
Acacia greggii 150 187+ 5
Cercidium floridum 300 229'+ 2
Cercidium micro phyllum 201 156+2
Parkinsonia aculeata 150 99 +2
Parkinsonia texana 100 84+ 1
merous factors other than host suitability. Hypotheses proposed to explain this
imperfect relationship between host use and host suitability variously implicate
predators and parasitoids (Lawton & McNeill 1979, Bernays & Graham 1988),
plant apparency (Feeny 1976, Rhoades & Cates 1976), phenology (Tahvanainen
1983), abundance (Wiklund 1982), or reliability (predictability) (Futuyma 1976,
Cates 1981), and the interaction between plant chemistry and the neuro-physi-
ological capabilities of insects (Ehrlich & Raven 1964, Smiley 1978, Jermy 1984,
Fox & Lalonde 1993). Currently, there is insufficient data available to reject any
of the above hypotheses as explanations for S. /imbatus’s failure to use P. texana.
The small size of P. texana seeds relative to the other palo verde species (Table
3) does not likely prevent S. /imbatus from using this host. Although survivorship
was lower and surviving adults were slightly smaller when reared on P. texana
than when reared on either A. greggii or C. microphyllum (Table 2), beetles reared
on P. texana survived better and were of similar size to beetles reared on C.
floridum. Also, Mimosestes amicus (Horn), a seed beetle that is substantially larger
than S. /imbatus throughout most of its distribution (VW. amicus biomass >2 x
S. limbatus biomass), successfully uses P. texana in nature (Nilsson & Johnson
1993), although surviving M. amicus adults reared from P. texana are generally
much smaller than those reared from other hosts.
Possibly, beetles may ignore P. texana in favor of Acacia berlanderii Bentham,
which is interspersed with P. texana throughout southern Texas and is heavily
attacked by S. limbatus. Acacia berlanderii is closely related to A. greggii, and
thus survivorship on seeds of this host is likely high, such that it is a physiologically
more suitable host than P. texana. However, A. berlanderii pods dehisce early in
the year relative to P. texana, such that its seeds are not available after July,
whereas P. texanais available into late autumn. Our data suggest that S. limbatus
should use P. texana when A. berlanderii is not available. However, in field
collections we have found S. limbatus eggs on A. berlanderii seeds, but not on P.
texana seeds, even when the two species are within meters of each other (C. W.
Fox, personal observation).
Predation and parasitism may significantly influence the patterns of host use
by S. limbatus. For example, the parasitoid Uscana semifumipennis Girault (Hy-
menoptera: Trichogrammatidae) attacks eggs of S. limbatus on C. floridum in
Arizona (Siemens & Johnson 1992). Rates of parasitism may be affected by host
plant species, such that eggs laid on P. texana may have lower success than eggs
laid on alternative hosts due to parasitism of eggs and larvae, selecting for females
that avoid P. texana. However, mortality due to parasitism is density-dependant
on C. floridum (Siemens & Johnson 1992). If mortality due to parasitism on hosts
1996 FOX ET AL.: STATOR LIMBATUS HOST PLANTS 35
in southern Texas is also density-dependant, then P. texana would represent an
ecological escape from parasitoids (and possibly predators) because other S. /im-
batus are not currently using it.
Alternatively, we must acknowledge that, although S. /imbatus has not yet been
collected from P. texana, this shrub may actually be a host for S. /imbatus and
that more extensive collections may detect its use of this plant. However, even
if S. limbatus uses P. texana, the frequency of use is clearly very low; S. limbatus
is easily collected on Acacia species in southern Texas but, despite extensive
collections, has not been collected from P. texana. The above hypotheses proposed
to explain the absence of S. /imbatus on P. texana need also be considered to
explain the low frequency of use of this host, if indeed future surveys should
detect S. limbatus on P. texana.
ACKNOWLEDGMENT
We thank C. D. Johnson, L. A. McLennan, and K. J. Waddell for helpful
comments on this manuscript. G. Zigler provided housing while collecting beetles.
L. A. McLennan collected seed size data. Financial support was provided in part
by USDA/CSRS Grant no. 9301887 to T. A. Mousseau and an NSF post-doctoral
fellowship in environmental biology (DEB-9403244) to C. W. Fox.
LITERATURE CITED
Bernays, E. A. & M. Graham. 1988. On the evolution of host specificity in phytophagous arthropods.
Ecology, 69: 886-892.
Cates,R.G. 1981. Host plant predictability and the feeding patterns of monophagous, oligophagous,
and polyphagous insect herbivores. Oecologia, 48: 319-326.
Dethier, V. G. 1954. Evolution of feeding preferences in phytophagous insects. Evolution, 8:
33-54.
Ehrlich, P. R. & P. H. Raven. 1964. Butterflies and plants: a study in coevolution. Evolution, 18:
586-608.
Feeny, P. P. 1976. Plant apparency and chemical defenses. Rec. Adv. Phytochem., 10: 1-40.
Fox, C. W. & R. G. Lalonde. 1983. Confusion of hosts and the evolution of insect diet breadths.
Oikos, 67: 577-581.
Fox, C. W, K. Waddell & T. A. Mousseau. 1994. Host-associated fitness variation in a seed beetle
(Coleoptera: Bruchidae): evidence for local adaptation to a poor quality host. Oecologia, 99:
329-336.
Fox, C. W., K. Waddell & T. A. Mousseau. 1995. Parental host plant affects offspring life histories
in a seed beetle. Ecology, 76: 402-411.
Futuyma, D. J. 1976. Food plant specialization and environmental predictability in Lepidoptera.
Am. Nat., 110: 285-292.
Hsiao, T. H. 1982. Geographic variation and host plant adaptation of the Colorado potato beetle.
pp. 315-324. In Visser, J. H. & A. K. Minks (eds.). Proceedings of the Sth International
Symposium on Insect-Plant Relationships. Pudoc, Wageningen, the Netherlands.
Isley, D. 1975. Leguminosae of the United State. II. Subfamily Caesalpinioideae. Mem. N. Y. Bot.
Gard., 25: 1-228.
Jermy, T. 1984. Evolution of insect/host plant relationships. Am. Nat., 124: 609-630.
Johnson, C. D. & J. M. Kingsolver. 1976. Systematics of Stator of North and Central America
(Coleoptera: Bruchidae). U.S. Dept. Agric. Tech. Bull., 1537: 1-101.
Johnson, C. D., J. M. Kingsolver, & A. L. Teran. 1989. Sistematica del genero Stator (Insecta:
Coleoptera: Bruchidae) en Sudamerica. Opera Lilloana, 37: 1-105.
Lawton, J. H. & S. McNeill. 1979. Between the devil and the deep blue sea: on the problem of being
a herbivore. Symp. Brit. Ecol. Soc., 20: 223-224.
Mitchell, R. 1977. Bruchid beetles and seed packaging by Palo Verde. Ecology, 58: 644-651.
36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Nilsson, J. A. & C. D. Johnson. 1993. Laboratory hybridization of Stator beali and S. limbatus,
with new host records for S. Jimbatus and Mimosestes amicus (Coleoptera: Bruchidae). South-
west. Natur., 38: 385-387.
Rhoades, D. F. & R. G. Cates. 1976. Toward a general theory of plant anti-herbivore theory. Rec.
Adv. Phytochem., 10: 168-213.
Siemens, D. H. & C. D. Johnson. 1992. Density-dependent egg parasitism as a determinant of clutch
size in bruchid beetles (Coleoptera: Bruchidae). Envir. Entomol., 21: 610-619.
Siemens, D.H., C.D. Johnson, and R.L. Woodman. 1991. Determinants of host range in bruchid
beetles. Ecology, 72: 1560-1566.
Siemens, D. H., C. D. Johnson, and K. V. Ribardo. 1992. Alternative seed defense mechanisms in
congeneric plants. Ecology, 73: 2152-2166.
Smiley, J. T. 1978. Plant chemistry and the evolution of host specificity: new evidence from Heli-
conius and Passiflora. Science, 201: 745-747.
Tahvanainen, J. 1983. The relationship between flea beetles and their cruciferous host plants: the
role of plant and habitat characteristics. Oikos, 40: 433-437.
Thompson, J. N. 1988. Evolutionary ecology ofthe relationship between oviposition preference and
performance of offspring in phytophagous insects. Entomol. Exp. Appl., 47: 3-14.
Waldbauer,G.P. 1962. The growth and reproduction of maxillectomized tobacco hornworms feeding
on normally rejected non-solanaceous host plants. Entomol. Exp. Appl., 54: 117-124.
Wiklund, C. 1974. Oviposition preferences in Papilio machaon in relation to the host plants of the
larvae. Entomol. Exp. Appl., 17: 189-198.
Wiklund, C. 1975. The evolutionary relationship between adult oviposition preferences and larval
host plant range in Papilio machaon. Oecologia, 18: 185-197.
Wiklund, C. 1982. Generalist vs. specialist utilization of host plants among butterflies. pp. 181-191.
In Visser, J. H. & A. K. Minks (eds.). Proceedings of the Sth International Symposium on
Plant-Insect Relationships. Wageningen, the Netherlands.
PAN-PACIFIC ENTOMOLOGIST
72(1): 37-38, (1996)
Scientific Note
GNATHAMITERMES PERPLEX US (BANKS)
(ISOPTERA: TERMITIDAE): A NUISANCE STRUCTURAL
TERMITE PEST IN SOUTHERN CALIFORNIA
The genus Gnathamitermes Light consists of four species (Weesner, F. M. 1970.
Biology of termites. Academic Press, New York. Vol. 2). These termites are
primarily of a southern Nearctic distribution and they occur largely in the south-
western United States and portions of Mexico. Two species, Gnathamitermes
perplexus (Light) and Gnathamitermes nigriceps (Light) were described from spec-
imens taken in western Mexico (Light, S. F. 1930. Univ. Cal. Pub. Ent., 5: 175-
214). Gnathamitermes tubiformans (Buckley) occurs in the arid and semiarid
regions of New Mexico, Texas, Arizona, and northern Mexico (Snyder, T. E. 1949.
Smithsonian Misc. Coll., 112; Allen, C. T., D. E. Foster & D. N. Ueckert. 1980.
Environ. Entomol., 9: 461-466). Gnathamitermes perplexus (Banks) occurs in
southern California, Nevada, Arizona, and Texas. In southern California, it occurs
Figure 1. Surface scarification of wood caused by Gnathamitermes per plexus.
38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
in the coastal areas of San Diego County and inland in the arid and semiarid
desert regions of the southern portions of the state.
Gnathamitermes tubiformans has received attention because of its supposed
conflict with man in rangeland management for cattle production (Bodine, M. C.
& D. N. Ueckert. 1975. J. Range. Mgmt., 28: 353-358; Ueckert, D. N., M. C.
Bodine & B. M. Spears. 1976. Ecology, 57: 1273-1280; Allen, C. T., D. E. Foster
& D. N. Ueckert. 1980; Schaefer, D. A. & W. G. Whitford. 1981. Oecologia, 48:
277-283). Ecologists have attempted to assess its role in energy flow, nutrient
turnover, and nutrient cycling in desert ecosystems (Johnson, K. A. & W. G.
Whitford. 1975. Environ. Ent., 4: 66-70; Schaefer, D. A. & W. G. Whitford. 1979.
Bull. Ecol. Soc. Amer., 60: 128; Schaefer & Whitford 1981).
Ecological data on the distribution, foraging behavior, and food preference of
G. perplexus are presented elsewhere (Light 1930; Light, S. F. 1934. Termites and
termite control. Univ. Cal. Press, Berkeley, CA; Weesner 1970; Haverty, M. L &
W. L. Nutting. 1975. Ann. Ent. Soc. Amer., 68: 533-536; Haverty, M. IL & W.
L. Nutting. 1975. Environ. Ent., 4: 480-486). Gnathamitermes spp. were not
known to attack structures and they were considered to have no structural eco-
nomic significance (Light 1930, 1934).
On 26 Oct 1990, G. perplexus was found within a structure on Kalmia Street,
Murrieta, California where it created a nuisance problem and caused superficial
aesthetic damage. On 27 Sep 1994, it was discovered within a structure on west
Franklin Street, Lake Elsinore, California. At this location, G. perplexus caused
appreciable superficial scarification and aesthetic damage to baseboards, wall-
paper, drywall, ceiling tiles, and door frame moldings (Fig. 1).
Both of these infestations were of such intensity and persistence that it took
one complete subterranean termite treatment and several retreatments per prop-
erty to control them.
Gnathamitermes spp. were not previously reported to cause aesthetic damage
within structures. The two incidences reported here involving G. perplexus rep-
resent the first cases of a species of this genus entering structures and causing
nuisance problems and aesthetic damage.
Acknowledgment. —I thank John Chapman, Stoy Hedges, Rusty Bracho, and
Ken Hobbs for reading the manuscript and offering suggestions for improvement.
Hanif Gulmahamad, Terminix International, 9559 Center Avenue, Suite N.
Rancho Cucamonga, California, 917 30.
PAN-PACIFIC ENTOMOLOGIST
72(1): 39-40, (1996)
Scientific Note
THE DISTRIBUTION OF AQUATIC HETEROPTERA
(NOTONECTIDAE) IN HAWAII
Lentic habitats on the main Hawaiian islands were sampled for hemipterans,
particularly backswimmers (Heteroptera: Notonectidae), during Jul and Aug 1991
and Aug 1992. The primary habitats were ponds, stock troughs (concrete and
metal), and cisterns associated with stock troughs. Sampling was conducted on
Hawaii, Oahu, and Maui in 1991 and 1992, and on Kauai in 1992. During the
course of the sampling effort, three species of notonectids were collected, two
previously known in Hawaii and a single specimen of a genus that had not been
reported for the State and the Western Hemisphere. This latter specimen has since
been reported in the literature (Polhemus, D. A. 1995. Bishop Museum Occasional
Papers, 42: 42-43).
Of the two species of notonectids that had been recorded from the state, Buenoa
pallipes (Fabr.) was recorded around the turn of the century (Zimmerman, E. C.
1948. Insects of Hawaii. Volume 3: Heteroptera. University of Hawaii), whereas
Notonecta indica L. is far more recent. This was the first collection of the third
species, Anisops kuroiwae Matsumura, in the Hawaiian Islands. It is entirely
possible that these species have colonized the islands as the result of human
activities, perhaps as eggs in aquatic plants used in the aquarium trade. Even so,
all but A. kuroiwae have successfully exploited available still-water habitats on
all major islands surveyed. In their successful colonization of various islands after
the initial event, man-aided or not, they owe no small debt to the presence of live
stock. Most of the localities where notonectids were found in 1991 and 1992 were
either ponds that had been augmented to hold more water for cattle or metal/
concrete/plastic stock troughs.
The single male specimen of A. kuroiwae was collected on Maui in 1991 (Pol-
hemus 1995) and is notable as it is the first confirmed record for the Western
Hemispere. The recorded distribution of this species extends from India through
Burma, mainland China, Taiwan, the Philippines, Iriomote Island, and Okinawa
Island (Brooks, G. T. 1951. Univ. Kansas Sci. Bull., 34: 301-519). Brooks (1951)
reports that this genus is restricted to the Eastern Hemisphere, with one “‘some-
what questionable” record from Unalaska, the only previous record from the
Western Hemisphere. It is impossible to tell if this record of Anisops from Hawaii
represents a colonization or a human mediated introduction. In either event, a
single male specimen was found and preserved in alcohol. Since this specimen
was collected, others have collected additional specimens of Anisops in two hab-
itats on the island of Lanai (D. A. Polhemus, personal communication).
Notonecta indica was collected on the islands of Hawaii, Oahu, and Maui in
1991 and 1992 (Polhemus 1995). None was found on Kauai in 1992. This species
was found in natural ponds and man-made habitats (metal or concrete stock
troughs). This species was abundant on the island of Hawaii in 1991 as hundreds
were observed in several ponds. These ponds were dry in 1992, but 8 adults and
40 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
numerous nymphs were collected from another pond not observed in 1991. Five
adults and 50 nymphs were taken from two stock troughs on Oahu in 1991. One
of the troughs remained in 1992 and numerous adults were observed. Numerous
stock troughs were sampled on Maui, west and south of Haleakala National Park,
each with adults and nymphs in both 1991 and 1992.
Buenoa pallipes was collected on the islands of Hawaiiin 1991, Oahu and Maui
in 1991 and 1992, and Kauai in 1992 (Polhemus 1995). On the island of Hawaii,
this species was found with N. indica in only one pond. Even though numerous
Notonecta were found in the stock troughs on this island, no Buenoa were found
in this habitat. This species was found in two locations on Oahu, near Puu O
Mahukona Heian State Monument, and Makakilo. The solitary specimen near
the Monument was collected in a shallow muddy pool just below the road from
the Monument. On Maui, specimens were collected from numerous stock tanks.
Aquatic habitats on Kauai were sampled onlyin 1992, and only Buenoa specimens
were found.
The solitary specimen of A. kuroiwae was collected in a concrete stock trough
on Maui along with 10 specimens of N. indica. The head and pronotum are
deposited in the entomological collection of the Bishop Museum, Honolulu as a
voucher specimen. The remainder of the specimen was used to obtain DNA for
molecular systematic studies of the Notonectidae.
Acknowledgment. —Two anonymous reviewers made helpful corrections to this
manuscript. Dan Polhemus assisted with the initial identification of the Anisops
specimen, and an anonymous reviewer clarified a synonomy.
Eric Larsen, Biology Department, Villanova University, Villanova, Pennsylvania
19085-1699.
PAN-PACIFIC ENTOMOLOGIST
72(1): 41-42, (1996)
Scientific Note
NORTHERN RANGE EXTENSION FOR
HAEMATOSIPHON INODORUS (DUGES)
(HEMIPTERA: CIMICIDAE)
Herein we report the occurrence of a hematophagous ectoparasite, Haemato-
siphon inodorus (Dugés), in raptor nests and on nestlings at the Snake River Birds
of Prey National Conservation Area (NCA), near Kuna, Idaho. Commonly known
as the Mexican chicken bug, H. inodorus was previously reported south of 37°
latitude from western Oklahoma to southern California and south to central
Mexico (Usinger R. L. 1966. Monograph of Cimicidae. Horn Shafer Co., Balti-
more, Maryland). The presence of H. inodorus in southwestern Idaho (c. 43°10’
N, 116°30' W) represents a new northern latitudinal distribution and a range
extension of over 800 km from the nearest previously identified population (Red
Mountain, California).
We conducted reproductive surveys of prairie falcons (Falco mexicanus Schle-
gel) and golden eagles (Aquila chrysaetos L.) from 1991-1994. We noticed an
increase from 1991 to 1992 in the number of ectoparasitized nests and in the
degree of infestation. Because we suspected that the bugs contributed to prairie
falcon nest failure in 1992 and 1993, we collected specimens for identification
from prairie falcon nestlings, from cavities in basalt cliffs used by nesting prairie
falcons, and from an active golden eagle nest. Cimicid bugs collected from seven
nests during the 1992-94 breeding seasons were subsequently identified as H.
inodorus and are in the possession of M. E. McFadzen.
Prairie falcons, golden eagles, and other raptor species, as well as California
condors (Gymnogyps californianus Shaw), turkey vultures (Cathartes aura Weid),
and domestic fowl, are known hosts of this ectoparasite (Grubb, T. G., W. L.
Eakle & B. N. Tuggle. 1986. J. Wild. Dis. 22: 125-127).
Since 1967, biologists have noted the presence of “‘bedbugs’ in raptor nests at
the NCA (M. Kochert, personal communication). More recently, cimicid bugs
collected from prairie falcon nests at the NCA were reported as Oeciacus vicarius
(Horvath), the cliff swallow bug (Sitter, G. 1983. M.S. Thesis. Univ. Idaho. Mos-
cow). Although active cliff swallow (Hirundo pyrrhonota Vieillot) colonies were
near some of the prairie falcon nests from which we obtained bug specimens, O.
vicarius was not found in any of our samples.
Little is known about the ecological impact of H. inodorus on raptors. However,
these parasites appear to have detrimentally influenced reproductive success. High
levels of H. inodorus in raptor nests have been blamed for nestling mortality
(Platt, S. W. 1975. Wilson Bull. 87: 557; Grubb et al. 1986; McFadzen, M. E. &
J. M. Marzluff, unpublished data) and for low nestling mass and hematocrit
(McFadzen, M. E. & J. M. Marzluff, unpublished data). Additionally, hemato-
phagous ectoparasites significantly increase costs associated with reproduction
(Moller, A. P. 1993. J. Anim. Ecol. 62: 309-322), which may consequently de-
crease fitness.
42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Record. —USA. Idaho. ADA CO.: nr Kuna, June/July 1992-94, M. E. McFadzen and M. S. Vekasy,
ex. prairie falcon and golden eagle nestlings and nests.
Acknowledgment. — Thanks are extended to Greenfalk Consultants and the Rap-
tor Research and Technical Assistance Center, U.S. National Biological Service,
Boise, Idaho, for providing field support, and to W. Hansen and A. Roe, Utah
State University, Logan, and D. Baker, University of California, Davis, for iden-
tification of some cimicid specimens. C. Schaefer, University of Connecticut,
Storrs, and J. M. Marzluff, Sustainable Ecosystems Institute, Boise, Idaho, pro-
vided helpful comments on the manuscript. This note is a result of a cooperative
research project between the U.S. Bureau of Land Management and the Idaho
Army National Guard. Funding for the work was provided by the Idaho Army
National Guard through the U.S. Army Chemical Research, Development, and
Engineering Center to Greenfalk Consultants, contract # DAAD 05-90-C-0135.
Mary E. McFadzen,!* Mark S. Vekasy,!? Theresa Y. Morishita,’ and John H.
Greve,’ /Greenfalk Consultants, 8300 Gantz Ave., Boise, Idaho, 83709. ?Present
address: 441 Thatcher St., Boise, Idaho, 83702. 7Department of Preventive Med-
icine, Ohio State University, 1900 Coffey Road, Columbus, Ohio, 43210-1092.
#Department of Veterinary Pathology, Iowa State University, Ames, Iowa, 5001 1-
1250.
PAN-PACIFIC ENTOMOLOGIST
72(1): 43-47, (1996)
PROCEEDINGS OF THE PACIFIC COAST
ENTOMOLOGICAL SOCIETY, 1990
FouR HUNDRED AND SEVENTY-FOUTH MEETING
The 474th meeting of the Pacific Coast Entomological Society was held 19 January 1990 in the
Morrison Auditorium of the California Academy of Sciences with President Robert Dowell presiding.
The meeting was called to order and minutes of the 473rd meeting were read and accepted. Five new
members were proposed and accepted, 3 regular, 1 student and 1 sponsoring; Barbara Wilson, Shawn
O’Kief, Dr. Spangler, Glen Connor, and Bob Sautter.
Several announcement were made. Barbara Wilson announced that the California Department of
Health Services will have 2 vacancies to fill in their vector surveillance and control program, one in
Santa Rosa and one in Berkeley. She noted that an examination for the positions will be given no
later than June 1990, and that the state of California is an affirmative action employer. Dr. Dowell
announced that the next meeting will be on 23 February, at U.C. Berkeley. A map will appear in the
next Bits & PCES. The March meeting will be held on 16 March at U.C. Davis. The April meeting
will be held on 21 April at U.C. Berkeley and the program will be an “Old Timers Day.”
Kirby Brown showed several slides of a very attractive stellate scale, family Coccidae which arrived
in a shipment in Florida. President Dowell introduced the speaker for the evening. Dr. Jeffrey Granett,
chairman of the Department of Entomology at U.C. Davis. Dr. Granett’s presented an excellent slide
lecture titled ““Grape Phylloxera in California: No Sex in the Vineyard.” Dr. Granett’s talk covered
the pest status, biology, research and consequences of Phylloxera. Interestingly there are only females
present in California, no males. However, there are males present in other parts of the world such as
the eastern U.S. and Europe. Dr. Granett outlined two types of reproductive behavior, asexual if living
in the soil and sexual if living only on the leaves. Phylloxera is restricted to species of grapes. Under
stress the nymphs develops into the winged form and lay male and female eggs. The species native
to the eastern U.S, infects several species and forms galls on the leaves but the roots are fairly resistant.
In the European strain, the root is not resistant but the leaves are. A club shaped formation on the
roots near a feeding site is very characteristic. Galls on leaves form all over the plant and harm the
plant but do not kill it, but if the population size is large enough on the roots it will kill the plant. Dr.
Granett gave a detailed history of phylloxera’s arrival in Europe and North America and discussed
the ocurrance of two biotypes and several host races. Dr. Granett’s talk gave a interesting and ento-
mological glimpse of the trials and tribulations of the wine industry.
The meeting was adjorned at 9:10 PM. Refreshments were served in the Entomology Department
of the Academy.—L. S. Saul, Recording Secretary.
The following 36 persons were present. 29 members: K. W. Brown, F. E. Cave, D. K. Dabney, R.
V. Dowell, J. T. Doyen, J. G. Edwards, S. V. Fend, C. W. Fox, C. D. Franklin, N. E. Gershenz, D.
W. Gray, D. F. Gross, J. E. Hafernik Jr., K. S. Hagen, P. S. Johnson, B. Keh, R. L. Langston, V. F.
Lee, D. L. Mead, W. W. Middlekauff, R. L. Penrose, J. A. Powell, L. S. Saul, W. E. Savary, J. T.
Sorensen, R. E. Stecker, P. H. Sullivan, C. Y. Takahashi, R. L. Zuparko. 7 guests: E. Amdt, V. M.
Barlow, J. Carter, M. D. Hagen, K. Hobson, S. Renkes, | illegible signature.
FouR HUNDRED SEVENTY-FIFTH MEETING
The 475th meeting of the Pacific Coast Entomological Society was held on Friday, 23 February
1990, in room 159, Mulford Hall at University of California Berkeley. The meeting was called to
order at 8:02 PM with President Robert Dowell presiding. The minutes of the 474th meeting were
read and accepted. David M. Pollack was proposed and accepted for student membership.
A motion to support the movement to designate the Monarch butterfly as the national insect of the
United States was proposed and accepted. Several announcements were made. Dr. Dowell announced
that the upcoming meeting will be on 16 March at U.C. Davis. A map will appear in the next Bits &
PCES. The speaker will be Dr. David Kistner giving a talk titled “Biological Glimpses of Australia.”
An unofficial announcement was made that there will be an Acarologist position opening at CDFA.
44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Rex Dufore, member of PCES and officer ofthe United Nations in Thailand says hello. Warren Savary
announced that Dr. Martin Muma, author of 195 papers passed away 1 December 1989. Harriet
Reinhard, long time butterfly enthusiast, is convalescing after surgery at Fairfield Convalescent Hos-
pital. The Xerces Society is looking for donations to publish a 45 page article by Harriet Reinhard
on “San Francisco Butterflies, Urban Survivors.” Historical records, food plants and current status
will be included. Anyone interested in making a donation should contact Melody Mackey Allen, 1050
SW Ash Street, Portland, Oregon 97204.
At 8:20 PM, Dr. Dowell introduced the speaker for the evening, Dr. John Hafernik, Professor of
Biology at San Francisco State University and past President of the Society. Dr. Hafernik gave a
fascinating slide lecture titled ““Mating Behavior of Jschnura Damselflies.” After a brief overview of
sexual selection theory was given. Dr. Hafernik discussed the natural history of five species of Ischnura
damselflies that occur in the Bay Area, Ischnura erratica, I. perparva, I. cervula, I. denticollis, and I.
gemina. Dr. Hafernik presented his research spanning the past 10 years on the mating behavior of
Ischnura gemina. His research concerned the variability of mating success of males versus females,
the effects of coloration on mortality and mating success and the ecological interactions that affect
mating systems in this genus.
The meeting was adjourned at 9:15 PM. Refreshments were served in the Entomology Museum in
Wellman Hall, U.C. Berkeley.—L. S. Saul, Recording Secretary.
The following 32 persons were present. 23 members: P. H. Arnaud Jr., L. G. Bezark, T. S. Briggs,
K. W. Brown, H. K. Court, R. V. Dowell, C. D. Franklin, D. W. Gray, D. F. Gross, A. Hom, N. N.
Hrebtov, R. L. Langston, V. F. Lee, W. A. Maffei, G. J. Mallick, N. D. Penny, K. J. Ribardo, R. G.
Robertson, L. S. Saul, H. I. Scudder, C. Y. Takahashi, D. Ubick, B. A. Wilson. 9 guests: M. M.
Amaud, J. E. Court, J. Granett, C. Hrebtov, D. Maffei, A. M. L. Penny, W. C. Rauscher, J. M. Ribardo,
J. Robertson.
FouR HUNDRED AND SEVENTY-SIXTH MEETING
The 476th meeting of the Pacific Coast Entomological Society was held on 16 March 1990 in Room
122, Briggs Hall at the University of California, Davis. The meeting was called to order at 8:00 PM
with President Robert Dowell presiding. The minutes of the 475th meeting were read and accepted.
22 members and 11 guests attended. No new members were proposed.
Several announcements were made. Dr. Dowell announced that the position open at the California
Department of Food and Agriculture will be filled with someone with experience in biosystematics.
No new business was discussed. The next PCES meeting will be held on 21 April 1990 at the University
of California at Berkeley. The program will be ‘“‘Old Timers Day’: Changes in Entomology. Retired
entomologists will relate their early experiences as entomologists. The program portion will run from
10:00 AM to 12:30 PM. and there will be a picnic lunch on the lawn. A map will appear in Bits &
PCES. The Insect Zoo’s 11th Annual Open House will be held on Sunday, 24 June 1990 from 10:30
AM to 4:30 PM, and will be an opportunity to reach thousands of people and let them know how
wonderful, interesting and important insects are.
Dr. Dowell introduced Dr. David Kistner professor at California State University at Chico. Dr.
Kistner presented an interesting slide lecture titled “Biological Glimpses of Australia.”’ Dr. Kistner’s
primary research interest is insects associated with ants and termites and has focused his attention on
the army ant genus /nictus and its myrmecophiles. This expedition to Australia spanned four and a
half months, 18,000 kilometers and was funded by the National Science Foundation and the National
Geographic Society. The nest structure and biology of several species of termites and their associated
termitophiles was discussed. Beetles from the families Staphylinidae, Histeridae, and Adaridae can
be found in termite mounds. Dr. Kistner discussed many interesting features of termite and termi-
tophile behavior and ecology including phoresy, the use of special glands and termite mimicing larvae.
The meeting was adjourned at 9:01 PM. Refreshments were served in the Bohart Museum. —L. S.
Saul, Recording Secretary.
The following 34 persons were present. 23 members: R. M. Bohart, R. M. Brown, D. A. Carmean,
R. V. Dowell, J. G. Edwards, C. D. Franklin, D. M. Gordon, D. W. Gray, D. F. Gross, L. S. Kimsey,
D. H. Kistner, V. F. Lee, J. M. Leong, D. L. Mead, N. D. Penny, J. A. Powell, R. G. Robertson, L.
S. Saul, R. E. Somerby, J. T. Sorensen, C. Y. Takahashi, R. W. Thorp, T. L. Tyler. 11 guests: V. M.
Barlow, M. Bohart, J. Garcia, J. Granett, S. Granett, R. Longair, D. P. Muth Jr., R. D. Oliver, A. M.
L. Penny, J. Robertson, G. Spaulding.
1996 PROCEEDINGS 45
FoUR HUNDRED AND SEVENTY-SEVENTH MEETING
The 477th meeting of the Pacific Coast Entomological Society was held on 19 October 1990 in
Room 122, Briggs Hall at the University of California, Davis. The meeting was called to order at 8:05
PM with President Robert Dowell presiding. The minutes of the 476th meeting were read and accepted.
Six new regular members including David Mills, Alan Robinson, and Dr. Robert Pemberton and 2
new student members were proposed and accepted.
Dr. Dowell announced that “Old Timer’s Day” was held on 21 April 1990 at Morgan Hall on the
U. C. Berkeley campus. The event was attended by 24 people and included the following speakers:
Louie Blanc, Dr. Edwards, Dr. Dean Furhman, Dr. Harry Laidlaw, Dr. Harry Lange, Dr. Mittlecoff,
Dr. Edward Ross, Dr. Harvey Scudder, and Dr. Arthur Smith. All the speakers gave some very
interesting and entertaining insights into what it was like working in the field of entomology during
the 30’s, 40’s, 50’s to the present.
The PCES sponsored a symposium at the AAAS Pacific Division meeting held at U.C. Davis in
June 1990 entitled ““The Ecology, Taxonomy, Evolution and Importance of Insect Biotypes, Host-
plant Races and Sibling Species’’; the symposium was very well received. Daniel Gross has accepted
a position with the U.S. Food and Drug Administration in Los Angeles, hence he will be resigning as
membership chairman. Vincent Lee will become acting chairman of membership. It was announced
that several other positions were still open: recording secretary, managing secretary, and president-
elect. Harvey Scudder announced that the africanized bee was discovered in Texas. Dr. Dowell
announced that the medfly program was being wrapped up in southern California, however that
mexican, oriental, and melon fruit flies have been caught and they will be treating for gypsy moth in
San Diego and Santa Barbara Countries. Cornell style cabinets and drawers are available for purchase
from the Entomology Department of the California Academy of Sciences.
Dr. Dowell introduced the speaker for the evening, Dr. Larry Orsak. Dr. Orsak presented an
interesting slide lecture entitled ‘““How to Keep from Being Eaten in Papau New Guinea— From an
Insect’s Point of View.” After a giving an overview of some of the cultural aspects of New Guinea,
Dr. Orsak described some of the biological flora and fauna of New Guinea. Using the WAU Institute
as a base and funding and volunteers from Earthwatch a reference moth collection was established.
The defenses of moths are being studied as possible biological indicators. Dr. Orsak discussed the
concept of apparency and the multiple defenses strategies used by moths.
The meeting was adjourned at 9:35 PM. Refreshments were served in the Bohart Museum.—L. S.
Saul, Recording Secretary.
The following 39 persons were present. 22 members: R. L. Aalbu, F. G. Andrews, T. D. Cuneo, D.
C. Dailey, R. V. Dowell, J. G. Edwards, E. M. Fisher, D. W. Gray, L. S. Kimsey, C. Y. Kitayama,
V. F. Lee, W. A. Maffei, L. J. Orsak, N. D. Penny, K. J. Ribardo, R. G. Robertson, L. S. Saul, H. I.
Scudder, R. E. Stecker, C. Y. Takahashi, G. W. Ulrich, B. M. Wilk. 17 guests: V. M. Barlow, C.
Greene, D. Kitaya, D. Limburg, C. Lupcho, D. Maffei, J. C. Marston, A. M. L. Penny, J. M. Ribardo,
C. Sala, K. Schwarz, D. R. Shaffer, B. Trimble, M. Williams, R. Worth, 2 illegible signatures.
FouR HUNDRED AND SEVENTY-EIGHTH MEETING
The 478th meeting of the Pacific Coast Entomological Society was held on 16 November 1990 in
Room 101, Morgan Hall at U.C. Berkeley. The meeting was called to order at 8:10 PM with President
Robert Dowell presiding. The minutes of the 477th meeting were read and accepted. One new regular
member Dr. William Tolzer and one new student member, Mr. Stephen J. Sulatycki were proposed
and accepted.
Several announcements were made. Curtis Takahashi announced that Hugh B. Leach died on
Thursday, 15 November 1990 of Parkinson disease. He was born on 10 May 1910 and came to work
at the Academy in 1947. He area of specialty was water beetles. He retired in 1974. President Dowell
announced that the December meeting will be held on the 14th and that he will speak on western
tiger swallowtails. Vincent Lee will be acting membership chairman. Dr. Gordon Frank and Dr. Howell
Daly offered to help recruit new members to PCES. Dr. Dowell announced that the Peach fly Dacus
zonatus was found recently in California.
President Dowell introduced the evening’s speaker Dr. Richard Karban from U.C. Davis. His talk
was titled ‘Local Adaptation of Thrips to Individual Plants and the Role of Sexuality.”” Dr. Karban
discussed two questions: 1. Is a natural population of herbivores structured according to the genotype
46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
oftheir host; and 2. Does sexual reproduction allow plants to escape the evolutionary tracking oftheir
adapted herbivores. The thrip populations studied were found on the long lived host plants Erigeron
glaucus in the Bodega Bay area. These plants can live up to eighteen years and stay green throughout
the entire season but most of the biomass is underground. They are also easy to manipulate and can
be grown from cuttings. Thrips are active year round but do not move very much from plant to plant
or from rosette to rosette. They have a winged and apterous form and have a short generation time
going from adult to adult in three weeks. Males are rare in the population and there is both sexual
and asexual reproduction. The number of thrips supported by different plants varied greatly from 10-
100-1000 thrips per plant. The same population levels were found experimentally as well. It was
determined that the host clone is very important in explaining abundance of thrips rather than
interspecific competition and climatic conditions. Morphological differences seem to environmentally
produced (i.e. height, leaf size). It was concluded that the population of thrips are adapted to individuals.
The meeting was adjourned at 9:15 PM Refreshments were served in the entomology museum.—
L. S. Saul, Recording Secretary.
The following 28 persons were present. 17 members: K. W. Brown, H. V. Daly, R. V. Dowell, J.
T. Doyen, J. G. Edwards, G. W. Frankie, C. D. Franklin, D. W. Gray, V. F. Lee, S. B. Opp, N. D.
Penny, A. B. Rackett, K. J. Ribardo, R. G. Robertson, L. S. Saul, C. Y. Takahashi, B. A. Wilson. 11
guests: D. Corey, L. Culp, R. Dake, S. Griffin, R. Karban, J. Knipe, A. M. L. Penny, J. M. Ribardo,
J. Robertson, S. Spisak, T. Williamson.
FouR HUNDRED AND SEVENTY-NINTH MEETING
The 479th meeting of the Pacific Coast Entomological Society was held on 14 December 1990 in
the Goethe Room of the California Academy of Sciences. The meeting was called to order at 8:10
PM with President Robert Dowell presiding. The minutes of the 478th meeting were read, corrected
and accepted. Two new regular members Dr. Brian J. Armitage and Dr. Chad M. Murvosh were
proposed and accepted. Annual reports were given by the following committees. Acting Membership
Chairman Vincent Lee reported that in 1990 there was a total of 461 members: 343 regular, five
family, two retired, 56 student, 42 sponsoring, 4 honored and 9 life members. A total of 18 new
members joined the Society in 1990; 12 regular, 5 student and 1 sponsoring. The Publication Com-
mittee discussed new guidelines for publications. Dr. Paul Amaud reported that the Historical Com-
mittee received material containing correspondence of Dr. E. Gorton Linsley and Dr. Donald G.
Denning’s Trichoptera correspondence. This has now been stored in 35 archival boxes. Dr. Amaud,
chairman of the Historical Committee was asked to confirm that there were 264 shares of stock and
while doing so discovered correspondence from William Seymour Edwards to butterfly specialist, W.
G. Wright from 1894. Dr. Amaud on behalf of Treasurer Sandra Shanks reported that all accounts
were in order. The Society as of 30 September 1990 had $112,000 in assets, $45,000 in the C. P.
Alexander Fund and $30,000 in the Fall Memoir Fund. Dr. Norman Penny reported for Vannoy
Davis, Chairman of the Audit Committee, that all the accounts were in order. The Nominating
Committee proposed the following candidates: President-elect, Dr Norman Penny; recording Secretary,
Larry Bezark; Managing Secretary, Mr. Wesley Maffei: Membership Chairman, Mr. Curtis Takahashi;
and there were two replacements to the Publication Committee, Dr. Susan Opp and Dr. Alice Hunter.
All candidates were elected.
President Dowell reported the highlights of the Executive Board Meeting held on 14 December
1990. Seventeen years of changes have now been incorporated into the by-laws and will be published
in the January 1991 issue (Vol. 67: number 1) of The Pan-Pacific Entomologist. The Society will
support efforts to obtain additional budgetary resources through Congress for the USDA Lab. A motion
to approve moneys for exceptional speakers was accepted. It was decided that PCES would not sponsor
a symposium at the 1991 AAAS meeting in Utah but would sponsor one at the 1992 AAAS meeting
in Santa Barbara. Several announcements were made. The Society has started a journal exchange with
Revista Nicaraguense de Entomologia and it will be available for study in the Academy Library. Ron
Stecker and Keve Ribardo will be traveling in April to Copper Canyon in Mexico and requested
information on the area from any members. Several notes were given. Dr. Donald Burdick from Cal.
State Fresno demonstrated the Mideo microscopy system and showed how the system could be used
to store images of type and holotype specimens. By putting this information onto video tapes, this
information can be shared without having to send out the type specimens. Leslie Saul showed a group
slide of the “Old Timers Day” participants. Dr. Edward Smith gave a note on new discoveries of
1996 PROCEEDINGS 47
fossils in the Gilboa area of New York which were extracted by acid maceration of rock dated at 414
million years. Large arachnoid predators were found. Dr. Dowell announced the upcoming meetings
and speakers: 18 January, Dr. Robert Page; 15 February, Dr. John Alcock; and 15 March, Dr. Jerry
Powell.
President-elect Leslie Saul introduced the evening’s speaker Dr. Robert Dowell of the California
Department of Food and Agriculture. Dr. Dowell gave a fascinating presidential address titled ““Host
Plant Relations in the Western Tiger Swallowtail.” The natural history of Papilio rutulus was sum-
marized. Data on larval food preference was collected and growth and survival rates of larvae reared
on over 20 different native and exotic food plants was analyzed and reported. It was determined that
the highest survival rate was achieved when reared on Lombardy poplar. A relationship between
survivorship, developmental time and pupal weight was discovered.
The meeting was adjourned at 9:45 PM. Refreshments were served in the entomology museum.—
L. S. Saul, Recording Secretary.
The following 38 persons were preserit. 28 members: P. H. Arnaud Jr., L. G. Bezark, T. S. Briggs,
D. J. Burdick, J. S. Chinn, H. K. Court, P. R. Craig, D. K. Dabney, R. V. Dowell, J. T. Doyen, J. G.
Edwards, N. E. Gershenz, D. W. Gray, R. L. Langston, V. F. Lee, W. A. Maffei, G. J. Mallick, D. L.
Mead, N. D. Penny, K. J. Ribardo, L. S. Saul, W. E. Savary, J. T. Sorensen, G. S. Spicer, R. E. Stecker,
C. Y. Takahashi, D. Ubick, T. J. Zavortink. 10 guests: M. M. Arnaud, L. Baker, D. Bohmann, J. E.
Court, T. F. Hlavac, L. Lubin, A. M. L. Penny, W. C. Rauscher, J. M. Ribardo, R. Saint John.
PAN-PACIFIC ENTOMOLOGIST
72(1): 47-54, (1996)
PROCEEDINGS OF THE PACIFIC COAST
ENTOMOLOGICAL SOCIETY, 1991
FouUR HUNDRED AND EIGHTIETH MEETING
The 480th meeting of the Pacific Coast Entomological Society was held on 18 January 1991 in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Leslie Saul presiding, and 26 members and 11 guests in attendance.
The minutes of the meeting held 14 December 1990 were read and corrected.
The following members were elected to membership in the Society: Student Members; James E.
Baxter and Wayne R. Owen; Regular Members; John A. DeBenedictis, Andre Macedo and Thomas
O. Robbins.
Vince Lee announced an upcoming showing of insect drawings done by Debbie S. Brennan of the
Academy.
President Saul gave a pitch for membership renewals and announced a job opening for a production
supervisor for a butterfly facility, and enticed the membership to participate in an upcoming San
Francisco Zoo run by extolling the virtues of the walking stick T-shirt given away to participants.
The speaker of the evening was Dr. Robert Page of the University of California at Davis, who
summarized his research on the influence of genetics on honeybee worker behavior and organization
within the hive, with a talk entitled “Self Organization and Adaptation in Insect Societies.”
Refreshments were served in the Entomology Department Conference room.—Larry G. Bezark,
Recording Secretary.
The following 37 persons were present. (26 members): P. H. Arnaud Jr., L. G. Bezark, T. S. Briggs,
D. K. Dabney, W. A. Doolin, S. V. Fend, N. E. Gershenz, J. E. Hafernik Jr., A. Hom, B. Keh, R. L.
Langston, V. F. Lee, W. A. Maffei, G. J. Mallick, N. D. Penny, J. M. Ribardo, K. J. Ribardo, R. G.
Robertson, L. S. Saul, H. I. Scudder, E. L. Smith, R. E. Stecker, P. H. Sullivan, C. Y. Takahashi, D.
Ubick, and S. P. Welles Jr.; (11 guests) M. M. Arnaud, D. Maffei, B. Page, M. Page, R. Page, A. M.
L. Penny, W. C. Rauscher, S. Renkes, J. Robertson, A. Sternfeld-Dunn, and D. Sternfeld-Dunn.
48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
FouR HUNDRED AND EIGHTY-FIRST MEETING
The 481st meeting of the Pacific Coast Entomological Society was held on 15 February 1991 in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Leslie Saul presiding, and 36 members and 37 guests in attendance.
The minutes of the meeting held 18 January 1991 were read and approved.
The following members were elected to membership in the Society: Student Members; Paul da Silva
and Richard A. Worth; Regular Members; Rodney L. Crawford and Kingston Leong.
Ron Stecker introduced Elias Castillo of the National Geographic Society who will be leading an
upcoming expedition into Northern Mexico’s Copper Canyon.
Leslie Saul asked for recommendations for future Spring or Fall programs and workshops.
Vince Lee announced two seasonal positions with the San Mateo Mosquito Abatement District that
will be available from May through September. He also announced a journal exchange for the Japanese
“Elytra” which is not to be confused with the Spanish “Elytron” which we already receive as part of
another exchange.
President Saul described a macrovideo system, similar to the one that Don Burdick exhibited
recently, that is available at the San Francisco Zoo for various projects. She also mentioned the need
for a new recording secretary for next year and the immediate need for a refreshment chairperson.
The speaker of the evening was Dr. John Alcock of Arizona State University, who presented us
with a fascinating overview of insect mating strategies, illustrated with carpenter bee and pompilid
wasp examples in a talk entitled: ‘“Hilltopping Mating Systems of Insects: A Continuing Puzzle.”
John’s refreshing style, coupled with misleading audience participation brought everyone into the
subject. After the talk, many good questions were posed and Dr. Alcock tied up the loose ends.
Refreshments were served in the Entomology Department Conference room.—Larry G. Bezark,
Recording Secretary.
The following 73 persons were present. (36 members): J. R. Anderson, P. H. Amaud Jr., L. G.
Bezark, T. S. Briggs, K. W. Brown, P. Buickerood, R. Buickerood, D. K. Dabney, P. G. da Silva, L.
H. Davis, H. E. M. Dobson, J. G. Edwards, C. W. Fox, N. E. Gershenz, J. E. Hafernik Jr., R. L.
Langston, V. F. Lee, J. Leong, D. L. Mead, J. A. Powell, B. E. Rackett, J. M. Ribardo, K. J. Ribardo,
R. G. Robertson, L. S. Saul, W. E. Savary, H. I. Scudder, S. S. Shanks, R. E. Stecker, C. Y. Takahashi,
R. W. Thorp, D. Ubick, S. C. Williams, R. A. Worth, T. J. Zavortink, and R. L. Zuparko; (37 guests)
J. Alcock, S. E. Alcock, M. M. Arnaud, E. Amdt, V. M. Barlow, L. Bergey, V. H. P. Bueno, H.
Chilstrom, D. Corey, L. Culp, J. R. Cure, R. Dake, M. A. Garcia, J. Hauptman, K. Hobson, A.
Johnson, K. Jones, J. Knipe, C. Lijo, L. MacTague, L. McPheron, D. Miller, D. Nebenyahl, M. J.
O’Dowell, J. O’Keeffe, E. Pearson, B. Peterson, W. C. Rauscher, J. Robertson, D. Schmidt, K. Schwarz,
M. Seyeh, S. Spisak, A. L. Symors, K. Verhousek, U. L. Yee, and M. Zavortink.
FouR HUNDRED AND EIGHTY-SECOND MEETING
The 482nd meeting of the Pacific Coast Entomological Society was held on 15 March 1991 in the
Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco, with President
Leslie Saul presiding, and 46 members and 14 guests in attendance.
The minutes of the meeting held 15 February 1991 were read and approved as amended.
The following members were elected to membership in the Society: Student Members; Ms. Elizabeth
Amdt, John M. Collier and William A. Titherington; Regular Members; Mr. David J. Bohmann, Dr.
John D. Lattin, Dr. Gerhard Ringel, Dr. Thomas R. Unruh, and Mr. Josef O. C. Wiley.
Vince Lee had copies of the meeting report of the recent International Commission on Zoological
Nomenclature gathering in Maryland. A number of radical code changes have been proposed.
Paul Amaud mentioned that a series of insect drawings done by D. Brennan of the Academy were
upstairs for observation.
Larry Bezark showed the 3rd edition of the Young Entomologists Society, Resource Guide, which
lists entomological equipment, books and novelties. He also announced the availability of a newsletter
on scarabs, called (suprisingly enough) Scarabs.
President Saul discussed the upcoming 4th annual Lepidopterists’ Society meeting 1-4 August in
Tucson. She also mentioned a call for papers for the Xerces Society meeting being held 18-22 June
at the University of Wisconsin, in Madison, in conjunction with the Conservation Biology Society.
The Tropical Lepidopterology Society is still going strong with a beautifully illustrated full-color journal
and many field activities including collecting expeditions to Brazil and Africa. A field course on the
1996 PROCEEDINGS 49
Biology of Butterflies, is being held 29 June-S July near Pike’s Peak through the Colorado Outdoor
Education Center. Also mentioned was the California Native Plant Society meeting for March (which
you obviously missed, if you are hearing about it now!), and the one year moratorium on mining in
the Chiricahua Mountains of Arizona.
The speaker of the evening was Dr. Jerry A. Powell, of the University of California at Berkeley,
who summarized many years of field studies and taxonomic work on a wide variety of insects as they
related to the Antioch Dunes system, in a talk entitled “Changes in the Insect Fauna at the Antioch
Dunes During 60 Years of Human Exploitation.’ Due to habitat destruction and over SO years of
human intervention, this area has undergone catastrophic alterations and as a result more than 40%
of the insects known from these dunes have not been collected in recent times. In 1980, this area was
afforded wildlife refuge status and as a result the endangered riodinid butterfly that lives there has
increased its population. This very well organized and extensively researched presentation by Dr.
Powell, generated a lot of questions about endangered species and habitat management.
Refreshments were served in the Entomology Department Conference room.—Larry G. Bezark,
Recording Secretary.
The following 60 persons were present. (46 members): R. L. Aalbu, F. G. Andrews, P. H. Arnaud
Jr., J. F. Barthell, L. G. Bezark, F. L. Blanc, D. J. Bohmann, D. S. Brennan, T. S. Briggs, K. W. Brown,
R. M. Brown, P. Buickerood, R. Buickerood, J. R. Clopton, J. M. Collier, H. K. Court, D. K. Dabney,
P. G. da Silva, J. A. DeBenedictis, T. D. Eichlin, S. S. Ferguson, W. E. Ferguson, E. M. Fisher, N. E.
Gershenz, D. M. Gordon, J. E. Hafernik Jr. A. Hom, D. H. Kavanaugh, R. L. Langston, V. F. Lee,
W. A. Maffei, D. L. Mead, J. A. Powell, W. J. Pulawski, A. E. Rackett, J. M. Ribardo, K. J. Ribardo,
R. G. Robertson, L. S. Saul, W. E. Savary, P. H. Sullivan, C. Y. Takahashi, W. A. Titherington, D.
Ubick, S. C. Williams, and R. L. Zuparko; (14 guests) M. M. Arnaud, R. Berlin, F. Blanc, J. E. Court,
K. Davey, B. Deutsch, M. Hannaford, K. Hobson, A. K. Johnson, D. Kammerer, D. Miller, F.
Ratmelis, W. C. Rauscher, and J. Robertson.
FouUR HUNDRED AND EIGHTY-T HIRD MEETING
The 483rd meeting of the Pacific Coast Entomological Society was held on 19 April 1991 in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Leslie Saul presiding, and 19 members and 9 guests in attendance.
The minutes of the meeting held 15 March 1991 were read and approved.
The following members were elected to membership in the Society: Student Member; David S.
Guertin; Regular Members; Carl D. Barrentine, William Chapco, Barbara Deutsch, David A. Guinn,
Ian D. Hodkinson and John Lane.
Susan Opp from Cal State Hayward, introduced several students that accompanied her from the
University, Larry Bezark exhibited photographs from the Museum Dedication at San Jose State
University (8 April 1991). The Entomology Museum was dedicated to J. Gordon Edwards.
Warren Savary showed a newsletter from the Oakland Museum Nature Sound Society which de-
scribed their many recordings including those of stoneflies drumming, as well as upcoming Society
events.
The speaker of the evening was Dr. Susan Opp of Hayward State University, who presented us with
an interesting look into the mating behavior and strategies of the apple maggot fly in a talk entitled
“Polygamy in the Apple Maggot Fly: Influence on Male and Female Reproductive Success.”’ This
research focused on the variance in mating success for each sex relative to multiple matings.
Refreshments were served in the Entomology Department Conference room.—Larry G. Bezark,
Recording Secretary.
The following 28 persons were present. (19 members): P. H. Arnaud Jr., B. T. Berke, L. G. Bezark,
T. S. Briggs, D. K. Dabney, N. E. Gershenz, A. Hom, V. F. Lee, W. A. Maffei, S. B. Opp, N. D.
Penny, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, L. S. Saul, W. E. Savary, E. L. Smith, C. Y.
Takahashi, and D. Ubick; (9 guests) M. M. Arnaud, L. Culp, R. Dake, S. Griffin, J. Knipe, R. Pelleter
Jr., A. M. L. Penny, K. Reynolds, and S. Spisak.
FouR HUNDRED AND EIGHTY-FOURTH MEETING
The 484th meeting of the Pacific Coast Entomological Society was held on 17 May 1991 in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Leslie Saul presiding, and 33 members and 9 guests in attendance.
50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
The minutes of the meeting held 19 April 1991 were read and approved, as amended.
The following persons were elected to membership in the Society: Student Member; Douglas E.
Kain; Regular Members; Harry Brailovsky, Luis M. Chong L., Daniel L. Gustafson, Thomas M.
Mowry, and Robert M. Nowierski.
Warren Savary introduced Luis Salzarno from the Food and Drug Administration in San Francisco.
A report from the publication committee indicated that the cost to publish volume 66 (1990) of
the Pan-Pacific Entomologist exceeded the revenue from dues, and as a result a dues increase will be
forthcoming in January 1992.
Ed Smith announced the new “Life Through Time,” exhibit at the Academy, and recommended a
visit.
Vince Lee announced the sale of Schmitt boxes and half-unit trays, available in the Entomology
Department. He also announced that the second issue of volume 67 of the journal would soon be
delivered, and that the third issue will contain the annual dues envelope; prompt payment is always
appreciated.
Warren Savary told the members about positions with the Food and Drug Administration in San
Francisco.
President Saul announced a Unified School District open house; mentioned the San Francisco Zoo
12th Annual open house and that WTN in London was to do a piece on the Zoo. Also discussed was
a Japanese Insectorium outside Tokyo.
Tom Briggs showed slides of Banksula harvestmen; cave dwellers which have recently been found
on talus slopes on San Bruno Mountain by Darryl Ubick.
Ed Smith discussed recent articles by Jarmila Kukalova-Peck and showed drawings of an early
hemipteroid insect; and several drawings from his atlas of insect anatomy showing many structures
of primitive insects. He also showed slides of Bittacus chlorostigma and the grassy oak-woodland
habitat in which it lives in Chico’s Bidwell Park. Although the adults are numerous and commonly
collected in the Spring, no one has yet discovered the whereabouts of the larvae.
The speaker of the evening was Dr. Norman Penny of the California Academy of Sciences, who
spoke on the “Insects of the La Amistad Biosphere Reserve in Costa Rica.” In his illustrated talk,
Norm discussed field stations and diversity studies on insects and plants in the tropics, often showing
trails that simply end somewhere in the middle of the jungle at property boundaries.
Refreshments were served in the Entomology Department Conference room.—Larry G. Bezark,
Recording Secretary.
The following 42 persons were present. (33 members): R. L. Aalbu, B. T. Berke, L. G. Bezark, R.
L. Brett, T. S. Briggs, P. Buickerood, R. Buickerood, J. S. Chinn, J. M. Collier, H. K. Court, D. K.
Dabney, J. G. Edwards, F. Ennik, S. V. Fend, E. M. Fisher, N. E. Gershenz, J. E. Hafernik Jr., V. F.
Lee, W. A. Maffei, G. J. Mallick, N. D. Penny, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, R. G.
Robertson, E. S. Ross, L. S. Saul, W. E. Savary, H. I. Scudder, E. L. Smith, C. Y. Takahashi, D.
Ubick, and S. P. Welles Jr.; (9 guests) J. Arciniega, L. W. Berke, J. E. Court, S. Hauguee, A. M. L.
Penny, S. Renkes, J. Robertson, L. Solerzano, and C. Stewart.
FouR HUNDRED AND EIGHTY-FIFTH MEETING
The 485th meeting of the Pacific Coast Entomological Society was held on 20 September 1991 in
the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Leslie Saul presiding, with a whopping 57 members and 53 guests in attendance.
The minutes of the meeting held 17 May 1991 were read and approved.
The following persons were elected to membership in the Society: Student Members; Darren A.
Pollack, Carolyn E. Warren, Vonny (sometimes also known as Chaz) Barlow, Kerri A. Schwarz, and
Steve Toarmino. Regular Members; Marian J. Barksdale, Stephen H. Bullock, Leland M. Humble,
C. Don MacNeill, Michael J. Martinez, Fred Punzo and Nobuyo Yoshida.
Bob Dowell announced that the Society will sponsor a symposium entitled ‘““The Status of Native
California Arthropods,” at the AAAS meeting in Santa Barbara in June of 1992.
Leslie Saul told the members about the availability of unit trays and Cornell cabinets at the Academy;
of course for a price. She also mentioned that a butterfly book, “Butterflies of Southeast Arizona,” is
now available.
Larry Bezark announced an open house and unveiling of exhibits at the Bohart Museum of Ento-
mology, at UC Davis, on 11 October 1991, at 4:30-7:30 pm in rooms 381 & 394 of Briggs Hall.
1996 PROCEEDINGS 51
Larry Bezark gave a note on an interesting population of the primitive tabanid, Stonemyia californica
(Bigot), collected in El Dorado county, in Summer 1991. These files are not haematophagous, but are
pollen feeders. Males are dominant in collections and most specimens have a spur vein. Each of the
seven flies collected from this location lacked the spur vein, and three of the individuals were females.
The speaker of the evening was Dr. Daniel Janzen, of the University of Pennsylvania, who presented
the lecture ““How to Inventory a Tropical Country’s Entomofanua with Local Human Resources.” He
discussed preserving the biodiversity of Costa Rica by developinghuman resources, the ongoing process
of maintenance of the country’s national parks and an inventory of insects and other groups, and the
integration of the entomological data into a form that is available to and used by society. He stressed
that the collection of insects itself, is not really the end product as seen by funding agencies, but is
merely the extension of the human resource development, that made the collection possible.
Refreshments were served in the Entomology Department Conference room.—Larry G. Bezark,
Recording Secretary.
The following 110 persons were present. (57 members): R. L. Aalbu, F. G. Andrews, P. H. Arnaud
Jr., V. M. Barlow, L. G. Bezark, T. S. Briggs, K. W. Brown, P. Buickerood, R. Buickerood, J. S. Chinn,
J. R. Clopton, P. R. Craig, T. D. Cuneo, D. K. Dabney, J. A. DeBenedictis, W. A. Doolin, R. V.
Dowell, J. G. Edwards, S. V. Fend, S. S. Ferguson, W. E. Ferguson, E. M. Fisher, J. Garcia, N. E.
Gershenz, J. E. Hafernik Jr., L. P. Kite, J. Lane, V. F. Lee, J. M. Leong, W. A. Maffei, J. D. McCarty,
D. L. Mead, S. B. Opp, J. A. Powell, N. D. Penny, R. L. Penrose, A. E. Rackett, J. M. Ribardo, K.
J. Ribardo, R. G. Robertson, L. S. Saul, W. E. Savary, K. A. Schwarz, W. D. Shepard, E. L. Smith,
R. E. Stecker, C. Y. Takahashi, S. P. Taormino, R. W. Thorp, W. A. Titherington, D. Ubick, C. E.
Warren, J. S. Wasbauer, M. S. Wasbauer, S. C. Williams, B. A. Wilson, and R. A. Worth; (53 guests)
M. M. Armaud, J. D. Barackman, M. M. Barackman, C. B. Barr, M. Barzman, C. Benesh, M. H.
Chapman, L. Culp, G. Enfiajian, C. Geiger, S. Griffin, J. Guardi, J. J. Hafernik, W. Hallwachs, D.
Janzen, L. Jasinskyj, J. Y. Kim, J. Knipe, N. Lewin, D. Limburg, D. Maffei, W. May, D. Miller, R.
Myatt, M. O’Malley, B. Orr, V. T. Parker, E. Pearson, A. M. L. Penny, L. Randall, H. G. Real, S.
Renkes, K. Reynolds, J. Robertson, N. E. Robinson, R. Roche, A. Royer, R. Saint John, D. Schmidt,
A.R. Seaborg, D. M. Seaborg, A. Smyth, L. Solerzano, G. Spaulding, S. Spisak, D. Stark, A. Stroganoff,
D. Ugles, M. Vasey, N. Warnock, B. A. Zlotnick, and 2 illegible signatures.
Four HUNDRED AND EIGHTY-SIXTH MEETING
The 486th meeting of the Pacific Coast Entomological Society was held on 18 October 1991 in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco, with
Leslie Saul presiding, and 39 members and 24 guests in attendance.
The minutes of the meeting held 20 September 1991 were read and approved.
The following persons were elected to membership in the Society: Student Member; Ms. Gail M.
Getty, and Regular Members; Mr. Donald Mahoney, and Ms. Robin K. Roche
Norm Penny of the Academy, introduced Dr. Brett Ratcliffe of the University of Nebraska, who
was visiting the Academy and working on the scarab collection.
Dr. Bill Ferguson will be collecting in Chiapas, Mexico next year and solicited information from
members who had recently been to the area.
He also had in the coffee room after the meeting, an initiave sponsored by the Audubon Society
and the Sierra Club, which dealt with Forest Trees.
President Saul appointed an auditing committee, chaired by Vannoy Davis and including Paul
Arnaud. They will give their report at the annual meeting in December.
Warren Savary of the Food and Drug Administration told about a job in the Los Angeles area at
the GS 5, 7, or 9 level for an entomologist to deal with insects in stored products.
Curtis Takahashi announced that H. T. Harvey & Associates in San Jose will need a part time
worker to sort insects to family. This job is to be in Concord.
Paul Arnaud read a brief obituary about Herman Real who died on 15 October 1991. Ron Stecker
showed a couple of slides of Herman and others taken at the September 1991 meeting and mentioned
that he had a video of Herman also from the September meeting.
Leslie Saul announced that the Academy still was selling off some schmidt boxes and unit trays.
Curtis Takahashi showed a Cybister (Dytiscid beetle) from the Tar Pits in Southern California. The
specimen was purchased in a curio shop.
52 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
The speaker of the evening was Dr. Edward S. Ross, of the California Academy of Sciences, who
presented a lecture entitled ‘“‘Close Encounters Along Amazon Forest Trails.’’ He showed many insect
slides from recent and past field trips to Ecuador.
Refreshments were served in the Entomology Department Conference room.—Larry G. Bezark,
Recording Secretary.
The following 63 persons were present. (39 members): P. H. Amaud Jr., V. M. Barlow, L. G. Bezark,
T. S. Briggs, R. M. Brown, J. A. Chemsak, J. R. Clopton, D. K. Dabney, J. G. Edwards, S. S. Ferguson,
W. E. Ferguson, E. M. Fisher, M. Garcia-Vidal, N. E. Gershenz, P. S. Johnson, D. H. Kavanaugh, B.
Keh, R. L. Langston, V. F. Lee, C. D. MacNeill, T. C. MacRae, J. D. McCarty, D. L. Mead, N. D.
Penny, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, E. S. Ross, L. S. Saul, W. E.
Savary, H. I. Scudder, R. E. Stecker, C. Y. Takahashi, S. P. Taormino, C. E. Warren, J. S. Wasbauer,
M. S. Wasbauer, and S. P. Welles Jr.; (24 guests) M. M. Arnaud, L. J. Boutin, A. S. Brick, J. Cruz,
L. Cruz, B. Deutsch, D. D. Giuliani, K. S. Hom, J. Johnston, A. Jung, J. Jung, J. Jung, T. Kipping,
L. B. Mak, R. Morgan, N. Nealley, B. Ng-Jung, A. M. L. Penny, W. A. Rauscher, B. Rice, J. Robertson,
S. Simmons, L. Solerzano, and L. M. Wolcott.
FouR HUNDRED AND EIGHTY-SEVENTH MEETING
The 487th meeting of the Pacific Coast Entomological Society was held on 22 November 1991 in
the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Leslie Saul presiding, and 23 members and 15 guests in attendance.
The minutes of the meeting held 20 September 1991 were read and approved, with the following
amendments: 1) Dr. Edward S. Ross’ presentation was dedicated to the memory of Herman Real, and
2) Helen Court was also a member of the auditing committee.
The following persons were elected to membership in the Society: Student Member; Ms. Mary H.
Chapman, Mr. Yu-Feng Hsu and Mr. Michael Prentice, and Regular Members; Mr. Jeff B. Knight
and Dr. Phillip S. McNally.
Vince Lee of the Academy, presented an Entomation flier he had received from Mark F. O’Brien.
This flier described several Macintosh programs of an entomological nature that were available for
purchase. Copies of the flier were made available to those interested, in the Entomology Conference
room.
President Saul made the following announcements: First—she announced the receipt of a letter from
the Entomological Society of America about endorsing the Monarch Butterfly as the National insect.
This letter was soliciting letters of support and also had attached with it, a joint resolution listing
those organizations that already endorsed it. The letter and resolution were available for viewing in
the Entomology Conference room.
Second —President Saul briefly talked about her visit to the butterfly sanctuary at Cairns, Australia
and brought a packet of information which was available for inspection during the social hour.
Third—President Saul presented information about the Entomological Collections Network meet-
ings that were to be held in Reno, Nevada on December 7th and 8th preceding the National ESA
meetings. A program of information was also available for inspection during the social hour.
Dr. Ed Smith showed an unusual fruit (possibly a cucumber relative?) which he had found in the
street earlier in the day and asked members for assistance in its identification before presenting it to
the Botany Department.
The speaker of the evening was Dr. Robert Full, of U. C. Berkeley, who presented a lecture entitled
“Inspirations From Insects: The Design of Legged Robots.”’ Dr. Full showed slides and a video tape
about some robots that had been developed by various researchers and himself. He also discussed
the basic principles of motion and presented information that compared and modeled the walking
motion of various animals, arthropods and robots.
Refreshments were served in the Entomology Department Conference room following the main
speakers’ presentation. — Wesley A. Maffei, Acting Recording Secretary.
The following 38 persons were present. (23 members): R. L. Aalbu, P. H. Amaud Jr., T. S. Briggs,
K. W. Brown, R. M. Brown, D. J. Burdick, D. K. Dabney, W. A. Doolin, J. Garcia, P. S. Johnson,
B. Keh, R. L. Langston, V. F. Lee, W. A. Maffei, G. J. Mallick, S. B. Opp, N. D. Penny, J. M. Ribardo,
K. J. Ribardo, L. S. Saul, W. E. Savary, H. I. Scudder, and E. L. Smith; (15 guests) M. M. Amaud,
H. Barter, L. J. Boutin, L. Culp, M. Fish, R. J. Full, W. Hamersky, D. A. O’Leary, A. M. L. Penny,
W. A. Rauscher, J. Rogers, A. Shinnick, G. Spaulding, S. Spisak, and 1 illegible signature.
1996 PROCEEDINGS 53
FouR HUNDRED AND EIGHTY-EIGHTH MEETING
The 488th meeting of the Pacific Coast Entomological Society was held on 20 December 1991 in
the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Leslie Saul presiding, and 29 members and 9 guests in attendance.
The minutes of the meeting held 22 November 1991 were read and approved.
The following persons were elected to membership in the Society: Regular Members; Dr. J. A.
Slater, and Mr. Matthew Fish.
Norm Penny of the Academy introduced Margaret Thayer and Al Newton from the Field Museum
of Natural History in Chicago. Both are currently doing work on staphylinid beetles.
The following committee reports were given: Curtis Takahashi reported for the Membership Com-
mittee, that for 1991 there were 327 Regular, 45 Student, 22 Regular Retired, 6 Regular Family, 25
Sponsoring, 9 Life and 1 Honored members. The total for 1991 is 435 members. Curtis mentioned
that the Committee was soliciting additional names for Honored Member status.
Paul Arnaud reported for the Historical Committee that correspondence or photos of Hugh B. Leech,
Herman G. Real, James W. Tilden, and Thomas W. Davies. In the last few years several hundred
archival boxes of historical materials have been transferred to the archives section of the Academy.
It would be desirable to prepare a list of the entomologists’ who are represented in the archives.
Larry Bezark reported for the Nominating Committee, the following slate of candidates for 1992:
President, Norman Penny, President-Elect, Susan B. Opp, Recording Secretary, Keith Dabney, Man-
aging Secretary, Wes Maffei, Treasurer, Roberta Brett. There were no additional nominations from
the floor, and subsequently, these candidates were unanimously approved for office in the Society for
1992.
Larry Bezark reported that the Treasurer’s report was in the mail. H. Vannoy Davis had prepared
the Society’s report which would be read at the next meeting.
Helen Court gave the Auditing Committee report, stating that Mr. Davis had completed his audit
and the Society’s tax forms and that everything was in order.
Dr. Smith suggested that the Society formally thank Mr. Davis for his excellent service to the Society.
President Saul reported that the Executive Board met and that an Assistant Treasurer position was.
created. This will be an appointed not an elected position. Julie Parinas has been doing a great deal
of the bookkeeping work and the Society wishes to thank her. Julie will continue to serve the Society
in this official capacity. How the California sales tax affects the Society, and the fact that the Society
needs to upgrade it computer technology were also discussed at the Board meeting.
The Society would also like to thank Vince Lee of the Academy for his tremendous contributions
throughout the year. Although Vince currently does not hold an official position, he still performs
quite a few services for the Society.
Norm Penny announced that the Academy still had drawers and units for sale.
Vince Lee made the following announcements: A new CAS publication on Neuroptera, by Norm
Penny is now available for purchase.
Order forms were available for a Macintosh program called CLADOS version 1.0.
He also announced a summer class at Sagehen Creek given by Phil Ward of UC Davis. Entitled
“Insect Diversity and Natural History in the Sierra Nevada, this field course runs from 21 June to 26
July of 1992.
A position for an aquatic insect systematist working with Trichoptera is available at the Royal
Ontario Museum; the position starts in July of 1992.
Leslie Saul announced that Art Evans of the Los Angeles County Museum will be leading an
expedition from 1-6 February to Monarch butterfly overwintering sites.
Notes and Exhibits: Ed Smith of the Academy showed slides of primitive insects from his Atlas
and slides of the Life Through Time exhibit at CAS and briefly discussed the entire phylogeny of
insects as seen by Kukalova-Peck and others.
Larry Bezark talked briefly about a collecting trip to the rain forests of Rondonia, Brazil and had
two cases of insects from that trip. He discussed mimetic assemblages which included endomychid
and cerambycid beetles and scutellarid bugs.
President Saul then handed off the gavel to incoming President Norm Penny who introduced Leslie,
of the San Francisco Zoological Society, as the speaker of the evening. Her talk was entitled “In Search
of Insect Emissaries and Some Thoughts on Their Conservation.” Leslie discussed the need to educate
the public about the good that can be found.in insects and arthropods, the features that make insects
54 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
good emissaries and the role we can play in conserving their habitats. She showed slides of a few
recent collecting expeditions, to Borneo and Papua New Guinea.
Refreshments were served in the Entomology Department Conference room following the meeting. —
Larry G. Bezark, Recording Secretary.
The following 38 persons were present. (29 members): R. L. Aalbu, P. H. Arnaud Jr., V. M. Barlow,
L. G. Bezark, T. S. Briggs, H. K. Court, D. K. Dabney, J. G. Edwards, N. E. Gershenz, J. E. Hafernik
Jr., A. Hom, P. S. Johnson, B. Keh, R. L. Langston, V. F. Lee, W. A. Maffei, G. J. Mallick, N. D.
Penny, J. M. Ribardo, K. J. Ribardo, E. S. Ross, L. S. Saul, W. E. Savary, E. L. Smith, R. E. Stecker,
C. Y. Takahashi, S. P. Taormino, R. W. Thorp, and C. E. Warren; (9 guests) M. M. Arnaud, J. E.
Court, M. Fish, C. Greene, A. F. Newton Jr, A. M. L. Penny, W. A. Rauscher, J. Sigg, and M. K.
Thayer.
PAN-PACIFIC ENTOMOLOGIST
72(1): 55-56, (1996)
PACIFIC COAST ENTOMOLOGICAL SOCIETY
NOTES TO THE FINANCIAL STATEMENTS
YEAR ENDED SEPTEMBER 30, 1991
SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES
Accounting Method
Income and expenses are recorded by using the cash basis of accounting.
Capital Expenditures
Annual capital expenditures of $5000 or less are charged to expense.
Marketable Securities
American Telephone & Telegraph Co. and Pacific Telesis Group common stocks
are carried at market value. Increases and decreases in value are reflected in
income.
Income Tax
The Society is exempt from Federal Income and California franchise tax.
As Chairman of the Auditing Committee, and in accordance with the Society’s
bylaws, I have reviewed the financial records of the Society but have not made
an audit of them.
During the course of this review, nothing was noted which indicated any material
inaccuracy in the financial statements.
H. Vannoy Davis
Chairman of the Auditing Committee
PACIFIC COAST ENTOMOLOGICAL SOCIETY
STATEMENT OF INCOME, EXPENDITURES AND
CHANGES IN FUND BALANCES
YEARS ENDED SEPTEMBER 30, 1991 AND 1990
99 Te §e 920
Income
Des atid SUbSCrIpEONS 4m. ok, gins skew otere oo ule ele a scale Steal scely name). $ 17,375 $ 11,689
Reprints and miscellaneous ............. 0... c cece cece cee ences 18,159 12,539
[PTAC do gs a oe en Re a eT in ee Pe se SOT Te coh» 5,407 6,117
DIV IGENdS rhs 4 fe A Ree yea ht ee Se a 4 Lee en r22 616
Increase (Decrease) in value of capital stock:
American Telephone & Telegraph Company ..................... 530 (1,120)
Pacific Telesis Group
Pie ETE Se Le BA Oe hn tes ne ae eaten ee (627) (231)
WOta EIN CONISE se eee ee es OG le a DO DS A $ 41,566 $ 29,610
56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
Expenditures
Publication costs—Pan-Pacific Entomologist .................00000. $ 41,561 $ 16,818
Postage, newsletter and miscellaneous expenses .................... 1,462 1,652
Total Bxmenditres (0055 2) 2h eg ea he Oi cee tee ade a $ 43,023 $ 18,470
Increase (Decrease) in fund balances ............ 0.0.0 cc cece cece eee $ (1,457) $ 11,140
Fund balances October 1, 1990 and 1989 ......... 0.0... ccc cee eee 112,001 100,861
Fund balances September 30, 1991 and 1990 ......................... $110,544 $112,001
STATEMENT OF ASSETS
AS OF
SEPTEMBER 30, 1991 AND 1990
1991 1990
Cash in bank
OTMAET C1AL ACCOUT oe cy canine tn oy arta epic ees he tei ie eae $ 5,144 $ 6,379
Undeposited dividend checks ............ 0... cc cece eee eee es 183 319
Certificates of Deposit and Money Fund:
General Fund— Wells Fargo Bank .............. 0.000 cece cee ee eee 9,219 13,656
C. P. Alexander Fund—Capital Preservation Fund ................ 47,630 44,990
Fall Memoir Fund— Wells Fargo Bank .................. 00000 eee 34,313 32,505
Potal-cach Inibanic. = 2 h.95 oe ees rh oe, Se a aaron GS OaAS APD Le $_ 96,489 $_ 97,849
Capital Stock (at market value)
American Telephone & Telegraph Co., 80 shs. .................005. 3,000 2,470
Pacific Telesis Group, 264 shs. ..... 00.0... c ccc cece cece tees
11,055 11,682
14,055 14,152
$110,544 $112,001
See accompanying notes to the financial statements
PAN-PACIFIC ENTOMOLOGIST
72(1): 57-58, (1996)
1990 SPONSORING MEMBERS OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY
Phillip A. Adams
Robert P. Allen
William F. Barr
Richard M. Bohart
Paula & Robert Buickerood
Donald J. Burdick
Leopoldo E. Caltagirone
Arthur L. Chan
Kenneth W. Cooper
J. Gordon & Alice Edwards
William E. & Stephenie S. Ferguson
George R. Ferguson
Wayne C. Fields Jr.
John G. Franclemont
E. Eric Grissell
John E. Hafernik Jr.
Kenneth S. Hagen
Alice S. Hunter
Benjamin Keh
Robert J. Lyon
Lowe B. Mak
Robert L. Mangan
David G. Marqua
Gordon A. Marsh
Woodrow W. Middlekauff
Calvert E. Norland
Harry W. Oswald
Richard L. Penrose
Robert W. L. Potts
Jacqueline L. Robertson
Norman E. Gershenz & Leslie S. Saul
Evert I. & Marion E. Schlinger
Harvey I. Scudder
Terry N. Seeno
Frank E. Skinner
Edward L. Smith
Roy R. Snelling
Patrick H. Sullivan
Marius S. & Joanne S. Wasbauer
David B. Weissman
Barbara “A. Wilson
Sandra S. Shanks & Thomas J. Zavortink
1991 SPONSORING MEMBERS OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY
Robert P. Allen
Paula & Robert Buickerood
Arthur L. Chan
Steve H. Dreistadt
J. Gordon & Alice Edwards
John G. Franclemont
E. Eric Grissell
John E. Hafernik Jr.
Kenneth S. Hagen
Franklin T. Hovore IV
Robert L. Mangan
David G. Marqua
Gordon A. Marsh
Calvert E. Norland
Harry W. Oswald
Richard L. Penrose
Robert W. L. Potts
Norman E. Gershenz & Leslie S. Saul
Evert I. & Marion E. Schlinger
Harvey I. Scudder
Frank E. Skinner
Edward L. Smith
Roy R. Snelling
David B. Weissman
Sandra S. Shanks & Thomas J. Zavortink
58 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(1)
1992 SPONSORING MEMBERS OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY
Robert P. Allen Calvert E. Norland
Paula & Robert Buickerood Richard L. Penrose
Steve H. Dreistadt Norman E. Gershenz & Leslie S. Saul
J. Gordon & Alice Edwards Warren E. Savary
John G. Franclemont Evert I. & Marion E. Schlinger
E. Eric Grissell Harvey I. Scudder
John E. Hafernik Jr. Frank E. Skinner
Kenneth S. Hagen Edward L. Smith
Franklin T. Hovore IV Roy R. Snelling
David G. Marqua David B. Weissman
Gordon A. Marsh Sandra S. Shanks & Thomas J. Zavortink
1993 SPONSORING MEMBERS OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY
Robert P. Allen Gordon A. Marsh
Ernest Anderson Calvert E. Norland
Fred G. Andrews Harry W. Oswald
Paul Belluomini Richard L. Penrose
Paula & Robert Buickerood Norman E. Gershenz & Leslie S. Saul
Barbara Deutach Warren E. Savary
Steve H. Dreistadt Harvey I. Scudder
E. Eric Grissell Frank E. Skinner
Charles E. & Teresa Meikle Griswold Edward L. Smith
John E. Hafernik Jr. David B. Weissman
Franklin T. Hovore IV Thomas J. Zavortink
1994 SPONSORING MEMBERS OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY
Robert P. Allen Richard L. Penrose
Ernest Anderson J. Michael Poellot
Paula & Robert Buickerood Albert E. Rackett
Bryan K. Eya Norman E. Gershenz & Leslie S. Saul
E. Eric Grissell Warren E. Savary
Charles E. & Teresa Meikle Griswold Harvey I. Scudder
John E. Hafernik Jr. Frank E. Skinner
Franklin T. Hovore IV Edward L. Smith
Calvert E. Norland David B. Weissman
Harry W. Oswald Thomas J. Zavortink
PAN-PACIFIC ENTOMOLOGIST
72(1): 59-60, (1996)
The Pan-Pacific Entomologist Reviewers
Volumes 70 and 71
Aide, M. Feminella, J. McCafferty, W.
Akre, R. Fisher, E. Michener, C.
Alarie, Y. Foottit, R. Neff, J.
Alcock, J. Froeschner, R. Nishida, T.
Alexander, B. Gagne, R. O’Brien, C.
Allred, D. Garrison, R. Opp, S.
Andrews, F. Gaulet, H. Ordway, E.
Atkinson, T. Gibson, G. Page, R.
Arnaud, P. Giesbert, E. Peck, S.
Asquith, A. Gill, R. Penny, N.
Asquith, M. Goeden, R. Penrose, R.
Bari, A. Griswold, T. Perring, T.
Baumann, R. Hagen, K. Peters, W.
Beers, E. Hall, R. Powell, J. R.
Bellinger, P. Halstead, J. Pratt, H.
Berlocher, S. Hamilton, K. Prokopy, R.
Berner, L. Helms, K. Purcell, A.
Bert, T. Henneberry, T. Rees, N.
Bezark, L. Henry, T. Resh, V.
Black, W. Heyden, S. Rice, R.
Bohart, R. Holldobler, B. Robbins, R.
Borkent, A. Horn, D. Rust, M.
Brower, L. Hovore, F. Rust, R.
Cane, J. Huber, J. Sasakawa, M.
Chemsak, J. Hunter, A. Saul, L.
Cheng, L. Joern, A. Schaeffer, C.
Christiansen, K. Johnson, K. Schmid, J.
Clement, S. Jones, V. Schoenly, K.
Cole, B. Kaneshiro, K. Schuh, R.
Dahlsten, D. Kaspari, M. Schwartz, M.
Daly, H. Krombein, K. Shapiro, A.
Danforth, B. Krysan, J. Sheppard, W.
Deitrich, C. Lattin, J. Smetana, A.
DeVries, P. Lange, H. Smith, D.
Dingle, H. Lawrence, J. Sorensen, J.
Dobel, H. Leong, K. Spangler, J.
Dowell, R. Lewis, V. Starmer, W.
Doyen, J. Liquido, N. Summers, C.
Ehler, L. Lockwood, J. Tassano, T.
Erwin, T. Mackauer, M. Tauber, M.
Evanhuis, N. Markov, T. Thorpe, R.
Faulkner, D. Masner, L. Torchio, P.
60
Triplehorn, C.
Trumbo, S.
Turner, C.
Unnithan, G.
van den Berg, N.
Vaughn, C.
Vargas, R.
Vocheroth, J.
THE PAN-PACIFIC ENTOMOLOGIST
Voegtlin, D.
Walker, G.
Wagner, M.
Wasbauer, M.
Wharton, R.
Wheeler, Q.
Whitfield, J.
Williams, S.
Vol. 72(1)
Wood, D.
Wood, T.
Woodley, N.
Woolley, J.
Wrubel, R.
Zavortink, J.
Zedler, P.
Zoller, B.
PAN-PACIFIC ENTOMOLOGIST
Information for Contributors
See volume 66(1): 1-8, January 1990, for detailed general format information and the issues thereafter for examples; see below for
discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English,
but foreign language summaries are permitted. Manuscripts not meeting the format guidelines may be returned. Please maintain a copy
of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy-editing.
Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 x 11 in,
nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS
OF ANY FIGURES TO THE 8.5 x 11 IN PAGE. Number pages as: title page (page 1), abstract and key words page (page 2), text
pages (pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last.
List the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right corner. The
title must include the taxon’s designation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words; use
five to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.
Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases
underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet
1988, Dorly et al. 1989), (Burton in press) and (R. F. Tray, personal communication). For multiple papers by the same author use:
(Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith
1987: table 3).
Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194-198; if you do not have access to that
volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are
applicable. These requirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED
(INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author
of each species after its first mention.
Data Formats. — All specimen data must be cited in the journal’s locality data format. See volume 69(2), pages 196-198 for these
format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before
submitting manuscripts for which these formats are applicable.
Literature Cited. — Format examples are:
Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.
Blackman, R. L., P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics
provide some answers? pp. 233-238. In Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics
and taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985.
SPB Academic Publishing, The Hague, The Netherlands.
Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.
Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.
Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 x 181 mm while maintaining label
letter sizes of at least 1 mm; this reduction must also allow for space below the illustrations for the typeset figure captions. Authors
are strongly encouraged to provide illustrations no larger than 8.5 x 11 in for easy handling. Number figures in the order presented.
Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of
the manuscript, and (5) journal. FFGURE CAPTIONS must be on a separate, numbered page; do not attach captions to the figures.
Tables. — Keep tables to a minimum and do not reduce them. Table must be DOUBLE-SPACED THROUGHOUT and continued
on additional sheets of paper as necessary. Designate footnotes within tables by alphabetic letter.
Scientific Notes. — Notes use an abbreviated format and lack: an abstract, key words, footnotes, section headings and a Literature Cited
section. Minimal references are listed in the text in the format: (Bohart, R. M. 1989. Pan-Pacific. Entomol., 65: 156—-161.). A short
acknowledgment is permitted as a minor headed paragraph. Authors and affiliations are listed in the last, left indented paragraph of
the note with the affiliation underscored.
Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for
pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include
reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page
charge fee notice with acknowledgment of initial receipt of manuscripts.
THE PAN-PACIFIC ENTOMOLOGIST
Volume 72 January 1996 Number 1
Contents
ROSS, D. W.—Phenology of pandora moth (Lepidoptera: Saturniidae) adult emergence and
eee sc Omisiomalne com Al (Olen iia ae, as. 2ikeene. Me Bele Le SM ee J
BRUNNER, J. F.—Discovery of Colpoclypeus florus (Walker) (Hymenoptera: Eulophidae) in
APPle OnCharaasoleyy aan CMOl ia ewe ed et ee Pe
HALBERT, S. E., J. B. JOHNSON, P. L. GRAVES, P. M. MARSH & D. NELSON —<Aphidius
uzbekistanicus (Hymenoptera: Aphidiidae) established tr [Gabo ccc cceeceeceeeeeeeeeeeeeeeeeee
HALAJ, J., D. W. ROSS, R. R. MASON, T. R. TORGERSEN & A. R. MOLDENKE—
Geographic variation in arboreal spider (Araneae) communities on Douglas-fir in western
Oey CTEM ec a Wc es el ay se Me peal bm Lale dn ines ey Toned ty rc :
ZUPARKO, R. L.—Hymenoptera reared from Plagiotrochus suberi (Hymenoptera: Cynipidae)
galls in California
FOX, C. W., A. D. HARBIN & T. A. MOUSSEAU —Suitability of a non-host palo verde for
development of Stator limbatus (Horn) (Coleoptera: Bruchidae) larvae
SCIENTIFIC NOTES
GULMAHAMAD, H.— Gnathamitermes per plexus (Banks) (Isoptera: Termitidae): A nuisance
structural termite pest in southern California
LARSEN, E.—The distribution of aquatic Heteroptera (Notonectidae) in Hawaii .......0.............
McFADZEN, M. E., M. S. VEKASY, T. Y. MORISHITA & J. H. GREVE—Northern range
extension for Haematosi phon inodorus(Dugés) (Hemiptera: Cimicidae)
Pacific Coast Entomological Society, Proceeding for 1990
Pacific Coast Entomological Society, Proceeding for 1991
Pacific Coast Entomological Society, financial statement for 1990, 1990 oe ceestenceseeeee
Pacific Coast Entomological Society, Sponsoring Members 1990-1994
PAN-PACIFIC ENTOMOLOGIST REVIEWERS, Volumes 70 and 71
The
PAN-PACIFIC
ENTOMOLOGIST
Volume 72 April 1996 Number 2
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
(ISSN 0031-0603)
The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
The Pan-Pacific Entomologist
EDITORIAL BOARD
R. V. Dowell, Editor R. M. Bohart
R. L. Penrose, Associate Editor J. T. Doyen
R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.
Julieta EK Parinas, Treasurer Warren E. Savary
Published quarterly in January, April, July, and October with Society Proceed-
ings usually appearing in the October issue. All communications regarding non-
receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary;
and financial communications should be addressed to: Julieta F Parinas, Treasurer;
at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad-
emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
Application for membership in the Society and changes of address should be
addressed to: William Hamersky, Membership Committee chair, Pacific Coast
Entomological Society, Dept. of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, CA 94118-4599.
Manuscripts, proofs, and all correspondence concerning editorial matters (but
not aspects of publication charges or costs) should be sent to: Dr. Robert V.
Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture,
1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con-
tributors, and volume 66(1): 1—8, January 1990, for more detailed information.
Information on format for taxonomic manuscripts can be found in volume 69(2):
194-198. Refer inquiries for publication charges and costs to the Treasurer.
The annual dues, paid in advance, are $25.00 for regular members of the So-
ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for
institutional subscriptions or sponsoring members. Members of the Society receive
The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes
are available; see 67(1): 80 for current prices. Make checks payable to the Pacific
Coast Entomological Society.
Pacific Coast Entomological Society
OFFICERS FOR 1996
Wojciech J. Pulawski, President Vincent E Lee, Managing Secretary
Julieta E Parinas, Treasurer Stanley E. Vaughn, Recording Secretary
THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per
year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate
Park, San Francisco, CA 94118-4599. Second-class postage is paid at San Francisco, CA, and addi-
tional mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological
Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
This issue mailed 8 May 1996
The Pan-Pacific Entomologist (ISSN 0031-0603)
PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.
PAN-PACIFIC ENTOMOLOGIST
72(2): 61-69, (1996)
GEOGRAPHICAL DISTRIBUTION OF
BRACHYSTOMELLINAE
(COLLEMBOLA: NEANURIDAE)
JupItH NAJT! AND WANDA M. WEINER?
‘Laboratoire d’Entomologie et EP go du CNRS, MNHN,
45, rue Buffon, F-75005 Paris, France;
Institute of Systematics and Evolution of Animals,
Polish Academy of Sciences,
Slawkowska 17, PI-31106 Krakow, Poland.
Abstract.—The subfamily Brachystomellinae, Collembola, currently includes 14 genera occurring
mostly on the Southern Hemisphere with the sole exception of Brachystomella, which is dis-
tributed worldwide. The genera Bonetella, Salvarella and Subclavontella are endemic for Aus-
tralia. Rapoportella occurs primarily in the Neotropical region, with a single species known from
the Australian region, although Cassagnella, reported also from the southern part of South
America, is well developed in Australia. Brachystomellides and Setanodosa are represented in
the Neotropical and the Australian regions. The other five genera: Micronella, Parastomella,
Raponella, Winterella and Folsomiella occur exclusively in the Neotropical region. Probrachys-
tomellides is distributed in the Ethiopian region. Thus, both the greatest number of genera (as
many as 10), and maximum species diversity (57 species), are to be found in the Neotropical
regions.
Key Words.—Insecta, Collembola, Brachystomellinae, biogeography, zoogeographic regions
TAXONOMIC CONSIDERATIONS
The subfamily Brachystomellinae is distributed on all continents and includes
15 genera: Bonetella Stach, 1949; Brachystomella Agren, 1903; Brachystomellides
Arlé, 1959; Cassagnella Najt & Massoud, 1974; Folsomiella Bonet, 1930; Mas-
soudella (= Australella, Stach, 1949), Ellis & Bellinger, 1973; Micronella Arlé,
1959; Parastomella Rapoport & Rubio, 1968; Probrachystomellides, Weiner &
Najt, 1991; Raponella Najt, 1988; Rapoportella (= Probrachystomella Rapoport,
1962) Ellis & Bellinger, 1973; Salvarella Greenslade & Najt, 1987; Setanodosa
Salmon, 1942; Subclavontella Stach, 1949; and Winterella Massoud, 1967. A
dubious genus, Guacharia Jackson, 1927, similar to Brachystomella with seven
ocelli was distinguished on the basis of immature specimens (Type species: G.
trinitata = B. septemoculata Denis, 1931?).
We examined types of brachystomelline species from the collection of the Na-
tional Museum of Natural History in Paris, and concluded that some of the genera
contain species which are either transitional or difficult to classify as belonging
to one of the closely related genera. Thus, within one population of Brachystomella
surendrai Goto, 1961, specimens may have normal or reduced mucro. Brachys-
tomella minimucronata Palacios-Vargas & Najt, 1981 has its mucro strongly
reduced and Brachystomella cyanea (Rapoport, 1962) has the whole furca reduced
to two mamelons. Brachystomella emder Greenslade & Najt, 1987 possesses all
characters of the genus but it is the only species known so far with spiniform setae
on dens, which is typical of the genus Brachystomellides.
62 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Among the species of the genus Brachystomellides, only B. neuquensis Cassag-
nau & Rapoport, 1962, has globular maxillae with numerous teeth, as found in
the genus Brachystomella; the remaining three species have a globular maxilla
with only one or two teeth. Winterella arlesiana Massoud, 1967, has a pair of
reduced mandibles, without a molar plate. This is a primitive character, which
also occurs in Probrachystomellides nicolaii Weiner & Najt, 1991. In all other
genera the mandibles are absent.
As regards chaetotaxy, most of the genera demonstrate a distinct paurochaetosis.
Plurichaetosis is developed only in three genera: less conspicuously in Cassagnella,
abundantly in Bonetella and Salvarella.
Massoudella and Brachystomellides are identical in all generic characters, in-
cluding the presence of crosier-like structures on labium, as described by Massoud,
1967. For this reason we propose the following synonymy: Brachystomellides =
Massoudella syn. nov. Consequently, the subfamily Brachystomellinae should
contain 14 genera.
To better understand the geographical distribution of this group we have per-
formed a phylogenetic analysis based on morphological characters (to be published
separately), the results of which were applied in this work. The analysis enabled
us to distinguish two sister lineages: brachystomellian and rapoportellian groups,
each descended from a common ancestor.
Geographic Distribution.—In 1967 Massoud presented an outline of the geo-
graphic distribution for ten genera of Brachystomellinae. The principal lineage
includes five genera, Brachystomella, Folsomiella, Setanodosa, Micronella and
Winterella, and gives rise to two Neotropical lineages: one with Rapoportella, the
other with Brachystomellides. The fourth lineage, exclusively Australian, includes
three genera: Subclavontella, Bonetella and Massoudella.
Rapoport (1971) published a study of the geographical distribution of Neo-
tropical and Antarctic Collembola, considering only a few representatives of Bra-
chystomellinae. The author postulated that Folsomiella, Micronella, Parasto-
mella, Brachystomellides, Winterella and Rapoportella were endemic to the Neo-
tropical region. The occurrence of Brachystomellides neuquensis in Argentina and
Chile and the discovery of a new (although not described) species from this genus
in Peru (Winter 1962) gave evidence of the existence of contacts between the two
major stocks: Paleantarctic (relegated to the Araucanian subregion) and Neotrop-
ical (probably Afro-Brazilian or Holotropical). The genus Brachystomella was
reported to occur commonly in two regions: Palaearctic and Neotropical, but
Rapoport apparently overlooked its occurrence in the Australian region. Setan-
odosa is distributed in two regions: Australian and Neotropical. Rapoport con-
sidered the Holotropical region as the whole space between the tropics of Cancer
and Capricorn, claiming that Brachystomella contorta Denis, 1931 was widely
distributed over this vast area.
More than twenty years have passed since the publication of the two papers,
which discussed, among other things, the distribution of Brachystomellinae. New
genera and new species have since been described and the way in which their
characters are analyzed has changed and with more attention paid to chaetotaxy.
We re-examine the geographical distribution of genera and of species using the
zoogeographic division of continents by Mroczkowski (1968). However, we note
that, even today, the available data are fragmentary.
1996 NAJT & WEINER: BRACHYSTOMELLINAE DISTRIBUTION 63
Table 1. Geographic distribution of the genera Brachystomellinae. Zoogeographic regions: Nea—
Nearctic; Pal— Palearctic; Neo— Neotropical; Eth—Ethiopian; Ori— Oriental; Aus— Australian.
Genus Nea Pal Neo Eth Ori Aus
Bonetella Stach, 1949
Brachystomella Agren, 1903 + +
Brachystomellides Arlé, 1959
Cassagnella Najt & Massoud, 1974
Folsomiella Bonet, 1930
Micronella Arlé, 1959
Parastomella Rapoport & Rubio,
+++4++H+
++4++4
1968 +
Probrachystomellides Weiner &
Najt, 1991 +
Raponella Najt, 1988 +
Rapoportella Ellis & Bellinger,
1973 + +
Salvarella Greenslade & Najt,
1987 +
Setanodosa Salmon, 1942 + +
Subclavontella Stach, 1949 +
Winterella Massoud, 1967 +
ZOOGEOGRAPHY OF BRACHYSTOMELLINAE
Distribution of Genera.—The distribution of the brachystomelline genera by
geographic regions is presented in Table 1. Only Brachystomella occurs in all
regions. Folsomiella, Micronella, Parastomella, Raponella and Winterella are
known only from the Neotropical region. One genus: Probrachystomellides belongs
to the South-African subregion in the Ethiopian domain. Three genera: Bonetella,
Salvarella, and Subclavontella are Australian. The distribution of Brachystomel-
lides and Cassagnella covers two regions: the Australian and the Neotropical.
Rapoportella is principally a Neotropical genus, but it is found in the southern
part of the Nearctic region. This is probably the result of a neotropical expansion
to the north (Sonoran province, see map 1: Mroczkowski 1968) prior to the rise
of the Mexican neovolcanic chain. Rapoportella is also represented in Australia.
Setanodosa occurs in the Neotropical and the Australian. Brachystomella is the
only genus with a wide distribution; all other genera are found only south of the
Tropic of Cancer.
Distribution of Species.—The distribution of species is presented according to
two sister groups (to be published separately).
The brachystomellian group contains ten genera: Brachystomella, Setanodosa,
Micronella, Probrachystomellides, Brachystomellides, Parastomella, Bonetella,
Winterella, Subclavontella and Folsomiella.
Brachystomella includes 52 know species (Table 2). Brachystomella parvula is
a typically Palaearctic species, which is found in the Nearctic region. It is our
opinion that all specimens from other parts of the world that have been identified
as belonging to this species should be re-examined. Brachystomella contorta is
dispersed in the Neotropical, the Ethiopian, the Oriental regions and in the Ha-
walian subregion (Australian region).
The distribution of Brachystomella platensis is rather peculiar: originally de-
64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Table 2. Geographic distribution of species of the genus Brachystomella. Zoogeographic regions:
Nea—Nearctic; Pal—Palearctic, Neo—Neotropical; Eth—Ethiopian; Ori—Oriental; Aus— Australian.
Species Nea Pal Neo Eth Ori Aus
Brachystomella Agren, 1903
agrosa Wray, 1953 a
baconaoensis Gruia, 1983 +
barrerai Palacios-Vargas
& Najt, 1981 +
chilensis Rapoport
& Rubio, 1965 +
christianseni Massoud, 1965 +
coatesi Weiner & Najt, 1991 +
contorta Denis, 1931 + + + +
curvula Gisin, 1948 +
cyanea (Rapoport, 1962) +
dianae Greenslade & Najt, 1987
disputa Greenslade & Najt, 1987
emder Greenslade & Najt, 1987
fungicola Womersley, 1933
gabrielae Najt
& Palacios-Vargas, 1986 +
georgensis Weiner & Najt, 1991 +
globulosa Cassagnau & Rapoport,
1962 +
grootaerti Najt, Thibaud
& Jacquemart, 1991 +
hawaiiensis Yosii, 1965 +
heo Christiansen
& Bellinger, 1992 +
hiemalis Y osii, 1956 +
honda Christiansen
& Bellinger, 1988 +
insulae Najt & Thibaud,
1988 +
kahakai Christiansen
& Bellinger, 1992 +
kiko Christiansen
& Bellinger, 1992 +
koreana Weiner & Najt,
1985 +
mauriesi Thibaud & Massoud,
1983 +
micromucronata Palacios-
Vargas & Najt, 1981 +
momona Christiansen
& Bellinger, 1992 +
montebella Najt
& Palacios- Vargas, 1986
nana Rubio & Najt, 1979
neomexicana (Scott, 1960)
nubila Gisin, 1957 +
parvula (Schaeffer, 1896) + +
pastoralis Greenslade
& Najt, 1987 +
perraulti Thibaud
& Najt, 1993 +
++4+4+
+++
1996 NAJT & WEINER: BRACHYSTOMELLINAE DISTRIBUTION 65
Table 2. Continued.
Species Nea Pal Neo Eth Ori Aus
platensis Najt
& Massoud, 1974 + +
quadrituberculata Stach,
1964 +
ronderosi Najt, 1973 +
septemoculata Denis, 1931 +
sexoculata Massoud, 1967 +
solidaria Greenslade
& Najt, 1987 +
stachi Mills, 1934 + +
subandinensis Massoud,
1967 +
surendrai Goto, 1961 +
taxcoana Palacios-Vargas
& Najt, 1981 +
terrafolia Salmon, 1944 +
tuberculata (Wahlgren,
1906) +
ultima Greenslade
& Najt, 1987 +
unguilonga Najt
& Thibaud, 1988 +
victoriensis Izarra, 1972 +
villalobosi Cassagnau
& Rapoport, 1962 +
zapatai Najt
& Palacios-Vargas, 1986 +
scribed from an eucalyptus grove in Argentina, and never found anywhere else
in South America, this species appears to be widely distributed in Australia
(Greenslade & Najt 1987). Originating from the Australian region, 1t has been
most probably, introduced into the Neotropics. Brachystomella stachi is known
from the Nearctic and the Neotropical regions. Brachystomella surendrai is an
Oriental species. Brachystomella curvula, B. nubila, B. hiemalis, B. koreana and
B. quadrituberculata are known only from the Palaearctic region. Brachystomella
honda is marine littoral and occurs in the south Nearctic and the north Neotropical
regions.
Sixteen species originate from the Australian region: seven from Australian,
five from the Hawaiian (Brachystomella hawaiiensis, B. heo, B. kahakai, B. kiko
and B. momona), and three from the Polynesian (B. perraulti, B. insulae, B.
unguilonga) subregions. Among these 16 species Brachystomella fungicola may
represent another, probably new genus. Brachystomella terrafolia described from
the New Zelandian subregion was reported also from India (Prabhoo 1971), but
the specimens belong perhaps to a new species. Of the remaining 24 species, 22
are exclusively Neotropical and two are Ethiopian (from the South-African sub-
region).
The genus Setanodosa has 14 species (Table 3), eight belong to the Neotropical
fauna, five to the Australian and one occurs in the Ethiopian and the Australian.
66 THE PAN-PACIFIC ENTOMOLOGIST
Vol. 72(2)
Table 3. Geographic distribution of species belonging to brachystomellian group (except Brachy-
stomella sp.—see Table 2). Zoogeographic regions: Nea—Nearctic; Pal— Palearctic; Neo— Neotropical,
Eth—Ethiopian; Ori— Oriental; Aus— Australian.
Species
Setanodosa Salmon, 1942
afurcata (Womersley, 1933)
capitata (Womersley, 1930)
clavata (Schaeffer, 1897)
decemoculata (Cassagnau
& Rapoport, 1962)
fueguensis Najt, 1973
granulata (Womersley, 1935)
kanalua Christiansen
& Bellinger, 1992
occidentalis (Arlé, 1959)
peruensis Massoud, 1967
quinseta Salmon, 1944
rosasi (Bonet, 1934)
serrata Massoud, 1967
steineni (Schaeffer, 1891)
tetrabrachta Salmon, 1942
Micronella Arlé, 1959
porcus (Denis, 1933)
checayensis Massoud, 1967
Probrachystomellides Weiner & Najt,
1991
nicolaii Weiner & Najt, 1991
Brachystomellides Arlé, 1959
compositus Arlé, 1959
geniculatus (Womersley, 1934)
micro pilosus Cassagnau
& Rapoport, 1962
neuquensis Cassagnau
& Rapoport, 1962
Parastomella Rapoport & Rubio, 1968
mylodontis Rapoport
& Rubio, 1968
Bonetella Stach, 1949
terricola (Womersley, 1933)
Winterella Massoud, 1967
arlesiana Massoud, 1967
Subclavontella Stach, 1949
acantha (Womersley, 1933)
subacantha Massoud, 1967
Folsomiella Bonet, 1930
caeca (Folsom, 1927)
albida (Arlé, 1959)
intermedia (Arlé, 1939)
nothofagutalis (Rapoport
& Rubio, 1963)
polylepiana Massoud, 1967
Nea
Pal
Eth
1996 NAJT & WEINER: BRACHYSTOMELLINAE DISTRIBUTION 67
Table 4. Geographic distribution of species belonging to rapoportelian group. Zoogeographic regions:
Nea—Nearctic; Pal— Palearctic; Neo— Neotropical; Eth—Ethiopian; Ori— Oriental; Aus— Australian.
Species Nea Pal Neo Eth Ori Aus
Rapoportella Ellis & Bellinger, 1973
bonariensis (Rapoport, 1962) +
boneti Massoud, 1963 +
karta Greenslade & Najt, 1987 +
lowriei Najt, 1984 +
margaritae Najt &
Palacios-Vargas, 1986 +
mucronata Najt & Massoud,
1974 +
punillensis Izarra, 1973 +
rapoporti (Massoud, 1963) +
sergioi (Najt, 1973) +
sigwalti Najt &
Palacios-Vargas, 1986 +
yolandae (Rapoport & Mafio,
1969) +
Raponella Najt, 1988
dodecophthalma (Rapoport &
Rubio, 1963) +
Cassagnella Najt & Massoud, 1974
alba Najt & Massoud, 1974 +
anomala (Womersley, 1933) +
Salvarella Greenslade & Najt, 1987
wallacei Greenslade & Najt,
1987 ae
The only species of Micronella are exclusively Neotropical. Probrachystomellides
nicolai is monotypic, at present known only from South Africa, and thus belonging
to the Ethiopian region. Brachystomellides includes four species, three in the
Neotropical region and one in Australia. Parastomella mylodontis, described from
Mylodon Cave in Chile, is monotypic and limited to the Neotropical region
(Patagono-Andean subregion). Bonetella terricola, also a monotypic genus, is known
only from the Australian region, and Winterella arlesiana from the Neotropics.
Subclavontella is represented by two species in the Australian region. Fol/somiella,
with five species, 1s characteristic for the Central and South-American continent:
the Neotropical region.
Within the brachystomellian group, only three species of Brachystomella are
distributed over two and one in four geographical regions. Each of the remaining
77 species has its distribution restricted to a single biogeographic realm.
The rapoportellian group includes four genera: Rapoportella, Raponella, Cas-
sagnella and Salvarella (Table 4). Ten species of the genus Rapoportella are Neo-
tropical and one is Australian.
Raponella dodecophtalma is monotypic in the Neotropical region. Cassagnella
contains two species, the Patagono-Andean C. a/baand the Australian C. anomala.
This latter species, characterized by elongated maxillae with lamellae, was orig-
68 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
inally described as belonging to the genus Brachystomella but according to Wom-
ersley (1939) this classification was incorrect. Salvarella wallacei is monotypic
and known only from the Australian region. The rapoportellian group has no
representatives in the Holarctic region.
DISCUSSION AND CONCLUSIONS
Only one of the 14 brachystomelline genera, Brachystomella, has a worldwide
distribution; the occurrence of five genera is limited to the Neotropical region,
three to the Australian region, and one to the Ethiopian region. Four genera occur
in the Neotropical and the Australian regions.
Of the 98 known species of Brachystomellinae, three are from two and one
from four regions. Among the remaining 94 species, 55 are characteristic of the
Neotropical, 28 of the Australian, five of the Palaearctic, one of the Nearctic,
three of the Ethiopian, one of the Ethiopian and Australian and one of the Oriental
regions. Thus, both the greatest number of genera (as many as 10) and maximum
of species diversity (57 species) are to be found in the Neotropical region.
Collembola, a sister group of insects, are known from the mid Devonian, from
about 400 millions y.b.p. This panchronic group has survived all geological epochs
without undergoing any basic change of its initial form. Their center of origin was
doubtlessly located in Pangaea. Collembola have developed a variety of adap-
tations that allowed them not only to conquer all habitats but also invade various
continents.
We postulate that the Brachystomellinae began with a Pangaean ancestor, and
that the strongest diversification and the most intense speciation took place on
Gondwana Land beginning from an ancestor of Brachystomella type. The original
group: Brachystomella, survived unchanged on the continent of Laurasia, but
Gondwana Land, isolated and drifting away, developed a variety of climates and
habitats which allowed for an evolutionary explosion in situ. As we have shown
above, Brachystomellinae are represented by a few species in the Holarctic region
and only the genus Brachystomella. The same genus occurs throughout the Southern
Hemisphere, where it has diversified into many species and also given origin to
the proliferation of genera endemic to the Southern Hemisphere.
ACKNOWLEDGMENT
We express our sincere gratitude to Dr. J. Casevitz-Weulersse for the critical
reading of the manuscript. The work was partly supported by the grant KBN
1828/4/91 from the Polish Committee for Scientific Research to W. M. Weiner.
LITERATURE CITED
Ellis, W. N. & P. F. Bellinger. 1973. An annotated list of the generic names of Collembola (Insecta)
and their type species. Monogrn. Ned. Entomol. Veren., 7: 1-74.
Greenslade, P. & J. Najt. 1987. Collemboles Brachystomellinae de 1l’Australie I. Les genres Bra-
chystomella et Rapoportella. Ann. Soc. Entomol. France (N.S.), 23: 435-453.
Massoud, Z. 1967. Monographie des Neanuridae, Collemboles Poduromorphes 4 piéces buccales
modifiées. Biol. Am. Austr. CNRS, 3: 1-399.
Mroczkowki, M. 1968. Distribution of the Dermestidae (Coleoptera) of the world with a catalogue
of all known species. Ann. Zool., 26: 15-190.
Prabhoo, N. R. 1971. Soil and litter Collembola of South India. I—Arthropleona. Oriental Insects,
5: 1-46.
1996 NAJT & WEINER: BRACHYSTOMELLINAE DISTRIBUTION 69
Rapoport,E.H. 1971. The geographical distribution of Neotropical and Antarctic Collembola. Pacific
Insects Monogr., 25: 99-118.
Winter, C. 1962. Zur 6kologie und Taxonomie der neotropischen Bodentiere. (II). Zur Collembolen-
Fauna Perus. Zool. Jb. Syst., 90: 393-520.
Womersley, H. 1939. Primitive insects of South Australia. silverfish, springtails and their allies.
Government Printer, Adelaide.
PAN-PACIFIC ENTOMOLOGIST
72(2): 70-81, (1996)
THE NATURAL HISTORY OF NICROPHORUS NIGRITA,
A WESTERN NEARCTIC SPECIES
(COLEOPTERA: SILPHIDAE)
DEREK S. SIKES!
Department of Biology, University of California,
Santa Cruz, California 95064
Abstract. — Nicrophorus nigrita Mannerheim is an atypical Nearctic burying beetle due to its lack
of dorsal, elytral maculations. Aspects of this species’ natural history were investigated and
compared to those of Nearctic congeners. Adults from a central Californian coastal population
were found to be crepuscular and active year-round, with minimal activity during winter. The
sex ratio of wild-trapped N. nigrita was female-biased while laboratory-raised broods were slightly
male-biased. Adult male pronotal width was greater than that of females (mean + SD) (5.84 +
0.74 vs. 5.67 + 0.66). A minimum population size of 4565 individuals was calculated for Big
Creek Canyon. Analysis of mouse carcass transect data indicated that N. nigrita adults located
dead mice more successfully in moist, cool, redwood-forested canyons than in six other habitat-
types. Vertebrate scavengers, flies and ants were the most common competitors of N. nigrita for
mouse carcasses. The reproductive biology of this species differed only slightly from known
Nicrophorus biology. Carcass mass strongly predicted the mean pronotal width of the offspring
in a brood. Nicrophorus nigrita differs from Nearctic congeners in the lack of elytral maculations,
the greater length of time required to complete development from larva to adult and an apparent
lack of reproductive diapause. It only shares year-round activity with Nicrophorus mexicanus
Matthews.
Key Words.—Insecta, Nicrophorus nigrita, Silphidae, Nicrophorinae, burying beetle, carrion,
Acari, Parasitidae, Poecilochirus carabi, P. subterraneus, Histiostomatidae, Pelzneria
Approximately 85 species of Nicrophorus are described, 20 of which are found
in the New World and 15 of these are found north of Mexico (Peck & Anderson
1985). Ecological aspects of six old world and 11 new world Nicrophorus species
have been investigated (Pukowski 1933, Anderson & Peck 1985, Kozol et al.
1988, Scott & Traniello 1990, Robertson 1992, Trumbo 1994 and citations there-
in), but the bionomics of western and southern Nearctic species remains poorly
known. All the well-studied Nicrophorus species occur in regions that experience
harsh winters and the biology of species from environments lacking harsh winters,
such as the central coast of California, is poorly understood.
Nicrophorus biology, particularly reproductive behavior, is well described (Pu-
kowski 1933, Trumbo 1994, and citations therein). In brief, interspecific and
intrasexual competition occur for dominance of small (< 100 g) carcasses. Car-
casses dominated by Nicrophorus adults are buried by a male-female pair and
used as a food resource for developing offspring. The larvae develop within a
brood chamber which also houses the parents and the carcass. Both parents tend
the larvae by regurgitating food, maintaining the brood chamber and defending
the brood against predators or competing congeners. Numerous aspects of burying
beetle behavior, such as biparental care, communal breeding, brood parasitism,
1 Present Address: Department of Ecology and Evolutionary Biology, University of Connecticut,
Storrs, CT 06269.
1996 SIKES: NATURAL HISTORY OF NICROPHORUS NIGRITA El
etc. have been recently explored from a behavioral ecology perspective (Bartlett
1988; Bartlett & Ashworth 1988; Miiller et al. 1990; Scott 1990; Trumbo 1990a,
1991, 1994; Trumbo & Wilson 1993; Trumbo & Fiore 1994; Eggert & Miiller
1992; Scott & Gladstein 1993).
Nicrophorus nigrita Mannerheim, occurs along the Pacific coast, from British
Columbia to Baja California and inland to Nevada, (Anderson & Peck 1985, Peck
& Anderson 1985). It is the most common silphid in California (Miller & Peck
1979). These large, black beetles are present in most Californian ecosystems, yet
current knowledge is limited to data gathered from museum specimens.
The lack of dorsal, elytral maculations on N. nigrita adults is a species-level
trait unique among Nearctic Nicrophorus. Anderson & Peck (1986) suggest a
thermoregulatory hypothesis, supported by the evidence that melanic morphs of
Nicrophorus guttula Motschulsky, Nicrophorus defodiens Mannerheim and Nicro-
phorus investigator Zetterstedt are found in regions of reduced levels of solar
radiation, where N. nigrita is common. If N. nigrita is adapted to environmental
conditions unique for Nearctic burying beetles, as suggested by its dark coloration,
then other traits associated with this adaptation should exist. This investigation,
in addition to providing new natural history information, attempts to test this
prediction.
I investigated the hypothesis that N. nigrita displays additional traits associated
with its lack of elytral maculations, and documented this species’ life history.
During 1991 and 1992, I collected information on abundance, phenology, sex
ratio, morphology, diel periodicity, population size, carrion community species
composition, habitat preference and reproductive biology from a central Califor-
nian, coastal population of N. nigrita.
MATERIALS AND METHODS
Study Site.— Research was conducted from 19 Apr 1991 to 7 Mar 1992 at the
University of California Landels-Hill Big Creek Reserve. The reserve is located
within the Santa Lucia coastal mountain range, Monterey County, California
(36°4' N, 121°36’ W). The topography is convoluted with mountain ridges reaching
elevations above 600 m. Extreme climatic variation and steep elevational gra-
dients in the reserve are associated with many diverse communities. Twelve plant
communities exist within the reserve, with redwood (Sequoia sempervirens (D.
Don) Endlicher), oak (Quercus spp.), bay (Umbellularia californica (Hooker &
Arnott) Nuttall), and coastal sage scrub (Artemisia spp., Baccharis sp., Ceanothus
sp. et al.) being members of the more common communities (Bickford & Rich
1985).
Live Trapping. —Traps were positioned throughout the reserve to sample beetle
abundance over time and to obtain sex ratio and body size data. Eight 1 m?
hanging live-catch pitfall traps (Wilson et al. 1984), were modified by adding a
funnel inside the collecting entrance, fashioned from a 12 oz. plastic Solo (R) cup
minus its bottom, to prevent beetles from escaping. Traps were opened on 19
Apr 1991 and were rebaited at ten day intervals with 250 g of rotten chicken that
had been contained for 2—5 days at ambient temperature. Starting in mid October,
traps were rebaited at monthly intervals and trapping ceased 7 Mar 1992.
Population abundance and phenology were evaluated by recording trap catches
at ten day intervals and by marking, releasing and recapturing beetles. Captured
72 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
N. nigrita were marked by cutting a triangular piece, approximately 1 mm? in
size, from the posterior margin of an elytron (Wilson & Knollenberg 1987, Trumbo
1990b, Goldwasser et al. 1993) before releasing them at the trap site. I also
obtained phenology information from museum specimens of the California Acad-
emy of Sciences Entomology Collection in San Francisco and the Entomology
Collection of the University of California at Davis.
Sex ratio and morphology were evaluated by sexing trapped individuals and
by measuring their pronotal widths (to 0.05 mm). Sex determinations were based
on this species’ sexual dimorphism: the male’s foretarsi are wider and have more
pubescence than the female’s and the male has larger ridges above the eyes and
a larger clypeus than the female. Dried beetles obtained from the traps were
weighed to evaluate the relationship between pronotal width and mass (Bartlett
& Ashworth 1988, Scott 1990, Robertson 1992).
Diel periodicity was investigated by using a single trap located in a redwood-
forested canyon bottom, that was checked once for beetles an hour before and
once an hour after sun-up and sun-set. The data represent thirty days of obser-
vations, spanning 16 Jul to 30 Aug 1991.
Insect specimens of species not previously seen in the hanging traps were col-
lected and identified (Table 3). Only presence/absence data were recorded for
species other than N. nigrita. Voucher specimens of these species were deposited
in the Montana Entomology Collection, Montana State University. A long series
of adult N. nigrita specimens was deposited in the research collection at the
Landels-Hill Big Creek Reserve.
A minimum population size for this species within Big Creek Canyon (one of
four canyons sampled) was estimated using a sequential, Bayesian algorithm (Ga-
zey & Staley 1986, Kozol et al. 1988). To help meet assumptions of the algorithm,
data from a single trap run from 22 May to 21 Oct 1991 in Big Creek Canyon
were used.
Mouse Carcass Transect Studies. —Habitat preference was assessed by moni-
toring mouse carcass transects. Dead laboratory mice, (Mus musculus L.), thawed
10-15 h before use and weighing 15-35 g, were spaced on transects at 20 m
intervals in seven habitats (Table 4). Each mouse was attached to a flag marker
by 0.5 m of copper wire that prevented their removal by invertebrates, yet allowed
burial by beetles and subsequent carcass location. The mice were observed daily
for the first three days and every other day thereafter. I eventually recorded all
mice as being dominated, displaced or consumed. During these observations, I
hand collected representatives of the Coleoptera and Hymenoptera found on the
mouse carcasses (Table 3). Voucher specimens of these species were deposited in
the Montana Entomology Collection, Montana State University.
Laboratory Re production Studies. — Wild pairs of beetles obtained from hanging
traps were placed with freshly thawed mouse carcasses in plastic 2 liter repro-
duction chambers (RC). Each RC had a perforated lid, was filled with soil from
a redwood-forested, canyon-bottom floor and contained one pair of beetles and
a mouse carcass. The RCs were kept in a well aerated room within the reserve,
where the photoperiod and temperature fluctuated with external conditions. Lab-
oratory temperatures ranged from 13 to 20 °C. To test for seasonal reproductive
activity, wild-caught adults were placed in RCs on the following dates: 3 May, 5
Jul, 28 Jul, 24 Sep and 5 Nov 1991.
1996 SIKES: NATURAL HISTORY OF NJCROPHORUS NIGRITA 73
50
40
30
20
Abundance of Beetles /trap
10
100 158 217 275 333 27 85
Julian Day
Figure 1. Abundance of adult Nicrophorus nigrita captured in hanging pitfall traps; day 120 (1991)
= first trap capture = (30 Apr 1991); day 67 (1992) = last trap capture = (7 Mar 1992).
Quantitative data on the duration of parental care, duration of larval feeding
and the number of emerging progeny were obtained by observing activity within
the RCs daily. Fecundity data were obtained by counting the number of larvae
per brood on day 12. Sex ratios and pronotal widths of the emerging adults from
40 RCs were recorded.
Other aspects of Nicrophorus reproductive biology, such as the creation of a
brood ball and biparental care, were noted during the course of the study, but no
quantitative data were gathered. Phoretic mites were collected and identified for
comparative purposes. Mite specimens were deposited in the private collection
of B. M. O’Connor.
Reproductive Success.—I investigated the following three variables as possible
predictors of reproductive success: paternal pronotal width, maternal pronotal
width and mouse carcass size. Reproductive success was measured by two vari-
ables: the number of offspring, and the mean pronotal width of the offspring in
a brood. I also determined the relationship between offspring size and days from
burial to eclosion.
RESULTS
Live-trapping. —The abundance of N. nigrita adults steadily increased through-
out the summer but decreased sharply with the onset of cooler weather and greater
precipitation in early November (Fig. 1). However, N. nigrita adults were captured
74 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Table 1. Pronotal width of Nicrophorus nigrita (mm) captured in hanging-pitfall traps.
Sex n Mean (SD) Min. Median Max. Var.
é 862 5.84 (0.74) 3.90 5.85? 7.85 0.55
g 1022 5.67 (0.66) 3.70 5.70? 7.52 0.44
@ Males significantly larger than females, Mann-Whitney test, H = 24.27, df = 1, P = 0.000001.
throughout the winter. Specimens of N. nigrita in the California Academy of
Sciences Entomology Collection and the UC Davis Entomology Collection had
been collected during each month of the year (n = 611). In 313 days of trapping,
2349 N. nigrita adults were caught. I marked and released 1497 N. nigrita adults,
of which 123 (8.2%), were later recaptured.
The sampled, wild population of N. nigrita had a female-biased sex ratio (0.84:
1.0, n = 1884; differs significantly from 1:1 x? = 13.58, df = 1, P < 0.05). The
combined F, sex ratio of 32 laboratory reared broods was male-biased (1.08:1.0,
n = 354; differs significantly from 1:1 x? = 7.59, df= 1, P < 0.05).
The mean pronotal width of males was greater than that of females (Table 1).
Pronotal width was found to be a positive predictor of dried beetle mass (n =
214, linear correlation r = 0.65, P < 0.001, Kendall’s Tau = 0.461, P = 1.47 x
10-23),
Nicrophorus nigrita adults were most often caught at dusk, with some catches
recorded at dawn and fewer recorded during the night (Table 2). No beetles were
caught during the day (Table 2).
The agyrtid, Necrophilus hydrophiloides Guérin-Méneville, was commonly cap-
tured in the hanging pitfall traps during the winter months when silphids were
rare (Table 3).
Within Big Creek Canyon a total of 623 beetles were marked during 23 capture-
mark-release events and 83 (13%) were recaptured. I calculated a minimum pop-
ulation size of 4565 beetles (95% confidence intervals of 1832 and 5866 individ-
uals).
Mouse Carcass Transect Studies. — Fifty-eight of 90 dead mice placed on tran-
sects were displaced by vertebrate scavengers (64%). Two transects placed over
100 m from a trail or road were the least disturbed by vertebrates (Table 4). Table
3 lists Coleoptera and Hymenoptera species found associated with mouse car-
casses.
Table 2. Diel Periodicity of Nicrophorus nigrita adults caught in a hanging pitfall trap.
# Beetles
Time intervals # Days Total hrs. obs? observed>
Daytime >06:30 h <20:00 h 6 81 0
PM crepuscular >20:00 h <21:00 h 8 8 18
Night >21:00 h <05:30 h 10 85 >
AM crepuscular >05:30 h <06:30 h 6 6 3
@ Total hrs. obs. is the total number of hours during the specified time intervals that were sampled
for beetle presence.
> Beetles were most often caught at sundown (x? = 405.4, df = 3, P < 0.0001).
° Four beetles trapped on a cloudless night of full moon, 29 Aug 1991.
1996 SIKES: NATURAL HISTORY OF NICROPHORUS NIGRITA 75
Table 3. Coleoptera and Hymenoptera species caught in hanging pitfall traps baited with 280 g
rotten chicken, and on mouse carcasses in a variety of habitats (see Table 4), May through Oct 1991.
Mouse
Order, family Species carcasses Traps
COLEOPTERA
Silphidae Nicrophorus nigrita Mannerheim xX xX
Nicrophorus guttula Motschulsky xX
Agyrtidae Necrophilus hydrophiloides Guérin-Méneville xX
Staphylinidae Creophilus maxillosus (L.) xX
Dermestidae Dermestes marmoratus Say x x
Dermestes talpinus Mannerheim xX
Megatoma sp. x
Leiodidae Catops simplex Say xX
Histeridae Saprinus prob. lugens Erichson xX
Melyridae Collops sp. xX
Cleridae Necrobia rufipes (De Geer) 4
HYMENOPTERA
Vespidae Vespula vulgaris (L.) xX
Braconidae Alysia nr. alticola (Ashmead) xX
Formicidae Solenopsis xyloni McCook x
Solenopsis molesta (Say) xX
Crematogaster (mormonum Emery or coarctata Mayr) x
M onomorium ergatogyna Wheeler x
Formica moki Wheeler xX
Camponotus nr. vicinus Mayr xX
Nicrophorus nigrita only buried mice in cool, mesic, redwood habitats (Table
4). Temperature fluctuations within these habitats were less severe than habitats
found near or on ridge tops (unpubl. weather station data).
Laboratory Reproductive Studies. —Wild N. nigrita adults were reproductively
active in the laboratory from 3 May to 5 Nov 1991. The means of the number
of offspring produced from three independent rearing attempts were 15.6 (SD =
11.7, n = 3 broods), 15.3 (SD = 6, n = 8 broods) and 12.6 (SD = 5.6, n = 40
broods). Male parents remained with the brood for a mean of 11.1 days (SD =
1.2, n = 6 broods) whereas females remained with the brood for a mean of 13.1
days (SD = 2.3, n = 7 broods). Paternal residence time was not significantly
shorter than maternal (t = — 1.98, df = 9.5, P = 0.076). Together, parents tended
offspring for a mean of 10.9 days (SD = 1.5, n = 8 broods).
Larvae began pupation a mean of 12.3 days (SD = 1.2, n = 3 broods) after the
adults had been placed with a dead mouse (initiated 3 May 1991). Approximately
80 days elapsed between placement of adults with a dead mouse and the emergence
of offspring from their pupal chambers (mean = 79.9, SD = 6.6, n = 10 broods,
initiated 28 Jul 1991).
Features of N. nigrita biology consistent with other known Nicrophorus spp.
include: construction of a brood chamber and a brood “‘ball’’; deposition of anal
compounds onto the carcass purported to inhibit growth of bacteria and fungi
(Pukowski 1933, Halffter et al. 1983); departure of the male parent prior to the
female parent; regurgitation of food to the larvae by the parents; stridulation by
the parents while tending the young; three instars of larval development; and the
76 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Table 4. The proportion of mouse carcasses displaced, dominated or consumed by vertebrates or
saprophagous arthropods in 10 locations, over 10 days, Aug and Sep 1991.
Approx. N. Vespula
Habitat elevation n nigrita Vertebrates Flies Ants vulgaris Other
rwd?/mesic W sl 90 m 10 0.10 0.10° 0.80
rwd/mesic N sl 90 m 10 0.10 0.90
rwd/mesic N sl 90 m 10 0.90 0.10
rwd/mesic N sl 90 m 5 1.00
rwd/mesic N sl 150 m 11 0.40 0.60
rwd/xeric S sl 90 m 10 0.70 0.10 0.204
grass/xeric hltp 300 m 10 0.40 0.40¢ 0.20
oak/xeric hltpf 480 m 5 1.00
Pinus/xeric S sl 678 m 9 0.89 0.11
coast scrub N sl 330 m 10 1.00
mean %: 6 69 16 6 1 Ps
a4 rwd = redwood.
> sl = slope.
© Transect laid >100 m from trail or road.
4 Solenopsis xyloni.
© Solenopsis molesta.
f hltp = hilltop.
presence of phoretic mites and nematodes. The nematodes occasionally bred to
enormous population levels on carcasses in the laboratory causing the beetles to
abandon the carcasses—a previously unreported competitive interaction.
Three species of phoretic mites taken from N. nigrita adults were identified as
Poecilochirus carabi Canestrini (Parasitidae), Poecilochirus subterraneus (Mueller)
(Parasitidae) and Pel/zneria sp. (Histiostomatidae). These mites were all removed
from adult beetles and were deutonymphs.
Re productive Success. —No adult offspring were produced by 8 (20%) of 40 RCs
begun on 28 Jul 1991. Within the 32 remaining broods, a total of 465 larvae were
counted on day 12 after initiation. In the predator-free, laboratory environment,
354 (76%) of these larvae survived to eclosion. A significant, positive relationship
was found between days to eclosion and pronotal width of emerging adults (n =
354, r? = 0.17, P < 0.01).
Mouse carcass mass predicted the pronotal widths of emerging adults but pa-
ternal and maternal size did not. Mouse carcass mass (using mouse tissue parcels
weighing 2.75—42 g) was significantly related to mean pronotal width of the off-
spring in a brood (n = 37 broods, r? = 0.419, P < 0.0001; Fig. 2). Carcass mass
was a weak positive predictor of the number of larvae found on day 12 (n = 37
broods, r? = 0.104, P = 0.052) but not a predictor of the number of emerging
adult offspring (n = 37 broods, r? = 0.0039, P = 0.71).
DISCUSSION
Live-trapping & Mouse Carcass Transect Studies.—The phenologies of N. ni-
grita and its hypothesized sister species, Nicrophorus mexicanus Matthews (Peck
& Anderson 1985), are unique among the documented phenologies of Nearctic
Nicrophorus because adults of these species remain active in low numbers during
1996 SIKES: NATURAL HISTORY OF NICROPHORUS NIGRITA 77
7.2
e
e e
‘= e
E. 6.6 " e e
oO $ e
© 6.0 bs
re) ea .
SG ©
AS
= e e
— &
g 5.5 A e a
5 : °
e
c
o @
o 4.9
= a
e
4.3
0 5 10 15 20 25 30 35 40 45
Mouse Carcass Mass (g)
Figure 2._ Relationship between mouse carcass mass and the mean pronotal width of the offspring
in a brood for Nicrophorus nigrita, (n = 37 broods, Kendall’s Tau = 0.42, P = 0.000265; linear
correlation: r = 0.648, P < 0.001, pronotal width = 0.0407 (carcass mass) + 5.29).
winter months (Terron et al. 1991). These two species are found in regions that
generally lack cold winters, a factor related to their year-round activity. Species
in the southern United States, such as Nicrophorus carolinus Linnaeus, may also
remain active during the winter months due to the mild winters of those regions.
Peck & Anderson (1985) and Anderson & Peck (1985) state that N. mexicanus
and N. nigrita adults are most active during fall, winter and spring—a finding not
corroborated by this investigation. Terron et al. (1991) document N. mexicanus
as being most active during the summer, and least active during the winter, which
is consistent with my observations of N. nigrita.
Terron et al. (1991), studying N. mexicanus, Trumbo (1990a), studying Nicro-
phorus orbicollis Say and Wilson & Knollenberg (1984), studying N. orbicollis and
N. defodiens found female-biased sex ratios at pitfall traps. Trumbo (1990a) found
1:1 sex ratios for Nicrophorus pustulatus Herschel and Nicrophorus tomemtosus
Weber, whereas male-biased sex ratios were found in Palearctic Nicrophorus spp.
studied by Easton (1979) and Springett (1967). Wilson & Knollenberg (1984) and
Trumbo (1990a) both reported laboratory raised broods were non-biased sug-
gesting intrinsic and/or extrinsic factors skew the sex ratios in N. mexicanus, N.
orbicollis, N. tomentosus and N. nigrita.
This study indicates N. nigrita adults are crepuscular (Table 2), whereas N.
nigrita had been reported to be a nocturnal species based on data from black light
captures (Peck & Kaulbars 1987). The method used here, provides data more
78 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
representative of natural behavior than black-light data (Wilson et al. 1984).
However, I recorded activity during a brightly moon-lit night (Table 2), suggesting
these beetles may prolong their period of activity under such conditions.
Nicrophorus defodiens is active crepuscularly in Michigan (Wilson et al. 1984),
whereas N. tomentosus is diurnal and two other species, N. orbicollis and Nicro-
phorus sayi LaPorte, were documented to be strictly nocturnal. My data support
the nocturnal and crepuscular activity pattern more commonly found in this genus.
The low proportion of transect mouse carcasses dominated by N. nigrita adults
(Table 4) may result from a number of causes. Vertebrate scavengers, most likely
foxes, consumed the majority of dead mice placed in the field. Similar results,
with vertebrate scavengers removing from 60% to 100% of small carcasses, are
not uncommon (Putman 1976, 1983). Nicrophorus nigrita adults were more suc-
cessful in using carrion in areas where vertebrate scavengers have less impact as
seen in the two transects located more than 100 m from a trail or road (Table 4).
Other arthropod species may outcompete N. nigrita for carrion within the study
region (Table 3). Itis unclear whether N. nigrita prefers to breed on large carcasses,
as some congeners have been shown to do (e.g. N. pustulatus; Peck 1986, Rob-
ertson 1992, Trumbo & Wilson 1993). If shown to occur, this would help explain
the scarcity of adults on small carcasses in the field.
Laboratory Reproductive Studies.—The reproductive biology of N. nigrita is
similar to other Nicrophorus. The greatest difference found, in addition to the
apparent lack of reproductive diapause, involved the time required to complete
development. Robertson (1992) reports that Canadian N. pustulatus complete
development from larva to adult in 22-27 d. Scott & Traniello (1990) report N.
orbicollis from New Hampshire require approximately 44 d from burial ofa carcass
to eclosion. Halffter et al. (1983), studying N. mexicanus, found development was
completed from larva to adult in 39 days. These records are significantly shorter
than the corresponding 73-85 d development time required for N. nigrita. How-
ever, these traits may be population-level adaptations, or simple plastic (non-
fixed) responses to the temperature of the environments or laboratories in which
the species were studied, rather than fixed, species-level traits. Robertson (1992),
Scott & Traniello (1990), and Halffter et al. (1983) did not state at what temper-
ature the larvae developed, but all three of these studies were conducted within
laboratories, which generally range from 15 to 25° C.
Thereproductive phenologies of many Nearctic Nicrophorus are known (Wilson
et al. 1984, Scott & Traniello 1990, Anderson & Peck 1985). Nicrophorus nigrita
displays a less constrained reproductive schedule than those of congeners that are
tightly associated with the seasonal changes present in the eastern and northern
Nearctic. Nicrophorus nigrita females are able to reproduce within a month of
eclosion (unpublished data) and no evidence of reproductive diapause was found,
although reproduction in the field during the months of December to April has
yet to be demonstrated.
The reproductive success of N. nigritais consistent with that of other congeners.
Nicrophorus orbicollis shows no correlation between female size and offspring
number (Scott & Traniello 1990), but does show a strong correlation between
carcass size and total brood mass (Scott & Traniello 1990, Robertson 1992,
Trumbo 1994). I found similar relationships for N. nigrita.
Scott & Traniello (1990) found a significant negative correlation between brood
1996 SIKES: NATURAL HISTORY OF NICROPHORUS NIGRITA 79
size and days to eclosion for N. orbicollis. This is consistent with my findings
showing offspring size to be positively correlated with days to eclosion; brood size
and offspring size have been shown to be inversely related (Scott & Traniello
1990).
The departure of the male parent prior to the female has been reported from
other Nicrophorus (Bartlett 1988; Miiller & Eggert 1989; Scott 1990; Scott &
Traniello 1990; Trumbo 1991). I did not establish that male N. nigrita adults
abandon their offspring prior to females, however, this is most likely a result of
small sample sizes rather than strong evidence against the pattern of early male
departure. Scott & Gladstein (1993) present a predictive model and analysis of
the sociobiological ramifications of the duration of paternal care in burying beetles.
The geographic and phylogenetic distance (Peck & Anderson 1985) between N.
nigrita and the more well-known eastern species predicted differences were likely
to be found in the phoretic mite fauna of N. nigrita. Of the three mite species
found on N. nigrita, P. carabi is Holarctically distributed and the most commonly
reported phoretic associate of Nicrophorus spp. (Brown & Wilson 1992). Poe-
cilochirus subterraneus is not documented to occur in the Nearctic but is wide-
spread in the Palearctic (B. M. O’Connor, personal communication). There are
no described Nearctic species within the genus Pelzneria (B. M. O’Connor, per-
sonal communication). The presence of a Palearctic species of phoretic mite (P.
subterraneus) on N. nigrita indicates a possible biogeographic Asian connection.
The diversity and complexity of the phoretic fauna found on Nicrophorus spp.
has attracted attention (Wilson 1983, Wilson & Knollenberg 1987, Schwarz &
Miiller 1992, Richter 1993) but many avenues of ecological and systematic re-
search remain open (Brown & Wilson 1992).
In conclusion, I suggest that the autapomorphic aspects of N. nigrita’s biology
are associated with this species’ thermoregulatory adaptation to survival in regions
of cool but relatively constant temperatures. This hypothesis was first proposed
by Anderson & Peck (1986) for the melanic forms of N. guttula, N. investigator
and N. defodiens found predominantly along the pacific coast. The autapomorphic
aspects of N. nigrita are the continual activity throughout the winter months
(possible synapomorphy with N. mexicanus), the prolonged time span to complete
development from larva to adult, the apparent lack of seasonally-fixed reproduc-
tive activity and the absence of bright elytral maculations. Future studies inves-
tigating behavioral and ecological correlates of Nicrophorus thermoregulation are
needed to test this hypothesis.
ACKNOWLEDGMENT
Iam grateful to J. T. Smiley for providing advice, supplies and help throughout
this study. I thank M. E. Sikes for her assistance in the field and encouragement,
D. S. Wilson, S. T. Trumbo and S. B. Peck for sharing information regarding
Nicrophorus, the collection managers N. Penny (CAS) and S. Heydon (UCDC)
for their assistance, reviewers R. S. Miller, M. A. Ivie, J. T. Smiley, D. K. Le-
tourneau, C. Seibert, and two anonymous reviewers whose input improved the
manuscript, P. Ward (Formicidae), S. Heydon (Braconidae), D. Carmean (Ves-
pidae), B. O’Connor (Acari) and M. A. Ivie (Coleoptera) for their identifications
and S. V. and A. Sikes and F. Arias for their technical support. Financial support
80 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
was provided by F. F. Bliss, J. T. Smiley, S. V. and S. B. Sikes and The Department
of Biology, University of California, Santa Cruz. This research was carried out
as part of a Senior Thesis in Biology at UCSC. This is contribution J-3007 of the
Montana Agricultural Experiment Station.
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4: 332-339.
Trumbo, S. T. & A.J. Fiore. 1994. Interspecific competition and the evolution ofcommunal breeding
in burying beetles. Am. Midl. Nat., 131: 169-174.
Trumbo, S. T. 1994. Interspecific competition, brood parasitism, and the evolution of biparental
cooperation in burying beetles. Oikos, 69: 241-249.
Wilson, D. S. 1983. The effect of population structure on the evolution of mutualism: a field test
involving burying beetles and their phoretic mites. Am. Nat., 121: 851-870.
Wilson, D.S. & W. G. Knollenberg. 1984. Food discrimination and ovarian development in burying
beetles (Coleoptera: Silphidae: Nicrophorus). Ann. Entomol. Soc. Am., 77: 165-170.
Wilson, D. S., W. G. Knollenberg & J. Fudge. 1984. Species packing and temperature dependent
competition among burying beetles (Silphidae: Nicrophorus). Ecol. Entomol., 9: 205-216.
Wilson, D. S. & W. G. Knollenberg. 1987. Adaptive indirect effects: the fitness of burying beetles
with and without their phoretic mites. Evol. Ecol., 1: 139-159.
PAN-PACIFIC ENTOMOLOGIST
72(2): 82-88, (1996)
A COMPARISON OF THE SEASONAL ACTIVITY OF
PTEROSTICHUS BEETLES (COLEOPTERA: CARABIDAE)
IN A COMMERCIAL APPLE ORCHARD IN
SONOMA COUNTY, CALIFORNIA
Eric W. RIppICcK! AND Nick J. MILLS
Laboratory of Biological Control,
Department of Environmental Science, Policy and Management,
University of California, Berkeley, California 94720
Abstract.— Adults of the predatory Pterostichus beetles were trapped in a commercial orchard
for two consecutive apple-growing seasons. Pterostichus (Dysidius) lustrans LeConte, Pterostichus
(Poecilus) cursitor LeConte, and two Pterostichus (Hypherpes) species, were occasionally more
active on the soil surface in plots under organic management of insect pests of apple, than in
plots under conventional management. But, the mean number of individuals captured in rep-
licated plots, between the two management schemes, was not significantly different on any
collection date. Adults of P. /ustrans, the predominant species of Pterostichus inhabiting the
orchard, were most active on the soil surface during May, June, and July 1991. In the 1992
season, P. /ustrans were most active during June and July, P. (Hypherpes) spp. were most active
during June, and P. cursitor were most active during July. The activity of Pterostichus beetles
may coincide with the time interval in which fifth-instar Cydia pomonella (L.) (codling moth)
larvae wander on the ground prior to pupation.
Key Words.—Insecta, Coleoptera, Carabidae, Pterostichus, predators
Carabid beetles are major components of the predator assemblages in managed
and unmanaged ecosystems (Kulman 1974, Luff 1987). Many species are predators
of insect pests in agroecosystems (Allen 1979, Clark et al. 1994). For example, in
apple orchards, adult carabids are important predators of apple maggot, Rhagoletis
pomonella (Walsh) (Allen & Hagley 1990) and codling moth, Cydia pomonella
(L.) (Hagley et al. 1982, Hagley & Allen 1988). Several Pterostichus species are
the most effective carabid predators of the codling moth (Riddick & Mills 1994).
However, carabid predation alone cannot reduce codling moth populations to a
level sufficient to prevent economic damage to fruit and eliminate the use of
insecticides.
Previous research has determined the effect of organophosphate insecticides on
carabid populations in orchards. Phosmet sprays reduced the numbers of Amara
spp. and Harpalus affinis (Schrank) adults captured in pitfall traps in an apple
orchard in Ontario, Canada; but not the number of Prerostichus melanarius (Illiger)
adults captured (Hagley et al. 1980). In the laboratory, phosmet killed Amara
spp. and H. affinis, but not P. melanarius (Hagley et al. 1980).
The use of a formulation of codling moth granulosis virus, a microbial insec-
ticide, was correlated with enhanced surface activity of Harpalus pensylvanicus
DeGeer adults in July, but not with the activity of Pterostichus (Hypherpes) sp.
adults in an apple orchard in Contra Costa County, California (Riddick & Mills
1 Current Address: Department of Entomology, 1300 Symons Hall, University of Maryland, College
Park, Maryland 20742, USA.
1996 RIDDICK & MILLS: CARABID SEASONAL ACTIVITY 83
1995). This suggests that some carabid species can become more useful in orchards
that are treated with microbial insecticides for the suppresion of insect pests.
The objective of this research was to compare the seasonal activity of Pteros-
tichus beetles in replicated plots ofa commercial apple orchard under conventional
or organic apple production. It is essential to determine the activity patterns of
the Pterostichus species so that their potential as control agents of insects pests
can be evaluated.
MATERIALS AND METHODS
The study site was a commercial apple orchard (Jewell Ranch) located in the
coastal region of northern California (Sonoma County), near Sebastopol, a major
apple growing district. The orchard was dry-farmed and contained standard trees
of Gravenstein, Red, and Golden Delicious varieties. The 6.5 ha experimental
section of this ~20 ha orchard was surrounded by non-experimental sections of
trees, which were treated with organophosphate insecticides, on three sides and a
residence on the remaining side. The experimental section was subdivided into
eight plots, which were originally designated in 1989 to monitor the effect of various
insecticides on the codling moth. Each 0.81 ha (2 acre) plot was comprised of 9
rows of trees and each containing ~20 trees. The average distance between trees
in a row was 7.3 m, and the average distance from the adjacent row was 6.4 m.
Details of the pest management practices in the commercial orchard have been
reported elsewhere (Vossen et al. 1994). During the 1991 and 1992 apple growing
seasons, synthetic organophosphate or granulosis virus insecticides (designed to
suppress the neonate stage of C. pomonella), were sprayed onto trees in replicated
plots. Conventional plots contained trees that were sprayed with organophosphate
insecticides and herbicides, whereas organic plots contained trees that were sprayed
with microbial insecticides and in which mating disruption pheromones (in dis-
pensers) were used.
Each treatment (conventional or organic) plot was replicated four times for a
total of eight 0.81 ha plots. Both treatments were systematically alternated within
the experimental section of the orchard such that the first replicate plot of the
conventional treatment was proceeded by the first replicate plot of the organic
treatment. Thus, four replicate plots of the conventional treatment were alternated,
spatially, with four replicate plots of the organic treatment. Although this design
lacked control plots, it provided a realistic method of comparing the activity of
carabid beetles in a commercial setting.
Insecticide treatments were applied during C. pomonella egg-hatch periods,
predicted from degree-day models (Pickel et al. 1986). Two to three generations
of C. pomonella per season have been common in northern California. In the
1991 season, chlorpyrifos (Lorsban®) was sprayed in the conventional plots on
14 May at a rate of 2.8 kg/ha and on 7 June at a rate of 3.4 kg/ha, and phosmet
(Imidan®) was sprayed on 9 July at a rate of 5.75 kg/ha. In the organic plots,
codling moth granulosis virus (CMGV/UCB.87) was sprayed on 11, 29 May; 13,
20, 27 July; and on 3, 10, 17, 24 August in applications of 1.5 x 10!* granular
inclusion bodies per ha, plus 0.45 liters/ha of NuFilmP sunscreen, and 5.7 kg/ha
of DriFlo molasses. Each formulation was combined with 378.5 liters of tap water,
then applied with a fan air-blast sprayer. In the 1992 season, chlorpyrifos was
sprayed again in the identical conventional plots of the previous season on 23
84 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
April (rate of 3.4 kg/ha) and phosmet on 2 June (rate of 5.75 kg/ha). Codling
moth granulosis virus was sprayed in the identical organic plots on 28 April and
15 May as described above.
Early in the season 1991-1992, the herbicide paraquat was used to suppress
weed growth at the base of the tree trunks in the conventional plots, but a hand
hoe was used to remove the weeds in the organic plots. Mating disruption pher-
omones in dispensers (Isomate C) were positioned on trees in the organic and
conventional plots on 23 Mar 1992 and again on 8 Jun 1992 at a rate of 1000
per ha.
The soil type in the commercial orchard is a Goldridge fine sandy-loam. The
soil was disked, and the remaining resident vegetation was removed at the be-
ginning of the apple growing season, in mid- to late April. After the end of the
season in October or November, the soil was cultivated and a cover crop of bell
beans (Vicia faba L.) and vetch (Vicia spp.) was seeded in all plots.
The seasonal activity of adult Pterostichus beetles inhabiting the commercial
orchard was compared between the treatment plots, with pitfall traps. Pitfall
trapping has become a standard technique for sampling carabids (Morrill 1975,
Halsall & Wratten 1988). Trap catch may estimate the activity of adult carabids
on the soil surface as well as reflect the density of carabid populations (Hokkanen
& Holopainen 1986). Traps were plastic cups (473 ml), with a 9 cm diameter
opening, which were sunk into the ground so that the rim was flush with the soil
surface. Leaf litter within 20 cm of the perimeter of each trap was removed and
the soil smoothed to prevent the litter from impeding the movement of carabids
around the traps (Greenslade 1964, Powell et al. 1985). Traps were filled to the
one-quarter mark with a solution of water and liquid detergent, which reduced
the surface tension of the water causing captured beetles to sink to the bottom of
the trap. A preservative was not used in the traps because of the risk that it would
alter the catch and sex ratio of the trapped species (Holopainen 1992).
Six pitfall traps were positioned in the central row of trees in each of the eight
plots in the 1991 season. The distance between traps was not recorded, but traps
were two to three trees apart within the same row. Traps were in place for con-
secutive days during each of five sampling periods in 1991, 30 April—-7 May, 4—
11 June, 3-9 July, 30 July-9 August, and 4-11 September. In the 1992 season,
three traps were positioned in the identical tree rows of the previous season. Traps
were in place for consecutive days during each of five sampling periods, 11-14
May, 5-8 June, 7-10 July, 28-31 July, and 18-21 August. Samples were collected
on the last day of each sampling period.
Trapped beetles of the genus Pterostichus were sorted to species or species groups
in the laboratory. Pterostichus beetles were selected because of their potential as
effective predators of codling moth (Riddick & Mills 1994). The species counted
were Pterostichus (Dysidius) lustrans LeConte, Pterostichus (Poecilus) cursitor
LeConte, and two Pterostichus (Hypherpes) species, namely, Pterostichus califor-
nicus (Dejean) and Pterostichus castanipes (Ménétriés) which are morphologically
similar, and not readily distinguishable at the time that this research was under-
taken. Pterostichus californicus and P. castanipes adults have been collected in
California only. Pterostichus cursitor adults have been collected in California and
Oregon, and P. /ustrans adults have been collected in British Columbia, south to
California and east to New Mexico and Colorado (Bousquet & Larochelle 1993).
1996 RIDDICK & MILLS: CARABID SEASONAL ACTIVITY 85
Trap data were converted to mean number of Pterostichus species per tree row
per trap per day in each treatment plot. All means were log-transformed before
subjection to the analysis of variance (ANOVA). Treatment means were consid-
ered significant if P < 0.05. All statistical analyses were performed with Stat-
graphics (STSC) software. Voucher specimens are located at the Laboratory of
Biological Control, University of California, Berkeley, and the Department of
Entomology, University of Maryland, College Park.
RESULTS
A total of 577 adult carabids were captured in pitfall traps in the 1991 apple
growing season. Pterostichus lustrans represented 22.2%, Pterostichus cursitor rep-
resented 11.8%, and the two Pterostichus (Hypherpes) species represented a com-
bined 11.3% of the total adult carabids. In the 1992 season, a total of 638 adult
carabids were captured, and P. /ustrans represented 10.3%, P. cursitor, 6.0%; and
P. (Hypherpes) spp., 2.3%.
The seasonal activity of all adult Pterostichus species was compared between
the conventional and organic treatment plots. There was no significant difference
in total beetle activity on the soil surface on any collection date between treatments
in the 1991 or 1992 season (Tables 1 and 2). Individually, P. /ustrans was most
active during early May, June, and July, but no clear pattern was discernable for
P. cursitor or P. (Hypherpes) spp. (Table 1). In 1992, P. ustrans adults were most
active during early June and early July, whereas P. cursitor adults were most active
during early July, and P. (Hypherpes) spp. adults were most active during early
June (Table 2).
DISCUSSION
During the first two sampling periods (7 May, 11 June) in the 1991 season, P.
lustrans adults were almost significantly more active in the organic plots than in
the conventional plots of the commercial orchard. Apparently, a number of these
beetles avoided contact with chlorpyrifos, phosmet or paraquat on the ground in
the conventional plots. Chlorpyrifos has been shown to have no significant effect
on the activity of Pterostichus spp. adults in treated fields of grass in comparison
to untreated fields (Asteraki et al. 1992) but it is toxic to P. melanarius and
Pterostichus chalcites Say adults (Bale et al. 1992, Reed et al. 1992). Phosmet did
not reduce the activity of P. melanarius in an apple orchard, nor did it kill P.
melanarius in the laboratory (Hagley et al. 1980).
Paraquat was applied in the conventional plots, but not in the organic plots.
Brust (1990) found that the activity of Pterostichus sp. adults was reduced for up
to 28 d after paraquat had been sprayed.
Codling moth granulosis virus and mating disruption pheromone did not appear
to affect Pterostichus beetle activity. The seasonal activity of P. (Hypherpes) spp.
adults in an organic managed section of a commercial apple orchard in Contra
Costa County was not altered by these alternative methods of suppressing codling
moth (Riddick & Mills 1995). In this study a complete randomized design was
used and each treatment (formulations of granulosis virus, Bacillus thuringiensis
Berliner plus oil, oil alone, no-spray control) was replicated three times. In the
present study, a no-spray treatment was not allowable. Consequently, these data
86
Table 1.
THE PAN-PACIFIC ENTOMOLOGIST
Mean (+SEM) no. of beetles per tree row per trap per day
Vol. 72(2)
Seasonal activity of Pterostichus beetles in organic and conventional plots on collection
dates in the commercial orchard, 1991 season.
Treatment 7 May 11 June 9 July 9 Aug. 11 Sept.
P. lustrans
Organic 0.24 (0.09) 0.17 (0.07) 0.12 (0.04) 0.02 (0.02) 0.01 (0.01)
Conven. 0.06 (0.03) 0.01 (0.01) 0.07 (0.05) 0.02 (0.01) 0.01 (0.01)
F; Pa 4.20; 0.09 4.99; 0.77 0.87; 0.40 0.48; 0.52 0.00; 1.00
P. cursitor
Organic 0.10 (0.04) 0.03 (0.02) 0.06 (0.03) 0.08 (0.03) 0.00 (0.00)
Conven. 0.02 (0.01) 0.01 (0.01) 0.03 (0.01) 0.05 (0.05) 0.00 (0.00)
F; P 3.71; 0.10 0.48; 0.52 0.90; 0.39 0.40; 0.55 —;-
P. (Hypherpes) spp.
Organic 0.01 (0.01) 0.09 (0.07) 0.10 (0.04) 0.03 (0.01) 0.05 (0.01)
Conven. 0.00 (0.00) 0.03 (0.01) 0.03 (0.03) 0.01 (0.01) 0.03 (0.02)
F; P 3.00; 0.13 0.74; 0.43 2.32; 0.18 2.07; 0.20 0.82; 0.41
@ Degrees of freedom = 1, 6; and 8 plots were sampled on each collection date. —; — = F and P
values not reported because no beetles were captured.
cannot reveal whether Prerostichus activity in unsprayed plots differed from that
observed in the conventional or organic managed plots.
Pterostichus lustrans adults were the most numerous of the Prerostichus beetles
in the orchard. The adults were most active on the soil surface during May, June,
and July 1991, and during June and July of 1992. Similar activity was observed
for P. lustrans in a nearby orchard that was not sprayed with pesticides (Riddick,
unpublished data). This pronounced activity may correlate with an increased
availability of suitable prey. The activity of Pterostichus beetles may coincide
with the time that C. pomonella larvae are wandering on the ground in search of
pupation sites. First generation larvae leave fruit during May or June in orchards
in California (Pickel et al. 1986), and thereafter become vulnerable to predation
by Pterostichus beetles on the ground.
The nightly rate of predation of tethered C. pomonella fifth-instar larvae by
carabids was 60% in early June in an organic managed block of an apple orchard
of semi-dwarf trees in Brentwood, California (Riddick & Mills 1994). Pterostichus
(Hypherpes) sp. (possibly P. californicus) adults were one of the most active carabid
species on the soil surface during this time interval. (Note that P. /ustrans and P.
cursitor were very rare in this Brentwood orchard.) When Pterostichus beetles are
active and abundant, their predation may help reduce the density of the upcoming
second generation of C. pomonella in modern orchards. But this assumption is
valid only when Pterostichus adults readily locate and capture unrestricted fifth-
instars on the ground.
The reduced activity of P. /ustrans in August in the Sebastopol orchard may
have resulted from inadequate moisture and intolerable temperatures at the soil
surface. The climate in the Sebastopol region is Mediterranean, characterized by
hot, dry summers and mild, wet winters (Altieri & Schmidt 1986). The lack of
ideal conditions for activity may have influenced the adults to remain beneath
the surface. In contrast, the reduced activity in August could have resulted from
1996 RIDDICK & MILLS: CARABID SEASONAL ACTIVITY 87
Table 2. Seasonal activity of Pterostichus beetles in organic and conventional plots on collection
dates in the commercial orchard, 1992 season.
Mean (+SEM) no. of beetles per tree row per trap per day
Treatment 14 May 8 June 10 July 31 July 21 Aug.
P. lustrans
Organic 0.03 (0.03) 0.64 (0.39) 0.47 (0.11) 0.05 (0.03) 0.00 (0.00)
Conven. 0.14 (0.05) 0.00 (0.00) 0.22 (0.08) 0.03 (0.03) 0.00 (0.00)
F; Pa 3.38; 0.12 3.77; 0.10 3.44; 0.11 0.43; 0.54 ——
P. cursitor
Organic 0.00 (0.00) 0.03 (0.03) 0.58 (0.19) 0.05 (0.03) 0.00 (0.00)
Conven. 0.00 (0.00) 0.00 (0.00) 0.25 (0.08) 0.03 (0.03) 0.08 (0.05)
F; P —— 1.00; 0.36 2.27; 0.18 0.43; 0.54 2.51; 0.16
P. (Hypherpes) spp.
Organic 0.05 (0.05) 0.25 (0.08) 0.03 (0.03) 0.00 (0.00) 0.00 (0.00)
Conven. 0.00 (0.00) 0.08 (0.05) 0.05 (0.05) 0.00 (0.00) 0.00 (0.00)
F; P 1.00; 0.36 2.63; 0.16 0.18; 0.69 -—;-— —3-
4 Degrees of freedom = 1, 6; and 8 plots were sampled on each collection date. —; — = F and P
values not reported because no beetles were captured.
increased growth of native vegetation in the commercial orchard. Although the
vegetation was sparse in all plots, its presence could have obstructed or altered
the speed of movement of P. /ustrans, especially if these beetles investigated the
base of plants when searching for potential prey.
In conclusion, although the activity of Pterostichus beetles was not significantly
different between the conventional and organic treatments at any sampling period,
the spraying of conventional pesticides must be minimized in the spring, the
season of heightened activity of these carabids.
ACKNOWLEDGMENT
We thank Sue Blodgett, former IPM Advisor, North Coast Counties, and Lucia
Varela, current IPM Advisor, North Coast Counties (University of California
Cooperative Extension, Santa Rosa, California) for providing information on the
spray schedules in the orchard; David Kavanaugh (Department of Entomology,
California Academy of Sciences, San Francisco) for his assistance with the carabid
identifications; and George Jewell, Jr. for allowing us to collect insects on his
farm. This research was supported in part by a Mentored Research Fellowship
(from the Graduate Division, University of California at Berkeley) awarded to
EWR.
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PAN-PACIFIC ENTOMOLOGIST
72(2): 89-91, (1996)
EUBRANCHIPUS BUND YI FORBES
(ANOSTRACA; CRUSTACEA), A NEW
RECORD FROM CALIFORNIA
D. CHRISTOPHER ROGERS
Jones and Stokes Associates, Incorporated,
2600 ““V”’ Street, Sacramento California 95818-1914
Abstract.— A new distributional record for Eubranchipus bundyi Forbes, (Anostraca), from Cal-
ifornia is reported along with a review of the previously known distribution. The two other
Eubranchipus species found in California are discussed.
Key Words. —Crustacea, fairy shrimp, Anostraca, Eubranchipus, disjunct, Eubranchipus bundyi
Belk & Brtek (1995) report nine species of the anostracan genus Eubranchipus
from North America. Eng et al. (1990) reported the first records of Eubranchipus
in California when they detailed the occurrence in the state of two species. This
report adds a third species of this genus, Eubranchipus bundyi Forbes to the list
of fairy shrimps known to occur in California.
In May 1990, I was sampling vernal pools in Siskiyou County in northern
California and on 8 May 1990, collected Eubranchipus bundyi from Grass Lake,
(T.44N, R.3W, S.22 and 23), on the north side of Highway 97, approximately 35
km northeast of the community of Weed. Grass Lake lies in the extreme end of
an eastern arm of the Shasta Valley.
Grass Lake covers approximately 3 square km, is nearly 3 km long and 1.5 km
wide. In places, the pool is 0.5 m deep. The pool is at an altitude of 1370 m and
is surrounded by coniferous forest. The pool contained sedges and perennial
grasses at the time I collected E. bundyi.
Grass Lake is formed in Esro Silt loam. This soil has very poor drainage. Water
levels in this pool are seasonally astatic, and fed by snow-melt runoff. The pool
is surrounded by soils containing high amounts of granite and lava rock.
Water quality data gathered by the California Department of Water Resources
for the central part of Siskiyou County in the 1950s (unpublished) showed the
surface water in Shasta Valley annual lakes with pH values greater than 9.0 and
electrical conductivity reaching a maximum extreme of 12,420 umhos/cm. So-
dium and chloride ion concentrations were greater than 2700 mg/L, and these
small basins had greatly elevated aluminum and boron levels. Additionally, due
to concentration through evaporation, basins such as Grass Lake have a high
mineral salt content. Grass Lake is also noted for having a tiger salamander
(Ambystoma tigrinum) Green population.
Eubranchipus bundyi occurred in areas of the pool that were sparsely vegetated,
and 30+ cm deep. They swam actively, but always several cm below the surface.
Clasping pairs were observed. No temperature data were taken. Mature adults
were taken with an aquarium dip net and fixed in 100% ethyl alcohol before being
transferred to a 70% ethanol solution.
The specimens were identified using Belk (1975) and Pennak (1978), and were
90 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
verified by Dr. Belk. Specimens are deposited at the California Academy of Science
(CASIZ #103358).
Eubranchipus bundyi is known from New Hampshire, Vermont, Illinois, Min-
nesota, South Dakota, and Nebraska in the central and eastern United States,
with unspecified reports from New York, Ohio, Indiana, Michigan, and Wiscon-
sin. In Canada, collections have been made in Quebec, Ontario, Manitoba, Al-
berta, and Yukon, with unspecified reports from Saskatchewan. In the western
United States, collections were made from Arizona, Utah, Wyoming, Washington,
and Alaska (Belk, personal communication, Belk 1975, Donald 1983, Hartland-
Rowe 1967, Daborn 1976, Forbes 1876, Maynard and Ross 1975). Further in-
vestigation may reveal populations in Oregon.
Eubranchipus bundyi is generally found in snow-melt-filled pools in coniferous
areas and appears to be more abundant in years when higher-than-average rain
occurs during the preceding year (Donald 1983).
Daborn (1976) and Broch (1965) report that upon hatching, the first free-
swimming stage of E. bundyi has the rudiments of the first nine or 10 swimming
legs. Daborn believes the first instar stages pass in the cyst. After eclosion, the
metanauplius undergoes a period of rapid development, completing the juvenile
stages in 12 days.
Belk (1977) reports temperature tolerances for E. bundyi; after acclimating
adults at 13.5° C for 19-27 hours, he found an LD50 for males at 29° C and an
LDSO for females at 32° C.
Eubranchipus bundyi is reported to co-occur on occasion with Eubranchipus
intricatus Hartland-Rowe, E. ornatus Holmes, Branchinecta lindahli Packard, B.
mackini Dexter, and B. paludosa Muller (Donald 1983, Hartland-Rowe 1967,
Maynard & Ross 1975).
Eubranchipus bundyi can be seperated from the other two recorded Californian
Eubranchipus by the apex of the distal segment of the second antennae of the
males, which in E. bundyi is trifid (whereas in E. serratus and E. oregonus it is
bifid or truncate). Eubranchipus bundyi can be distinguished from all other North
American Eubranchipus by the labrum, which has a large anteriorly produced
boss or knob between the bases of the second antennae (Belk 1975, Hartland-
Rowe 1967, Forbes 1876).
Eng et al. (1990) first reported the occurrence of Eubranchipus species (E.
oregonus Creaser and E. serratus Forbes) from the northern extremes of California.
Eubranchipus oregonus was recorded from a single locality along Deetz Road,
1.7 km, west of Interstate 5, nearly 8 km south of the community of Weed in
Siskiyou County. This is approximately 44 km southwest of the E. bundyi locality
at Grass Lake. The Deetz Road locale is at an altitude of 1770 m in the Cascade
Mountain Range. Like Grass Lake, this pool is surrounded by coniferous forest,
but the soils are clays in a wet meadow. The vernal pool is seasonally and annually
astatic. Other populations occur in western Oregon through Washington to British
Columbia, with disjunct populations in Oklahoma (Belk 1975, Eng et al. 1990,
Creaser 1930).
Eubranchipus serratus is also known from a single California locality, at McCoy
Flat, in Lassen County (Eng et al. 1990). This site is nearly 20 km from Susanville,
and nearly 6 km from Eagle Lake. McCoy flat is situated on the south side of
Highway 44 between Lassen Park and Susanville at an elevation of 1650 m, in
1996 ROGERS: EUBRANCHIPUS BUNDYI IN CALIFORNIA 91
the southeastern end of the Cascade Mountain Range. This pool is also fed by
snow melt and is ringed with conifers.
I attempted to collect E. serratus from this site in July 1990 (the previous E.
serratus collections from this site were made in December), and I found only
Streptocephalus sealii Ryder.
Belk (1977) measured temperature LDSOs for Arizona E. serratus of 30° C for
males and 32° C for females, and had eggs hatched at 5° C.
Outside California, E. serratus is known from British Columbia, Washington,
Oregon, Arizona, Illinois, Indiana, Kansas, Nebraska, Wyoming, Maryland, and
Virginia (Belk 1975 and 1977, Eng et al. 1990).
Eubranchipus bundyi is the twenty-third anostracan species recorded from Cal-
ifornia. This is one of ten species discovered in California in the last four years
(Eng et al. 1990, Fugate 1993, Thiery & Fugate 1994; one species has yet to be
described). The northern extremes are the least explored areas of California for
anostracans, and other locales, if not species, will certainly be found.
ACKNOWLEDGMENTS
The author would like to thank Denton Belk for all his help in reviewing both
this paper and the specimens, Clyde Eriksen for review and trouble shooting and
Cynthia Casanova for editing, reediting and patience.
LITERATURE CITED
Belk, D. 1975. Key to the Anostraca (Fairy Shrimps) of North America. Southwest. Natural., 20:
91-103.
Belk, D. 1977. Zoogeography of the Arizona fairy shrimps, (Crustacea: Anostraca). Ariz. Acad. of
Sci., 12: 70-78.
Belk, D. & J. Brtek. 1995. Checklist of the Anostraca. pp. 315-354. In Belk, D., H. J. Dumont &
G. Maier (eds.). Hydrobiologia; studies on large branchiopod biology and aquaculture, II. Vol.
298: 1-5.
Broch, E. S. 1965. Mechanism of adaption of the fairy shrimp Chirocephalus bundyi Forbes to the
temporary pond. Cornell Univ. Agric. Exp. Stn., Memoir 392.
Creaser, E. P. 1930. Revision of the phyllopod genus Eubranchipus, with the description of a new
species. Occas. Papers Mus. of Zool., Univ. Mich., No. 208.
Daborn, G. R. 1976. The life cycle of Eubranchipus bundyi Forbes (Crustacea: Anostraca) in a
temporary vernal pond of Alberta. Can. J. of Zool., 54: 193-201.
Donald, D. B. 1983. Erratic occurrences of anostracans in a temporary pond: colonization and
extinction, or adaption to variations in annual weather? Can. J. Zool., 61: 1492-1498.
Eng, L., D. Belk, and C. H. Eriksen. 1990. Californian Anostraca: distribution, habitat, and status.
J. Crust. Biol., 10: 247-277.
Forbes, S. A. 1876. List of Illinois Crustacea, with descriptions of new species. Bull. Ill. Muse. Natur.
Hist., 1: 25.
Fugate, M. 1993. Branchinecta sandiegonensis, a new species of fairy shrimp (Crustacea: Anostraca)
from western North America. Proc. Biol. Soc. Wash., 106: 296-304.
Hartland-Rowe, R. 1967. Eubranchipus intricatus N. SP., a widely distributed North American fairy
shrimp with a note on its ecology. Can. J. Zool., 45: 663-666.
Maynard, S.S.&S.V.Romney. 1975. The occurrence of four new anostracan (Crustacaea) phyllopods
in Utah, multispecies records, and some notes on their ecology. Utah Acad. Proc., Vol. 52,
Part 2: 7-11.
Pennak, R.W. 1978. Eubranchiopoda (fairy, tadpole and clam shrimps). Jn Fresh-water invertebrates
of the United States, 2nd ed., No. 44-368. Wiley, New York.
Thiery, A. & M. Fugate. 1994. A new American fairy shrimp, Linderiella santarosae (Crustacea:
Anostraca: Linderiellidae), from vernal pools of California, U.S.A. Proc. Biol. Soc. Wash., 107:
641-656.
PAN-PACIFIC ENTOMOLOGIST
72(2): 92-101, (1996)
THE MEXICAN AND CENTRAL AMERICAN SPECIES OF
LOPHOSTIGMA MICKEL, INCLUDING A NEW SPECIES,
NEW DISTRIBUTION RECORDS, AND
TAXONOMIC NOTES FOR THE GENUS
(HYMENOPTERA: MUTILLIDAE)
ROBERTO A. CAMBRA T. AND DIOMEDES QUINTERO A.
Museo de Invertebrados “‘G. B. Fairchild,’’ Estafeta Universitaria,
Universidad de Panama, Republica de Panama;
(DQA) Smithsonian Tropical Research Institute,
Unit 0948, APO AA 34002-0948
Abstract.—We present taxonomic comments on the genus Lophostigma Mickel, and a key for
the species from Mexico and Central America, including the new species, L. grisselli Cambra
& Quintero from Mexico, the northernmost distribution record for this Neotropical genus. Lo-
phostigma lebasi (Mickel), is synonymized with L. cincta (du Buysson). New distribution records
are given for: L. cincta (du Buysson), Costa Rica and Ecuador; L. subgracilis (Cameron), Mex-
ico, Guatemala and Costa Rica; L. acanthophora (Dalla Torre), Pert; L. caenodonta (Cameron),
Brazil; and L. cayennensis (André), Venezuela and Brazil.
Key Words.—Insecta, Mutillidae, Lophostigma, key, Lophomutilla, Mexico, Central and South
America, synonymy, taxonomy
The genus Lophostigma Mickel (1952) was established for nine sphaeropthal-
mine species, known only from females, distributed from the Province of Chiriqui
(8°50' N), in the Republic of Panama, to the northern half of South America
(above 10° S). Casal (1963) described an additional species, L. seabrai Casal,
from Nova Teutonia, Santa Catarina, Brazil (27°03' S), disjunct from the rest.
Females of Lophostigma range from 6 to 12 mm in length and are relatively rare
in collections. Casal (1963) confirmed this when he remarked that he had seen
only two specimens of Lophostigma out of some ten thousand specimens of Neo-
tropical Sphaeropthalminae studied.
During a recent visit to the Natural History Museum, London, R.A.C. examined
three specimens of Lophostigma from Mexico, and recognized two of them as
belonging to an undescribed species. The new species, with the northernmost
distributional record (21°15’ N) for the genus, is here described and illustrated.
In addition, we recently examined seven additional specimens of Lophostigma
from Costa Rica and one from Mexico. The Costa Rican specimens helped us to
understand and unravel the marked integumental color variations of L. subgracilis
and L. lebasi.
We have diligently amassed 53 specimens of Lophostigma that permit us to
present new distribution records for five species, discuss the affinities of the genus,
intraspecific variation, and to provide a key to the three species known from
Mexico and Central America: grisselli, cincta and subgracilis. We suspect syn-
onymies of the following species: L. seabrai Casal, 1963 with L. alopha Mickel,
1952; L. cayennensis (André), 1906, with L. iracunda (Cresson), 1902 (see Tax-
onomic Notes).
Format for the description of the new species follows Mickel (1952). Two new
1996 CAMBRA & QUINTERO: MEXICAN AND C. AMER. LOPHOSTIGMA 93
specific characters are used here: the outline of the thorax in lateral view (Figs.
3, 5), and the ratio of the distance from the hind margin of the eye to the posterior
angle of genal carina (EMGC) to the greatest diameter of the eye (EGD), or
EMGC/EGD (Fig. 6). Scanning electron microscopy was done with a JEOL model
JSM-5300LV SEM.
Depository Abbreviations.—British Museum of Natural History BM (NH); U.S.
Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM);
Universidad Central de Venezuela, Maracay IZAUCV); Snow Entomological
Museum, University of Kansas (SMUK); Instituto Nacional de Pesquisas da Ama-
zonia, Manaus, Brazil (INPA); Museo de Insectos, Universidad de Costa Rica,
San Pedro (MIUCR); Instituto Nacional de Biodiversidad, Heredia, Costa Rica
(INBio); Museo de Invertebrados “‘G.B. Fairchild’, Universidad de Panama
(MIUP).
Lophostigma grisselli Cambra and Quintero, NEW SPECIES
(Figs. 1—3, 7)
Types.—Holotype, female: MEXICO. N[orth] YUCATAN [State]: Temax, Gau-
mer col., BM (NH) [P. Cameron Coll. 1914-110 / 43 / Lophostigma sp. CEM
1955]. The latter is C. E. Mickel’s hand written det. label. Paratypes: same data
as holotype, one female; deposited: MIUP; 8 mi E[ast] of Mérida, 28 Jun 1966,
U. Kansas Mex. Exped., one female; deposited: Snow Entomological Museum,
University of Kansas.
Description—Holoty pe female.—Vertex with large yellow integumental spot; antennal tubercles,
mandible and clypeus light brown-red, scape dark red, rest of head black. Thorax black, except for
yellow propodeal spiracles. Abdomen black except sternum I and two distal segments red-brown;
tergum II with two suboval-transverse postmedian yellow integumental spots, separated by a distance
equal to one-third the width of a spot. Front, vertex and gena with deep, close punctures, not confluent.
Antennal scrobes strongly carinate dorsally. Antennal tubercles smooth, set distinctly apart. Anterior
margin of clypeus bituberculate in front of antennal sockets; mandible slender, edentate at tip and not
toothed within. Distance from posterior eye margin to posterior angle of genal carina 0.93 times
greatest diameter of eye. Gena carinate, carina not forming a tooth anteriorly, not extending to vertex
posteriorly. Front, vertex and gena with sparse, pale pubescence; scape and clypeal fringe with sparse,
fine pale pubescence. Thoracic dorsum distinctly convex in lateral view, with moderate, dense, but
not confluent punctures. Propodeum with short anterodorsal mesal longitudinal carina, distinctly higher
in middle. Anterolateral angles of pronotum defined by short, transverse carina; sides of pronotum
with moderate, close punctures. Lateral margins of mesonotum with pair of small teeth; propodeum
laterally with four sharp teeth. Sides of propodeum glabrous on anterior third, with strong, distinct,
deep punctures on posterior two-thirds. Meso- and metapleura glabrous, micropunctate. Dorsum of
thorax with sparse pale pubescence, and scattered long, erect pale hairs. Dorsum of propodeum with
pale micropubescence and scattered, long erect pale hairs. Mesopleura, metapleura and sides of pro-
podeum with sparse, inconspicuous, decumbent pale pubescence; posterior margin of mesopleura with
row of long, erect pale hairs. Legs dark red, almost black, clothed with pale pubescence. Tibial spine
on posterodistal internal angle slightly longer than the cylindrical process. Calcaria pale. Anterior face
of tergum I with small, sparse punctures; dorsal face with moderate, dense punctures. Tergum II with
anterolateral areas only slightly elevated, leaving a shallow depression in between, and crested with
short, parallel, interrupted carinae; with moderate, dense contiguous punctures throughout. Terga III-—
V with fine, close punctures. Pygidial area glabrous. Sternum I with a short, longitudinal, median
carina, higher anteriorly. Sternum II with moderate, close punctures. Sterna III-VI throughout with
fine, close punctures. Tergum I with sparse, erect, appressed pale pubescence. Tergum II with a short
anteromedian, longitudinal stripe of pale pubescence; posterior margin with short, dense, black pu-
bescence, except for small subtriangular spot of pale pubescence at middle; remainder of tergum II
94 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Figure 1. Lophostigma grisselli, NEW SPECIES. Dorsal habitus.
with sparse, erect and semierect pale hairs. Terga III—-V clothed with dense, pale pubescence. Sterna
covered with sparse pale hairs. Length: 8.6 mm.
Variation—The paratype deposited at MIUP has the frons, gena, thorax, ab-
dominal sterna II-IV, and legs dark red. These structures are black in the holotype
and the other paratype.
1996 CAMBRA & QUINTERO: MEXICAN AND C. AMER. LOPHOSTIGMA 95
oy
Figures 2-3. Lophostigma grisselli, NEW SPECIES. Figure 2. Propodeum, lateral view. Figure 3.
Thorax, lateral outline. Figures 4-5. Lophostigma cincta (du Buysson). Figure 4. Propodeum, lateral
view. Figure 5. Thorax, lateral outline. Abbreviations: ep = endophragmal pit; Ipp = lateral propodeal
punctures; mp = mesopleural punctures; pc = propodeal carina; pst = propodeal spiracle, tubercle.
Diagnosis——This species runs to L. simoni, from Venezuela, in the second part
of couplet 4 of Mickel’s key (1952). Both species have the dorsum and posterior
face of propodeum with a pair of broad, lateral stripes of appressed pale pubes-
cence. They can be separated as follows: L. grissellihas the propodeal side smooth
on the anterior third while in L. simoni the side is deeply punctate; L. grisselli
has an EMGC/EGD ratio of 0.92 + 0.03 (n = 3), largest in the genus (0.81 in
L. acantho phora, 0.59 in L. simoni); in lateral view the thorax of L. grisselli is
convex dorsally while that of L. simoni, as other species of Lophostigma we have
examined (specimens of L. seabrai and L. iracunda have not been seen) presents
a flat dorsal thoracic surface. In addition, L. grisselli has a large yellow integu-
mental spot on the vertex; in L. simoni the head is uniformly dark red but col-
oration is not a reliable character to separate species (see Taxonomic Notes).
Etymology.—Named in honor of our colleague and friend, Dr. E. Eric Grissell,
Systematic Entomology Laboratory, U.S. Department of Agriculture, USNM.
Lophostigma cincta (du Buysson), 1892
(Figs. 4-6)
Mutilla cincta du Buysson 1892: 58: Type locality: VENEZUELA. CARABOBO
STATE: San Esteban (Parque Nacional Miguel J. Sanz), M. E. Simon col.
96 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Figure 6. Lophostigma cincta (du Buysson). Head, lateral view. Abbreviations: af = antennal
foramen left, egd = eye greatest diameter; emgc = distance from eye posterior margin to genal carina.
Holotype female deposited MNHN, Paris, examined; Lophostigma cincta:
Mickel, 1952: 147 (in key).
Pseudomethoca lebasi Mickel 1937: 179-180. Type locality: COLOMBIA (no
additional data). D. Lebas col. Holotype female deposited Spinola Collection
No. 114, Torino, Italy; Lophostigma lebasi: Mickel 1952: 146-7 (in key, Pan-
1996 CAMBRA & QUINTERO: MEXICAN AND C. AMER. LOPHOSTIGMA eA
Figure 7. Distribution of species of Lophostigma. Solid symbols: L. grisselli (square), L. cincta
(triangle), and L. subgracilis (circle). Open symbols: L. cayennensis (triangle), L. acanthophora
(square), L. caenodonta (circle).
ama and Colombia); Cambra and Quintero 1992: 473 (six females, records for
Panama). NEW SYNONYM Y.
Notes on Synonymy.—The only recognized difference between L. lebasi and L.
cincta was coloration of the vertex. In /ebasi a large, yellow integumental spot is
present, while in cincta this spot is entirely ferruginous (“‘téte jaune’’ of du Buys-
son’s original description changes to “‘vertex entirely ferruginous”’ in Mickel’s
1952 key). As indicated in Taxonomic Notes, color is not a reliable character to
separate species in Lophostigma because of the large intraspecific variation. The
30 females examined from Panama have a yellow spot on the vertex but its size
is quite variable. No specimens have been seen from Colombia, and the type
remains the only specimen reported for that country. At the latitudinal extremes
of the distributional range of L. cincta we found a marked reduction or absence
of yellow spots on the vertex in six of the eight females examined outside of
Panama. Of five specimens from Costa Rica, two have the vertex solid black, and
one solid red. The Ecuadorian specimen has two very small yellow spots on the
vertex.
Distribution—Costa Rica, Panama, Colombia, Venezuela and Ecuador (Fig. 7).
Material Examined.—Specimens from Costa Rica and Ecuador represent new distribution records.
COSTA RICA. GUANACASTE PROVINCE: Estacion Maritza, 600 m, lado oeste Volcan Orosi, 27
Feb-10 Mar 1992, R. Vargas, 1 female (MIUP); Refugio Nacional Fauna Silvestre Rafael Lucas
Rodriguez, Palo Verde, 10 m, Mar 1991, U. Chavarria, 1 female (INBio); PUNTARENAS PROVINCE:
98 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Tres Rios, 18 Apr 1940, A. Bierig, 1 female (MIUCR); Osa Peninsula, 2.5 mi SW of Rincén (08°42'
N, 83°29’ W), 26 Feb 1967, C. Rettenmmeyer, | female (SMUK); SAN JOSE PROVINCE: 3 km S
of San Antonio de Escazt, 10-25 Dec 1987, W. Wcislo, 1 female (MIUP).: ECUADOR. NAPO
PROVINCE: Yuturi, 27 Feb 1990, S. Sandoval, 1 female (MIUP). PANAMA. COCLE PROVINCE:
Valle de Antoén, 13 Jul 1991, 27 Dec 1991, R. Contreras, 2 females (MIUP); DARIEN PROVINCE:
El Real, 4 Feb 1993, R. Cambra, 1 female (MIUP); Cruce de Mono, Estaci6n INRENARE, Parque
Nacional Darién, 9—23 Feb 1993, R. Cambra, 8 females (MIUP); PANAMA PROVINCE: Summit
Gardens, 9 Jan 1989, J. Bernal, 1 female (MIUP); Summit Gardens, 18 Dec 1994, D. Osorio, 1 female
(MIUP); Capira, 17 May 1991, 18 May 1991, 17 Jan 1992, R. Contreras, 3 females (MIUP); Capira,
La Julianita, Ollas Arriba, M. Castro, 31 Dec 1994, 2 females (MIUP); Potrero Rio Perequeté, Chorrera
Dist., 17 Jan 1992, R. Contreras, 2 females (MIUP); same loc, 7 Aug 1991, R. Contreras, 1 female
(MIUP); same loc, 27 Feb 1991, R. Cambra, 1 female (MIUP); Llano Largo, Chorrera Dist., 8 Apr
1990, A. Mena, 1 female (MIUP). VENEZUELA. FALCON STATE: Yaracal, 30 Apr 1989, L. J. Joly,
1 female IZAUCV); BARINAS STATE: Calderas, 1000 m, 8 May 1972, J. & B. Bechyne, 1. female
(MIUP).
Lophostigma subgracilis (Cameron), 1895
Sphaerophthalma subgracilis Cameron 1895: 339-340. Type locality: PANAMA.
Volcan de Chiriqui, 1220 m. Syntype females deposited BM (NH), examined;
Lophostigma subgracilis: Mickel 1952: 146-147 (in key); Cambra & Quintero
1992: 473.
Color variation—-The Teapa female has a black head with a small yellow
integumental spot on the vertex and a very thin, elongated spot of same color at
base of second tergite, and the thorax black. Two females from El Copé, Panama,
have the anteromedian part of the second tergite and the head integument uni-
formly black ferruginous, one with ferruginous thorax and the other with thorax
mostly black, except for lateral propodeum ferruginous. Two females from Costa
Rica: one with head mostly yellow-ferruginous, thorax black-ferruginous; other
female, yellow spot on vertex and rest of head black, thorax black.
Distribution—Mexico, Guatemala, Costa Rica and Panama (Fig. 7). Five of
the eight specimens studied (including the syntype) were collected on or near the
Cordillera Central, at elevations between 800 to 1200 m (personal observation).
On this evidence we recognize that L. subgracilis is the only species in the genus
that occurs primarily at altitudes higher than 800 m.
Material Examined.—Specimens from Mexico, Guatemala and Costa Rica represent new distribu-
tion records. MEXICO. TABASCO STATE: Teapa, Feb (H.H.S.), 1 female BM (NH). GUATEMALA.
ALTA V[ERA] PAZ DEPARTMENT: Cacao, Trece Aguas, Schearz and Barber, 1 female (USNM).
COSTA RICA. PUNTARENAS PROVINCE: Golfo Dulce, 24 km west of Piedras Blancas, 200 m,
Apr-May 1991, C. Hanson, 1 female (MIUCR); CARTAGO PROVINCE: Chitaria [alt. approx. 769
mJ], 17-20 Feb 1943, 1 female (MIUP). PANAMA. COCLE PROVINCE: El Copé, Div. Continental,
900 m, 21 Feb 1990, R. Cambra, 1 female (MIUP); same loc., 1-2 Sep. 1990, R. Cambra, 1 female
(MIUP).
Lophostigma acanthophora (Dalla Torre), 1897
Mutilla spinifera Smith 1879: 213, nec Olivier 1811: 59. Type locality: BRAZIL.
Para. Holotype female deposited BM (NH), London, examined.
Mutilla acanthophora Dalla Torre 1897: 6, new name for spinifera Smith; Lo-
phostigma acanthophora: Mickel 1952: 146 (in key).
Distribution—This species was known previously only from the holotype from
Brazil; Peri (Fig. 7).
1996 CAMBRA & QUINTERO: MEXICAN AND C. AMER. LOPHOSTIGMA 99
Material Examined.—The specimen from Pert represents a new distribution record. PERU. MADRE
DE DIOS DEPARTMENT: Reserva de Manu, Cocha Cashu, 27 Feb 1992, I. Bohorquez, 1 female
(MIUP).
Lophostigma caenodonta (Cameron), 1912
Ephuta ? caenodonta Cameron 1912: 415. Type locality: GUYANA (no additional
data). Holotype female deposited BM (NH), London, examined; Lophostigma
caenodonta: Mickel 1952: 146-148 (Gn key, redescription of type).
Distribution Guyana and Brazil (Fig. 7).
Material Examined.—Specimens from Brazil represent new distribution records. BRAZIL. MA-
RANHAO STATE: Isla Sao Luis, Foresta Sacavem, 1 Oct 1992, R. Cambra, 1 female (MIUP); Isla
Sao Luis, Foresta Sacavem, 1 Oct 1992, R. Cambra, 1 female (MIUP); Isla SAo Luis, Vila Maranhao,
29 Sep 1992, D. Quintero, 1 female (MIUP).
Lophostigma cayennensis (André), 1906
Ephuta cayennensis André 1906: 70—72. Type locality: FRENCH GUIANA. Cay-
enne. Holotype female deposited MNHN, Paris; Lophostigma cayennensis:
Mickel 1952: 146 Gn key).
Distribution—French Guiana, Venezuela and Brazil (Fig. 7).
Material Examined.—The specimens from Venezuela and Brazil represent new distribution records.
VENEZUELA. TERRITORIO FEDERAL AMAZONAS: Yavita, Pimichin, Apr 1952, Buchel, 1 female
(MIUP). BRAZIL. AMAZONAS STATE: Manaus, 24 Jun 1977, A. P. Luna Dias, 1 female (INPA);
same loc., 20 Jun 1977, 2 females (MIUP).
Three species of Lophostigma are now known from Venezuela: L. cayennensis, L. simoni and L.
cincta.
TAXONOMIC NOTES ON LOPHOSTIGMA
The genus Lophostigma is closest to Lophomutilla Mickel, presenting a re-
markable resemblance in most characters, except for the shape of the mandibles
(distally tridentate in Lophomutilla, edentate in Lophostigma) and the first abdom-
inal segment (nodose in Lophomutilla, disciform in Lophostigma). In addition,
the following combination of characters, although not autapomorphies, permits
the recognition of specimens of Lophostigma: a median longitudinal low carina
on the propodeum; an anteromedian elongated short pale pubescent spot on the
second abdominal tergum; and distinct parallel, longitudinal, interrupted carinae
on each side of the anteromedian area of pale pubescence on the second abdominal
tergum. The generic description by Mickel (1952) is quite thorough, but of the
above generic characters, the first two were not mentioned by Mickel.
We have recently reported the sex association of Paramutilla Mickel, 1973 as
the males of Lophomutilla Mickel, 1952 (Quintero & Cambra 1996). In this work,
we expanded our previous generic key (Cambra & Quintero 1992) to include the
mutillid taxa from South America. We expect that comparison of the males of
Lophostigma, with those of Lophomutilla will provide additional characters valu-
able to separate both genera.
The two groups of species in Mickel’s key (1952), were based on the degree
of elevation of the anterolateral areas of the abdomen (strongly or inconspicuously
elevated). We now recognize that L. grisselli and L. seabrai, which are found at
the distributional limits of the genus, north and south respectively, are interme-
100 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
diate with respect to the elevation of the anterolateral areas, thus the two groups
are not clearly defined.
The size of the integumental spots (one anteromedian and two transverse, post-
median) on the second abdominal tergum, and the coloration of the thorax are
variable in L. cayennensis, L. grisselli, L. subgracilis and L. cincta, and thus,
cannot be used to separate species. Lophostigma cayennensis is probably a junior
synonym of L. iracunda (Cresson), 1902 (species known only from the holotype
from Santarerm, Brazil), because it has been separated only by differences in
thoracic coloration and the size of the integumental spots on abdominal tergum
two.
Casal (1963) erroneously considered the presence of the median longitudinal
propodeal carina as the main difference between L. seabrai Casal and L. alopha
Mickel (present in the former, from Brazil, and absent in the latter, from Guyana).
However, the type of L. alopha has a faint carina, and the two species are oth-
erwise indistinguishable. Thus, L. seabrai probably is a junior synonym of L.
alopha.
KEY TO THE MEXICAN AND CENTRAL AMERICAN SPECIES OF
LOPHOSTIGMA (FEMALES ONLY)
1. Anterolateral area of second abdominal tergum slightly swollen, leaving a dis-
tinct, shallow medial depression (Fig. 1); median longitudinal carina on pro-
podeum highest in the middle; in lateral view, outline of thoracic dorsum
convex (Fig. 3); EMGC/EGD ratio: 0.92 (0.88 to 0.95) ..............
SOU En yee, rea ek ee evel en oe ee, eee eee ee grisselli NEW SPECIES
— Second abdominal tergum uniformly convex anteriorly, not swollen anterolat-
erally; median longitudinal carina on propodeum uniformly elevated along its
entire length; in lateral view, outline of thoracic dorsum flat (Fig. 5); EMGC/
Gi) -Tatiot OW OE MSS G5 ire ary toe Mere cat OMAN Re tee PR oy Od Shee: ape 8 Be one oe 2
2. Mesopleura and sides of propodeum densely punctate (Fig. 4); propodeal dor-
sum with a median longitudinal black pubescent band (Costa Rica through
We ZUG 1A wr CUAC OI): 2 ona deo er nC MeL ES tee, cyto os MMS ot ote cincta (du Buysson)
— A few scattered punctures on posterior margins of mesopleura and sides of
propodeum; propodeum covered with dense short white pubescence (Mexico,
Guatemala, Costa Rica, Panama) ................... subgracilis (Cameron)
ACKNOWLEDGMENT
We are grateful to the British Embassy in Panama, in particular to Ambassador
Thomas H. Malcomson, for securing transportation funds to England for R.A.C.;
to the Entomology Department personnel of the British Museum (Natural Histo-
ry), London, for providing R.A.C. with working facilities and assistance during
his visit. Our thanks to the Smithsonian Tropical Research Institute (STRI), in
particular to Ira Rubinoff, for providing research facilities. We highly appreciate
STRI Research Opportunities Fund No. 1234F20A-3400 to D.Q.A., that helped
us carry out field work in Brazil, and the great hospitality and facilities provided
by Patricia Albuquerque, Sao Luis, Maranhao, Brazil. We appreciate the loan of
unidentified mutillid specimens from John E. Lattke, Instituto Zoologia Agricola,
Universidad Central de Venezuela, Maracay, Robert Brooks, Snow Entomological
1996 CAMBRA & QUINTERO: MEXICAN AND C. AMER. LOPHOSTIGMA 101
Museum, University of Kansas, and Professor Giovanni Onnore, Universidad Pon-
tificia Catdélica del Ecuador, Quito. We also thank José Albertino Rafael, Instituto
Nacional de Pesquisas da Amazonia, Manaus, Brazil, Roger D. Hutchings H..,
INPA-Biological Dynamics of Forest Fragments Project-Smithsonian Institution,
Paul Hanson, Museo de Insectos, Universidad de Costa Rica, San Pedro, Jestis
Armando Ugalde Gémez, Instituto Nacional de Biodiversidad, Heredia, and Philip
J. Clausen, Department of Entomology, University of Minnesota, St. Paul, for
hospitality and loan of specimens. Recent collections of L. cincta from Darién
Province were made possible by funds Nos. 1-4500-91-12 & 04-05-94-01 from
Vicerrectoria de Investigaci6n y Postgrado, Universidad de Panama. Our thanks
to Arnold S. Menke, Karl V. Krombein, USNM, E. E. Grissell, U.S. Department
of Agriculture-USNM, and Annette Aiello, STRI, for reviewing the manuscript
and for their helpful suggestions. Special thanks to Isabelita Bohorquez, Univer-
sidad Nacional de San Marcos, Lima, Peru, for collecting and donating to us the
very rare female of L. acanthophora.
LITERATURE CITED
André, E. 1906. Nouvelles espéces de Mutillides d’Amerique (Hym.). Zeit. Hym. and Dipt., 6: 65-80.
Cambra T., R. & D. Quintero A. 1992. Velvet ants of Panama: distribution and systematics (Hyme-
noptera: Mutillidae). pp. 459-478. Jn Quintero A., D. & A. Aiello (eds.). Insects of Panama
and Mesoamerica: selected studies. Oxford University Press, Oxford.
Cambra T., R. & D. Quintero A. 1993. Timulla Ashmead (Hymenoptera: Mutillidae): new distribution
records and synonymies, and descriptions of previously unknown allotypes. Pan-Pacific Ento-
mol., 69(4): 296-310.
Cameron, P. 1894-96. Mutillidae. pp. 259-395. In Biologia Centrali Americana, Hymenoptera 2.
Cameron, P. 1912. The Hymenoptera of the Georgetown Museum. Part IV. The fossorial Hymenop-
tera. Timehri J. Royal Agri. Com. Soc. Brit. Guiana, 2: 413-416.
Casal, O. H. 1963 (1961). Mutillidae Neotropicales. 18. (Hymenoptera). Dos nuevas especies de
Argentina y Brasil. Rev. Soc. Entomol. Argentina, 24: 71-76.
Dalla Torre, C. G. 1897. Catalogus Hymenopterorum, 8 (Fossores): 1-749.
du Buysson, R. 1892. Voyage de M. E. Simon au Venezuela, Dec. 1887—April 1888. Hymenopteres.
Ann. Soc. Entomol. France., 61: 55-59.
Mickel, C. E. 1937. New World Mutillidae in the Spinola collection at Torino, Italy (Hymenoptera).
Rev. Entomol. (Rio de Janeiro), 7: 165—207.
Mickel, C. E. 1952. The Mutillidae (wasps) of British Guiana. Zoologica, 37(3): 105-150.
Olivier, A. G. 1811. Encyclopédie méthodique. Histoire naturelle. Insectes, 8: 51-66.
Quintero, D. & R. A. Cambra. 1996. Contribucion a la sistematica de las mutilidas (Hymenoptera)
del Pert, en especial las de la Estacién Biol6gica BIOLAT, Rio Manu, Pakitza. pp. 263-293.
In Wilson, D. E. & Sandoval, A. (eds.). La Biodiversidad del Sureste del Pert: Manu. Office
of Biodiversity Programs, Smithsonian Institution. Editorial Horizonte, Lima, Pert.
Smith, E 1879. Descriptions of new species of Hymenoptera in the collection of the British Museum.
London, pp. 189-227.
PAN-PACIFIC ENTOMOLOGIST
72(2): 102-103, (1996)
Scientific Note
RECENT CALIFORNIA RECORDS FOR THE SAWFLY
XIPHYDRIA MELLIPES HARRIS (HYMENOPTERA:
XIPHYDRITIDAE)
The Xiphydriidae, with 22 genera and 90 species, is a small family of wood-
boring sawflies that are found in most of the world except Africa (Smith, D. R.
1976. Trans. Am. Entomol. Soc., 102: 101—131). The only North American genus
is Xiphydria, with 10 species now occurring in the U.S. and Canada (Smith, D.
R. 1983. Proc. Entomol. Soc. Wash., 85: 860-861). In his revision of Xiphydria,
Smith (1976) states that X. mellipes Harris is the most widespread North American
Species in the genus occurring across Canada and the northern U.S. Although
(especially southern) British Columbia has. several records, his distribution map
(Fig. 1) and text show the only western U.S. records to be from NW Montana
and NE Oregon.
In the last decade X. mellipes has been recorded from non-quarantine situations
from several California locations (Fig. 1) in Fresno, Sacramento, Santa Clara and
San Mateo counties; therefore, its listed nearctic distribution warrants expansion.
Xiphydria mellipes is apparently restricted to Betula spp., although other ques-
tionable host records exist (Smith 1976: 118-119). It occurs in dead or decaying
logs, and commonly emerges from firewood brought into buildings (Smith 1976).
It bores galleries about 2.5 mm dia. in the partly decaying heartwood of standing
trees (Yuasa, H. 1922. Illinois Biol. Monographs, 7[4]: 1-172).
It is uncertain if the California records represent a recent range expansion, either
naturally or by movement of wood (e.g., firewood), or reflect a more accurate
sampling of the state’s fauna. Several Betula spp. have been planted in California
urban settings as ornamentals, but two native species do. exist in the state: Betula
glandulosa Michaux [north California] and Betula occidentalis Hook [north Cal-
ifornia, south through the Sierra Nevada to desert mountain ranges and the Great
Basin] (Hickman, J. C. [ed.] 1993. The Jepson manual, higher plants of California.
Univ. California Press, Berkeley; Griffin, J. R. & W. B. Critchfield. 1972. USDA
Forest Serv. Res. Paper, PSW-82/1972: map 13.). It seems most probable that X.
mellipes occurs indigenously in California but was undetected until recently. It
may have moved into urban settings naturally infesting aging ornamental Betula,
and/or with the movement of firewood from the northern or Sierran parts of the
state.
Xiphydria mellipes represents the second, but most confirmed, species of Xiph-
ydria occurring in California. A single (questionable) record for X. maculata Say,
an otherwise eastern species primarily associated with Acer spp., exists for Cal-
ifornia (Smith 1976: fig. 3); that record is “*... considerably out of the range of
maculata and is based on a single specimen labeled ‘Cal.’, ‘Collection T. Per-
gande.’’”’ (Smith 1976: 115).
Records —CALIFORNIA. FRESNO Co.: Fresno, 11. Apr 1991, N. Smith, “‘ex birch,’ 3 males, 4
females. NAPA Co.: Napa, 6 Jan 1982, ‘““Whitmer’’ collr., 1 male, 1 female. SACRAMENTO Co.: no
locality given, 16 Dec 1991, L. Allen, “‘reared Betula sp.,” 1 male, 1 female. SAN MATEO Co.:
1996 SCIENTIFIC NOTE 103
Figure 1. Distribution of Xiphydria mellipes, modified from Smith (1976: fig. 4). Black dots are
records in Smith (1976), white triangles are added California records.
Redwood City, 27 Feb 1992, “‘Eaton’’ collr., “in house,’ 1 female. SANTA CLARA Co.: no locality
given, late Mar or early Apr 1994, A. Clark, ‘‘found inside [residence] near/on window,”’ 1 female.
John T. Sorensen, Plant Pest Diagnostics Center, California Dept. of Food &
Agriculture, 3294 Meadowview Rd., Sacramento, California 95832-1448.
PAN-PACIFIC ENTOMOLOGIST
72(2): 104-109, (1996)
PROCEEDINGS OF THE PACIFIC COAST
ENTOMOLOGICAL SOCIETY, 1992
FouUR HUNDRED AND EIGHTY-NINTH MEETING
The 489th meeting of the Pacific Coast Entomological Society was held on 17 January 1992, at
8:00 PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Norman D. Penny presiding.
The minutes of the meeting held 20 December 1991 were read and accepted. Six persons were
proposed and elected as new members: Mr. Eric W. Riddick, Ms. Katherine M. Schick, Mr. Steven
J. Seybold, and Ms. Diane M. Stark as student members; Dr. James F. Cornell Jr. and Dr. Conley K.
McMullen as regular members.
Dr. Penny announced that a meeting of the Association of Biologists in Computing will be held 18
January 1992 at California State University, Hayward. He also announced that the Northern California
Lepidopterists will meet on 15 February 1992 and that the location will be listed in the February Bits
and PCES. Mr. Curtis Y. Takahashi presented a book titled The Illustrated Book of Insects which he
had purchased at a retail bookstore. Dr. Penny presented slides of many of the California species of
Mecoptera and briefly described the distributions and habits of each California species.
The featured speaker Dr. Wojciech J. Pulawski, California Academy of Sciences, presented a lecture
entitled ‘Wasp Collecting in Africa.”” He showed slides and discussed his study of the wasp genus
Gastrosericus (Sphecidae) and his discovery of range extensions for certain species. He stressed the
importance of field work by systematists and taxonomists as a means of discovering life history traits
of the taxa which they are studying.
The meeting adjourned at 9:40 PM, followed by a social hour in the entomology conference room. —
D. K. Dabney, Recording Secretary.
The following 41 persons were present. 32 members: P. H. Amaud Jr., T. S. Briggs, R. M. Brown,
P. Buickerood, R. Buickerood, J. S. Chinn, H. K. Court, D. K. Dabney, W. A. Doolin, J. G. Edwards,
B. K. Eya, S. S. Ferguson, W. E. Ferguson, R. L. Langston, V. F. Lee, W. A. Maffei, G. J. Mallick,
D. L. Mead, N. D. Penny, M. Prentice, W. J. Pulawski, A. E. Rackett, J. M. Ribardo, K. J. Ribardo,
R. G. Robertson, E. S. Ross, R. E. Stecker, C. Y. Takahashi, S. P. Taormino, D. Ubick, C. E. Warren,
and S. P. Wells; (9 guests) M. M. Arnaud, R. Bandar, J. E. Court, N. Doolin, S. Haugues, A. M. L.
Penny, C. Prentice, W. A. Rauscher, and J. Robertson.
FouUR HUNDRED AND NINETIETH MEETING
The 490th meeting of the Pacific Coast Entomological Society was held on 21 February 1992, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Norman D. Penny presiding.
The minutes of the meeting held 17 January 1992 were read and accepted. Six persons were proposed
and elected as new members: Ms. Caroline M. Crenshaw, Mr. William Hamersky, Ms. Stacie T.
Kawaguchi, and Ms. Barbara M. Rice as student members; Ms. Lisa J. Boutin and Mr. Luis A.
Solorzano as regular members.
Dr. Penny announced that there had been a business meeting and that the Society will buy a new
computer. Dr. Penny announced that the Society’s old computer is now for sale, and that he and
Vincent Lee will entertain offers. Dr. Ronald E. Stecker announced that Dr. Edgar Smith had died on
2 February 1992, read brief details concerning Dr. Smith’s career, and mentioned that Dr. Paul H.
Arnaud, Jr. will be publishing a more complete account of Dr. Smith’s career in the Society’s journal.
Dr. Penny announced that Dr. David H. Kavanaugh, California Academy of Sciences, has recently
published a study of the carabid beetles of the Charlotte Islands, British Columbia, and that this
monograph is available from Scientific Publications at the Academy.
Dr. Penny presented a note concerning Oliarces clara Banks (Neuroptera: Ithonidae), and mentioned
that it is the only New World species in this family.
The featured speaker Dr. J. Gordon Edwards, San Jose State University, presented a lecture entitled
“Insect Collecting in Australia and New Zealand.” His slides and discussion of his travels in these
1996 PROCEEDINGS 105
countries included accounts of insects, numerous other animals, and plants. Dr. Edwards also described
a brief trip to Tasmania.
The meeting adjourned at 9:25 PM, followed by a social hour in the entomology conference room.—
D. K. Dabney, Recording Secretary.
The following 60 persons were present. 46 members: P. H. Arnaud Jr., V. M. Barlow, L. G. Bezark,
F. L. Blanc, T. S. Briggs, K. W. Brown, R. M. Brown, P. Buickerood, R. Buickerood, J. R. Clopton,
J. S. Cope, H. K. Court, C. M. Crenshaw, T. D. Cuneo, D. K. Dabney, W. A. Doolin, J. G. Edwards,
B. K. Eya, S. V. Fend, S. S. Ferguson, W. E. Ferguson, M. P. Fish, P. S. Johnson, D. H. Kavanaugh,
S. T. Kawaguchi, R. L. Langston, V. F. Lee, W. A. Maffei, P. S. McNally, D. L. Mead, S. B. Opp, N.
D. Penny, A. E. Rackett, S. Renkes, K. J. Ribardo, B. M. Rice, R. G. Robertson, R. K. Roche, W.
E. Savary, H. I. Scudder, S. J. Seybold, R. E. Stecker, C. Y. Takahashi, S. P. Taormino, R. W. Thorp,
and C. E. Warren; (14 guests) A. Bandar, R. Bandar, C. Benesh, F. Blanc, J. E. Court, N. Doolin, B.
Kavanaugh, P. Krauny, D. Limberg, C. Miller, A. M. L. Penny, W. A. Rauscher, K. Reynolds, and
J. Robertson.
FouR HUNDRED AND NINETY-FIRST MEETING
The 49 1st meeting of the Pacific Coast Entomological Society was held on 20 March 1992, at 8:00
PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Norman D. Penny presiding.
The minutes of the meeting held 21 February 1992 were read and accepted. Two persons were
proposed and elected as new regular members: Mr. Floyd C. Baptista and Mr. Fred J. Moitoza.
Ms. Leslie S. Saul announced an open house of the San Francisco Zoological Society’s Insect Zoo
scheduled for 17 May 1992 and invited PCES members to attend or participate. Ms. Saul also
mentioned that the Insect Zoo was displaying Scolopendra gigantea (Chilopoda: Scolopendridae), a
26 cm (10%%”) centipede from Trinidad, and the Sexton Beetle Nicrophorus carinata (Coleoptera:
Silphidae). Dr. Penny announced the “Mariposa Butterfly Bonanza,” a short course scheduled for 8-9
May 1992 with John Lane of the Santa Cruz City Museum. He also mentioned “Bugfair,”’ an insect
exhibit scheduled for 6 June 1992 by the Sulphur Creek Nature Center in Hayward. Dr. Penny also
reported that the Entomology Department of the California Academy of Sciences was giving away
surplus large insect trays, and that on 5 April 1992 he and others would be driving to Tulare Co. to
collect insects from a flume trap with Dr. Donald J. Burdick and invited any interested members to
join the group.
The featured speaker Mr. Wesley A. Maffei of the Alameda County Mosquito Abatement District
presented a lecture entitled “Insect Iridescence and Morpho Colors.” He discussed insect colors in
general and his research on iridescence in Morpho butterflies.
The meeting adjourned at 9:12 PM, followed by a social hour in the entomology conference room.—
D. K. Dabney, Recording Secretary.
The following 24 persons were present. 19 members: V. M. Barlow, D. K. Dabney, W. A. Doolin,
J. G. Edwards, B. K. Eya, W. Hamersky, S. T. Kawaguchi, V. F. Lee, W. A. Maffei, D. L. Mead, N.
D. Penny, A. E. Rackett, R. G. Robertson, L. S. Saul, K. N. Schick, C. Y. Takahashi, S. P. Taormino,
D. Ubick, and R. A. Worth; (5 guests) N. Doolin, D. Maffei, A. M. L. Penny, J. Robertson, and J.
Schick.
FouR HUNDRED AND NINETY-SECOND MEETING
The 492nd meeting of the Pacific Coast Entomological Society was held on 24 April 1992, at 8:00
PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Norman D. Penny presiding.
The minutes of the 20 March 1992 meeting were read and accepted. Two persons were proposed
and elected as new regular members: Mr. J. Scott Cox and Mr. Andrew P. Jarman.
Ms. Leslie S. Saul announced a full-time permanent position at the San Francisco Zoological Society’s
Insect Zoo. Dr. Penny noted three full-time seasonal positions with the San Mateo Mosquito Abatement
District. He also mentioned that the organizers of ““Bugfair,” an insect exhibit scheduled for 6 June
1992 by the Sulphur Creek Nature Center in Hayward, are interested in having people display their
106 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
insect collections. Dr. Penny also announced a collecting trip to a flume site in Tulare County, and
the Xerces Society’s annual butterfly count scheduled for July.
Mr. Curtis Y. Takahashi announced insect-related exhibits scheduled forthe Sanborn Youth Science
Institute in Saratoga on 9 May 1992. Dr. Harvey I. Scudder mentioned that he has seen large groups
of migrating Painted Lady (Vanessa cardui) butterflies and that some local media had mistakenly
called them Red Admirals (V. atalanta). Dr. William E. Ferguson noted having seen Painted Lady
butterflies moving from the Wawona area in the southern Sierra, contrary to what he knew of their
migration patterns. Mr. Robert L. Zuparko requested information about collecting sites in the New
Orleans area. Ms. Saul showed slides of a spring trip to the deserts of Southern California and Arizona,
including those of desert wildflower bloom and the Sonoran Arthropod Studies center and remarked
on the center’s education and research projects.
The featured speaker Dr. Steven L. Heydon presented a slide lecture entitled “Summer Camp for
Professionals— Collecting in Rondonia, Brazil.” He discussed tour details, field stations, and some of
the insects he collected.
The meeting adjourned at 9:05 PM, followed by a social hour in the entomology conference room. —
D. K. Dabney, Recording Secretary.
The following 38 persons were present. 30 members: P. H. Amaud Jr., V. M. Barlow, L. J. Boutin,
T. S. Briggs, K. W. Brown, J. S. Chinn, D. K. Dabney, S. S. Ferguson, W. E. Ferguson, W. Hamersky,
A. Hom, A. S. Hunter, A. P. Jarman, D. H. Kavanaugh, V. F. Lee, G. J. Mallick, J. D. McCarty, N.
D. Penny, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, L. S. Saul, K. N. Schick,
H. I. Scudder, R. E. Somerby, C. Y. Takahashi, R. W. Thorp, D. Ubick, and R. L. Zuparko; (8 guests)
M. M. Arnaud, R. Bandar, S. L. Heydon, R. Johnson IV, A. M. L. Penny, J. C. Rackett, J. Robertson,
and W. C. Rauscher.
FouR HUNDRED AND NINETY-THIRD MEETING
The 493rd meeting of the Pacific Coast Entomological Society was held on 15 May 1992, at 8:00
PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Norman D. Penny presiding.
The minutes of the 24 April 1992 meeting were read, corrected, and accepted. There were no new
members proposed. Dr. Penny mentioned the AAAS Pacific Division annual meeting sceduled for
21-25 June 1992 in Santa Barbara.
Dr. Paul H. Arnaud, Jr. presented a note concerning Achias rothschildi (Diptera: Platystomatidae)
and displayed three specimens. He noted that with a head width of up to 52 mm, this species might
be the widest-headed fly known and that most of the 98 species of the genus occur in New Guinea.
The featured speaker Dr. Edward S. Ross of the California Academy of Sciences presented a slide
lecture entitled “Recent Encounters with Insects in South America.” He discussed his photographic
and observational examples of numerous evolutionary phenomena including camouflage and mimicry.
The meeting adjourned at 9:50 PM, followed by a social hour in the entomology conference room.—
D. K. Dabney, Recording Secretary.
The following 53 persons were present. 31 members: P. H. Amaud Jr., V. M. Barlow, B. T. Berke,
L. G. Bezark, L. J. Boutin, T. S. Briggs, R. M. Brown, P. Buickerood, R. Buickerood, H. K. Court,
D. K. Dabney, J. G. Edwards, S. V. Fend, S. S. Ferguson, W. E. Ferguson, M. A. Garcia-Vidal, A.
Hom, D. H. Kavanaugh, V. F. Lee, G. J. Mallick, J. D. McCarty, D. L. Mead, N. D. Penny, W. J.
Pulawski, E. S. Ross, H. I. Scudder, R. E. Stecker, C. Y. Takahashi, S. P. Taormino, W. A. Titherington,
and D. Ubick; (22 guests) V. Ahrens-Pulawski, M. M. Arnaud, A. Bandar, R. Bandar, L. Berke, L.
Bezark, J. E. Court, H. V. Davis, J. M. Donald, K. S. Hom, J. Johnston, A. Jung, N. Kostek, G. Y.
Leung, P. Myers, A. M. L. Penny, M. Porter, W. C. Rauscher, S. Renkes, S. Ross, and 2 unsigned
guests.
FouR HUNDRED AND NINETY-FOURTH MEETING
The 494th meeting of the Pacific Coast Entomological Society was held on 18 September 1992, at
8:00 PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Norman D. Penny presiding.
The minutes of the 15 May 1992 meeting were read and accepted. Ten persons were proposed and
elected as new members: Ms. Aljona Andrejeff, Mr. Charles M. Brandau, Mr. Ronald L. Cauble, Mr.
Larry V. Desmond, Dr. Judith Najt, Mr. Martin G. Rigby, and Mr. Ron Wilson as regular members;
and Mr. Damian S. Bruss, Ms. Paula M. Hartgraves, and Ms. Mozelle M. Langford as studentmembers.
1996 PROCEEDINGS 107
Dr. Edward L. Smith introduced Dr. Helmut Sturn of Universitat Hildesheim and mentioned that
Dr. Sturn has described several new local species of archaeognathans. Dr. Penny announced that
agricultural inspectors had recently found a Mediterranean fruit fly in Pasadena, CA. Mr. Curtis Y.
Takahashi added that authorities had confirmed seven medflies in Pasadena, as well as a Mexican
fruit fly. Dr. Penny also mentioned a report which appeared in newspapers during the summer involving
a brown recluse spider bite in southern California.
Dr. Smith presented a note concerning winglessness in primitive insects in relation to other arthopods
and discussed phylogenetic hypotheses based on morphological and other evidence from fossil and
extant species.
The featured speaker Dr. David H. Kavanaugh of the California Academy of Sciences presented a
slide lecture entitled “Collecting Undisturbed Hawaii.” He discussed his collecting and biogeographic
study of the carabid genus Bembidion (Coleoptera: Carabidae).
The meeting adjorned at 9:30 pm, followed by a social hour in the Entomology Department con-
ference room.—D. K. Dabney, Recording Secretary.
The following 49 persons were present. 29 members: P. H. Arnaud Jr., V. M. Barlow, T. S. Briggs,
R. M. Brown, R. L. Cauble, H. K. Court, D. K. Dabney, W. A. Doolin, J. G. Edwards, W. Hamersky,
D. H. Kavanaugh, S. T. Kawaguchi, B. Keh, V. F. Lee, N. D. Penny, J. A. Powell, M. Prentice, W.
J. Pulawski, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, W. E. Savary, H. I. Scudder,
E. L. Smith, L. A. Solorzano, R. E. Stecker, C. Y. Takahashi, and S. P. Taormino; (20 guests) V.
Ahrens-Pulawski, M. M. Arnaud, S. Arnaud, M. Caterino, J. E. Court, E. Goff, P. M. Hartgraves, M.
Hannaford, M. M. Langford, T. Meikle, R. Morgan, J. M. Parinas, A. M. L. Penny, C. Prentice, J.
Robertson, N. Kukich, H. Sturm, D. Tom, and 2 unsigned guests.
FouR HUNDRED AND NINETY-FIFTH MEETING
The 495th meeting of the Pacific Coast Entomological Society was held on 16 October 1992, at
8:00 PM, in the Goethe Room of the California Academy ofSciences, Golden Gate Park, San Francisco,
with President Dr. Norman D. Penny presiding.
The minutes of the 18 September 1992 meeting were read and accepted. Three persons were proposed
and elected as new members: Mr. Keith F. Dobry and Mr. Joseph Martin as regular members, and
Mr. Morgan J. Hannaford as student member.
Dr. C. Don MacNeill introduced Mr. Mike Collins. Mr. W. E. Savary introduced Mr. Al Olsen of
the U. S. Food and Drug Administration. Dr. Penny introduced Ms. Julieta Parinas and welcomed
Mr. Derham Giuliani of Big Pine, California.
Dr. Paul H. Arnaud announced that the Entomology Department at the Academy had received a
children’s pop-up book about spiders and had the book available for viewing during the social hour
following the meeting. Dr. Penny announced the publication and availability of the book Life in Amber
by G. Poinar.
The featured speaker Mr. K. J. Ribardo of the California Academy of Sciences presented a slide
lecture entitled ““Natural History Travels in Panama.” He discussed his recent collecting trip to Panama,
giving details on traveling and collecting in the area.
The meeting adjorned at 9 pm, followed by a social hour in the Entomology Department conference
room.—D. K. Dabney, Recording Secretary.
The following 49 persons were present. 32 members: P. H. Arnaud Jr., V. M. Barlow, T. S. Briggs,
K. W. Brown, J. S. Cope, H. K. Court, D. K. Dabney, C. E. Griswold, W. Hamersky, M. J. Hannaford,
Y.-F. Hsu, M. M. Langford, R. L. Langston, V. F. Lee, C. D. MacNeill, W. A. Maffei, M. J. O’Halloran,
S. T. O’Keefe, A. R. Olsen, S. B. Opp, N. D. Penny, A. B. Rackett, J. M. Ribardo, K. J. Ribardo, L.
S. Saul, W. E. Savary, K. A. Schwarz, H. I. Scudder, L. A. Solorzano, C. Y. Takahashi, D. Ubick,
and R. A. Worth; (17 guests) M. M. Arnaud, D. Carbon, M. Caterino, M. Collins, S. Cope, J. E. Court,
D. Giuliani, N. Kukich, T. Meikle, R. Morgan, D. O’Halloran, A. Olsen, J. M. Parinas, A. M. L.
Penny, J. Schweikert, D. Sear, and C. Tom.
FouR HUNDRED AND NINETY-SIXTH MEETING
The 496th meeting of the Pacific Coast Entomological Society was held on 20 November 1992, at
8:00 PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Norman D. Penny presiding.
The minutes of the 16 October 1992 meeting were read and accepted. Seven persons were proposed
and elected as new members: Mr. Paul Belluomini as a sponsoring member; Dr. Mary Carver, Dr.
108 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Maria Luisa Castillo, Dr. Walter H. Sakai, and Mr. Jere Schweikert as regular members; and Ms. Lee
A. Norton and Mr. Stanley E. Vaughn as student members.
Dr. Wojciech J. Pulawski introduced Dr. Eduardas Budrys from Lithuania. Dr. Ronald E. Stecker
introduced Mr. Lawrence Garibaldi, Ms. Lee A. Norton, Mr. Stanley E. Vaughn, and Mr. Dan Wolfe,
from San Jose State University. Mr. D. Keith Dabney introduced Mr. José Manuel Pino Moreno from
Mexico and announced that Mr. Pino would be offering two books about edible insects for sale during
the social hour.
Dr. Edward L. Smith announced the formation of a new entomological society The International
Society of Hymenopterists, and showed a copy of their journal with articles on Hymenoptera phylogeny
and on the use of SEM in entomology.
Mr. José M. Pino Moreno presented a note with slides describing the study of edible insects in
Mexico.
Dr. Norman D. Penny announced that the society has an IBM PC for sale. Dr. Penny also announced
that the society was seeking members to serve as officers in the positions of President, President Elect,
Treasurer, Managing Secretary, and Recording Secretary for 1993, and that he had formed a nominating
committee composed of Drs. Paul H. Arnaud Jr., John T. Doyen, and Marius S. Wasbauer.
The featured speaker Dr. Wojciech J. Pulawski of the California Academy of Sciences presented a
slide lecture entitled ““A Study of World Gastrosericus.” He discussed his approach to studying the
systematics of these wasps and included information on the use of natural history and biogeography
in systematics.
The meeting adjourned at 9:50 PM, followed by a social hour in the Entomology Department
conference room.—D. K. Dabney, Recording Secretary.
The following 46 persons were present. 35 members: V. M. Barlow, L. G. Bezark,-D. L. J. Boutin,
T. S. Briggs, R. M. Brown, P. Buickerood, R. Buickerood, J. S. Chinn, D. K. Dabney, S. S. Ferguson,
W. E. Ferguson, C. E. Franklin III, C. E. Griswold, W. Hamersky, A. P. Jarman, S. T. Kawaguchi,
M. M. Langford, R. L. Langston, V. F. Lee, G. J. Mallick, S. B. Opp, N. D. Penny, M. Prentice, A.
E. Rackett, J. M. Ribardo, K. J. Ribardo, J. Schweikert, H. I. Scudder, E. L. Smith, R. E. Stecker, C.
Y. Takahashi, D. Ubick, S. E. Vaughn, J. S. Wasbauer, and M. S. Wasbauer; (11 guests) V. Ahrens-
Pulawski, E. Budrys, L. Garibaldi, T. Meikle, L. A. Norton, J. M. Parinas, A. M. L. Penny, J. M.
Pino Moreno, W. Rauscher, K. Reynolds, and D. Wolfe.
FouR HUNDRED AND NINETY-SEVENTH MEETING
The 497th meeting of the Pacific Coast Entomological Society was held on 11 December 1992, at
8:00 PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Norman D. Penny presiding.
The minutes of the 20 November 1992 meeting were read and accepted. Five persons were proposed
and elected as new members: Ms. Virginia L. Scott and Prof. Dr. Helmut Sturm as regular members;
and Mr. Mikael Niehoff, Ms. Kathy Reynolds, and Mr. Daniel Wolfe as student members.
Committee members read reports from the auditing, historical, membership, nominating, and -
publication committees.
Mrs. Helen K. Court read the auditing committee report, announcing that the auditing committee,
under the chairmanship of Mr. H. Vannoy Davis, CPA, examined the financial records of the society
and found everything in order. A statement of Income Expenditures and Changes in Fund Balances
will be published in the proceedings.
Dr. Paul H. Amaud Jr. read the historical committee report, announcing that Dr. Kenneth E. Frick
had posthumously donated his correspondence files, and that Dr. Frick had had previously donated
his agromyzid collection to CAS in 1960.
Mr. Curtis Y Takahashi read the membership committee report, announcing that the society added
23 new regular members, 17 new student members, and | new sponsoring member, for a total of 41
new members, which is the lowest number in recent years.
D. K. Dabney read the 1992 slate of officers for Dr. J. T. Doyen, chairman of the nominating
committee, and proposed Dr. Susan B. Opp as President, Dr. Kirby W. Brown as President-elect, Dr.
Paul H. Arnaud Jr., as Treasurer, Mr. Vincent F. Lee as Managing Secretary, and Mr. Keve J. Ribardo
as Recording Secretary. All members present unanimously approved the slate. Dr. Penny read an-
nounced that the fourth number of The Pan-Pacific Entomologist for 1992 had been mailed and that
Dr. John E. Hafernik Jr. is the new chair of the publication committee.
Dr. Robbin W. Thorp announced that the 1993 Bohart Museum calendars featuring SEM photo-
graphs by the late Mr. Robert O. Schuster are available for sale.
1996 PROCEEDINGS 109
Dr. K. W. Brown presented a slide illustrated note about the varroa mite (Varroa), a pest of the
honey bee.
The featured speaker Dr. Norman D. Penny of the California Academy of Sciences presented the
presidential lecture entitled ‘“‘Use of Computers in Systematic Research and Collection Management.”
He discussed the use of computers in the Academy’s collection and in insect systematics, and then
opened the meeting to discussion of the topic.
The meeting adjourned at 9:00 PM, followed by a social hour in the Entomology Department
conference room.—D. K. Dabney, Recording Secretary.
The following 30 persons were present. 27 members: P. H. Arnaud Jr., K. W. Brown, R. M. Brown,
H. K. Court, D. K. Dabney, W. A. Doolin, W. E. Ferguson, J. E. Hafernik Jr., A. Hom, M. M.
Langford, R. L. Langston, V. F. Lee, W. A. Maffei, L. A. Norton, S. B. Opp, N. D. Penny, A. E.
Rackett, K. Reynolds, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, W. E. Savary, L. E. Serpa, R.
E. Stecker, C. Y. Takahashi, R. E. Thorp, and S. E. Vaughn; (3 guests) M. M. Arnaud, J. E. Court,
and A. M. L. Penny.
PAN-PACIFIC ENTOMOLOGIST
72(2): 109-114, (1996)
PROCEEDINGS OF THE PACIFIC COAST
ENTOMOLOGICAL SOCIETY, 1993
FoUR HUNDRED AND NINETY-EIGHTH MEETING
The 498th meeting of the Pacific Coast Entomological Society was held on 15 January 1993, at
8:00 PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Susan B. Opp presiding.
The minutes of the 11 December 1992 meeting were read and accepted. Two persons were proposed
and elected as new members: Dr. Diomedes Quintero Arias as a regular member, and Ms. Maria
Elena Resendiz Ruiz as a student member.
Dr. Opp thanked Dr. Norman D. Penny and congratulated him on a job well done as president of
PCES for 1992. She announced that the program committee consists of Ms. Leslie S. Saul and Dr.
Ronald E. Stecker. The historical committee will be chaired by Dr. David H. Kavanaugh.
Dr. Penny introduced Dr. Paul J. Johnson from the University of Wisconsin and Dr. Paul H. Arnaud
Jr. introduced Mr. Gary L. Peters from Oregon. Dr. Penny announced that the entomology department
of the California Academy of Sciences has insect drawers, unit trays, schmitt boxes, and envelope
boxes for sale.
Mr. Warren E. Savary of the U.S. Food and Drug Administration presented a note on the erotylid
beetle Dacne picta Crotch, a recently introduced pest of stored, dried shiitake mushrooms.
The speaker Dr. Carol Boggs of Stanford University presented a slide lecture entitled “Resources,
Reproduction, and Male Nutrient Donations in Insects.” Dr. Boggs described interconnections among
foraging, allocation, and life history strategies.
The meeting adjourned at 9:35 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 33 persons were present, (25 members), P. H. Arnaud, Jr., L. G. Bezark, T. S. Briggs,
R. M. Brown, J. S. Chinn, C. D. Franklin II, C. E. Griswold, J. E. Hafernik, Jr., W. Hamersky, P. J.
Johnson, M. M. Langford, V. F. Lee, S. B. Opp, N. D. Penny, G. L. Peters, K. Reynolds, J. M.
Ribardo, K. J. Ribardo, R. G. Robertson, L. S. Saul, W. E. Savary, J. Schweikert, L. A. Solorzano,
C. Y. Takahashi, and S. E. Vaughn; (8 guests) M. M. Arnaud, W. Baumgantl, C. Boggs, T. Meikle, J.
F. Parinas, A. M. L. Penny, W. E. Rauscher, and C. Tom.
110 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
FouR HUNDRED AND NINETY-NINTH MEETING
The 499th meeting of the Pacific Coast Entomological Society was held on 19 February 1993, at
8:00 PM, in the Goethe Room ofthe California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Susan B. Opp presiding.
The minutes of the 15 January 1993 meeting were read and accepted. Four persons were proposed
and elected as new members: Ms. Hariana Chilstrom as a regular member; Ms. Ana Maria Lira Penny
as a family member; and Mr. Leo T. Margolf and Mr. Mohammed A. Noor as student members.
Dr. Opp announced that Ms. Sandy S. Shanks needs live bumble bee queens. She also mentioned
that in March 1993, the Pacific Coast Entomological Society will hold its 500th meeting. She suggested
the Society celebrate this event by having a catered dinner at the California Academy of Sciences,
possibly in the Steinhart Aquarium.
Mr. Michael S. Caterino announced that Dr. Jarmila Kukalova-Peck would be speaking at the
University of California, Berkeley, on Monday, 22 February 1993, at 4:00 PM on fossil Paleozoic
insects and their contribution to understanding insect evolution. A flyer will be available in the
conference room after the meeting.
Dr. Norman D. Penny announced that a new preserve was being created near the Jatun Sacha
Biological Station in eastern Ecuador. Interested parties should contact Dr. Edward S. Ross of the
Entomology Department of the California Academy of Sciences for more information and about
contributions to help save this vanishing rainforest.
Ms. Leslie S. Saul announced that the San Francisco Insect Zoo had acquired and is displaying a
live Malaysian Flower Mantis, she encouraged everyone to see it.
Mr. Vincent F. Lee announced that Mr. Dexter Sear started an organization called IO Vision to
promote cultural entomology.
Dr. Ronald E. Stecker introduced Mr. Pyon Cho from Rangoon, Burma.
Mr. Curtis Y. Takahashi of the California State Department of Food and Agriculture presented a
note on the Sweet Potato Weevil, Cylas formicarius (Fabricius), which is an ‘“‘A”’ rated pest that has
recently been reported from Merced County where sweet potatoes are a 35 million dollar industry.
The speaker Dr. Douglas Whitman of Illinois State University presented a slide lecture entitled
“How Insects Defend Themselves Against Predators.” Dr. Whitman explained that because so many
other organisms eat insects, including insects themselves, they have evolved many kinds of strategies
to avoid being eaten, including chemical defense, mimicry, crypsis, and unpalatability.
The meeting adjourned at 9:15 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 49 persons were present. (34 members): P. H. Amaud, Jr., L. G. Bezark, T. S. Briggs,
M. S. Caterino, J. G. Edwards, E. M. Fisher, C. D. Franklin II, C. E. Griswold, W. Hamersky, A.
Hom, B. Keh, R. L. Langston, V. F. Lee, G. J. Mallick, S. T. O’Keefe, S. B. Opp, A. M. L. Penny,
N. D. Penny, A. B. Rackett, K. Reynolds, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, L. S. Saul,
K. A. Schwarz, J. Schweikert, R. E. Stecker, C. Y. Takahashi, S. E. Vaughn, C. E. Warren, D. W.
Whitman, D. Wolfe, R. A. Worth and R. L. Zuparko; (15 guests) M. M. Amaud, A. M. Castellanos,
P. Cho, R. Kawin, T. Meikle, R. Pelletier, Jr.. W. E. Rauscher, M. Slawinski, D. Whitman, K. S.
Whitman, L. S. Whitman, R. Whitman, T. Zaviezo, and 2 illegible signatures.
FIVE HUNDREDTH MEETING
The 500th meeting of the Pacific Coast Entomological Society was held on 19 March 1993, at 8:00
PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Susan B. Opp presiding.
The minutes of the 19 February 1993 meeting were read and accepted. Three persons were proposed
and elected as new members: Mr. Andreas Crameri and Dr. Robert D. Waltz as regular members,
and Mr. Robert Pelletier Jr. as a student member.
Dr. Sandy Ross announced that a new preserve at and near Cabanas Alifiahui, Tena, Ecuador was
being created. Members and friends of the Pacific Coast Entomological Society were encouraged to
donate or pledge towards preserving rainforest habitat at this species-rich site. Dr. Opp proposed that
the Society lend its moral support for the formation of the Alifiahui preserve. The motion was approved
by the majority of members present.
For the occasion of the 500th meeting of the Pacific Coast Entomological Society, Dr. Paul H.
1996 PROCEEDINGS 111
Arnaud, Jr. presented a series of slides and discussions on most of the 22 persons and the one Society
that were the founding Charter members of the Society. This included Mr. Carl Fuchs (1839-1914),
Prof. Vernon L. Kellogg (1867-1937), Dr. Frank E. Blaisdell (1862-1946), Dr. Edwin C. Van Dyke
(1869-1952), Prof. Henry C. Fall (1862-1939), Mr. Lucius E. Ricksecker (1841-1913), Dr. Hans
Herman Behr (1818-1904), Mr. W. G. W. Harford (1825-1911), Mr. Beverly Letcher (1865-1905),
Mr. Frederick W. Nunenmacher (1870-1946), Dr. William H. Ashmead (1855-1908), Mr. Edwin M.
Ehrhorn (1862-1941), Mr. James E. Cottle (1861-1935), Mr. Julius G. Grundel (1857-1933), Prof.
James J. Rivers (1824-1913), Dr. Leland O. Howard (1857-1950), the Grattum Naturalists Club of
Eldorado, California. Pictures and information were not available for C. A. Whiting, Newton B. Pierce,
C. W. Herr, George W. Harney, F. C. Clark, and L. Dustein.
Mr. Carl Fuchs had issued a call for the organization of an Entomological Society on August 7,
1901, and the first meeting was held at the California Academy of Sciences, on Market Street, in San
Francisco, on August 15, 1901, at 2 o’clock. The Society was first named the “‘California Entomology
Club” but the name was changed to the “Pacific Coast Entomological Society” at the society’s 5th
meeting held on August 16, 1902.
The speaker Dr. Edward S. Ross of the California Academy of Sciences presented a slide lecture
entitled “Surprise Insect Encounters in South America.” Dr. Ross showed a selection of slides dem-
onstrating the great beauty and almost infinite variety of plants and animals of the neotropical rain-
forest.
The meeting adjourned at 10:15 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 106 persons were present. (58 members): F. G. Andrews, P. H. Armaud Jr., R. L.
Brett, T. S. Briggs, R. M. Bohart, K. W. Brown, R. M. Brown, P. Buickerood, R. Buickerood, M. S.
Caterino, H. Chilstrom, J. R. Clopton, H. K. Court, A. Crameri, J. G. Edwards, T. D. Eichlin, S. S.
Ferguson, W. E. Ferguson, E. M. Fisher, C. D. Franklin II, C. E. Griswold, K. S. Hagen, W. Hamersky,
M. J. Hannaford, A. Hom, D. H. Kavanaugh, M. M. Langford, C. D. MacNeill, G. J. Mallick, L. A.
Norton, S. T. O’Keefe, S. B. Opp, R. Pelletier Jr., A. M. L. Penny, N. D. Penny, A. B. Rackett, V.
H. Resh, K. Reynolds, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, E. S. Ross, W. E. Savary, K.
A. Schwarz, J. Schweikert, W. D. Shepard, E. L. Smith, R. E. Somerby, J. T. Sorensen, R. E. Stecker,
C. Y. Takahashi, D. Ubick, S. E. Vaughn, J. S. Wasbauer, M. S. Wasbauer, D. Wolfe, R. A. Worth,
and R. L. Zuparko; (48 guests) J. Adams, M. M. Arnaud, A. Bandar, R. Bandar, C. B. Barr, M. E. R.
Bohart, S. Brown, K. Bunnell, J. E. Court, M. H. Crameri, H. V. Davis, J. Edwards, R. Ferguson, E.
Goff, T. Guenette, M. D. Hagen, C. Haigh, K. Kipping, T. Kipping, V. Kipping, S. Marino, D.
McGinnis, D. S. McInnes, T. C. Meikle, H. Nevins, C. Nufio, J. F. Parinas, G. Prlain, W. E. Rauscher,
J. Robertson, S. M. Ross, M. Rutherford, R. Schryrer, D. Smith, S. Smith, K. H. Sorensen, K. Sprague,
P. E. Stecker, B. Stewart, P. Svensson, R. Takumi, P. Thompson, C. Vaughn, R. Wilson, R. J. Worth,
B. Yew, T. Zaviezo, and C. Zuparko.
FIVE HUNDRED FIRST MEETING
The 5Ol1st meeting of the Pacific Coast Entomological Society was held on 16 April 1993, at 8:00
PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Susan B. Opp presiding.
The minutes of the 19 March 1993 meeting were read and accepted. Two persons were proposed
and elected as new members: Mr. Glenn K. Goodwin as a regular member and Ms. Teresa C. Meikle
as a family member.
The speaker Dr. Deborah Letourneau of the University of California, Santa Cruz, presented a slide
lecture entitled ““Trophic Cascades: Ant-Plants and Their Associates in Tropical Forests.” Dr. Le-
tourneau showed the important interrelationships between certain plants and their ants, in Costa Rica.
The meeting adjourned at 9:15 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 32 persons were present. (21 members): P. H. Arnaud Jr., V. M. Barlow, T. S. Briggs,
N. E. Gershenz, C. E. Griswold, W. Hamersky, R. L. Langston, V. F. Lee, L. A. Norton, S. B. Opp,
A. M. L. Penny, N. D. Penny, A. B. Rackett, K. A. Reynolds, L. S. Saul, W. E. Savary, J. Schweikert,
R. E. Stecker, C. Y. Takahashi, D. Ubick, and S. E. Vaughn; (11 guests) M. M. Amaud, P. Cho, C.
Craig, D. Farison, B. Gottlieb, L. Harnish, D. Letourneau, T. C. Meikle, W. E. Rauscher, M. Tokar-
zewski, and W. Wood.
112 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
FIVE HUNDRED SECOND MEETING
The 502nd meeting of the Pacific Coast Entomological Society was held on 17 September 1993, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Susan B. Opp presiding.
The minutes of the 16 April 1993 meeting were read and accepted. Sixteen persons were proposed
and elected as new members: Mr. Ernest Anderson as a sponsoring member; Dr. Harry E. Andersen,
Mr. Robert P. Apsey, Mr. Terry L. Bishop, Dr. Michael W. J. Crosland, Dr. Bryan N. Danforth, Dr.
William G. Eberhard, Dr. Hanif Gulmahamad, Mr. Donald G. Harrington, Mr. Katsuya Ichinose,
Mr. Richard G. Little, Dr. Robert E. Roughley, Dr. Deborah A. Waller, and Dr. Jiri Zidek as regular
members; and Mr. Darin B. Allred and Mr. John C. Herr as student members.
Dr. Opp introduced and welcomed Dr. Richard M. Bohart, Dr. J. Gordon Edwards, Dr. Kenneth
S. Hagen, Dr. Harry W. Lange, Dr. Edward S. Ross, and Dr. Elwood C. Zimmerman as honorary
members of the Society. She presented Mr. Vincent F. Lee with a special award in recognition of his
many years of service to the society.
Dr. Opp announced that the executive board voted in favor of raising member dues and publication
fees in the journal for the coming year.
Mr. Lee announced that duplicate reprints will be available for sale in the entomology department
conference room, following the meeting.
Dr. Opp called for members and interested students to consider sponsoring a symposium, workshop,
or contributed paper section at the 1994 American Association for the Advancement of Science Pacific
Division meeting which will be held 19-23 June at San Francisco State University.
Dr. Norman D. Penny circulated a container with several giant lacewings (Polystoechotidae) which
were recently collected in the Sierra Nevada.
The speaker Dr. Jenella Loye of the University of California, Davis, presented a slide lecture entitled
“Blood Feeding Maggots That Live With Birds.” Dr. Loye observed that birds avoid highly parasitized
nests and described how parasites utilize their food source and, in turn, the host adaptive response
to parasitism.
The meeting adjourned at 9:35 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 49 persons were present. (38 members): P. H. Arnaud Jr., L. G. Bezark, R. M. Bohart,
K. W. Brown, P. Buickerood, R. Buickerood, M. S. Caterino, H. K. Court, P. G. da Silva, D. K.
Dabney, W. A. Doolin, J. G. Edwards, C. D. Franklin II, P. H. Freytag, M. J. Hannaford, R. L.
Langston, V. F. Lee, L. A. Norton, S. B. Opp, A. M. L. Penny, N. D. Penny, W. J. Pulawski, A. B.
Rackett, K. A. Reynolds, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, E. S. Ross, W. E. Savary,
H. I. Scudder, W. D. Shepard, R. E. Stecker, C. Y. Takahashi, J. E. Tobler I, S. E. Vaughn, J. S.
Wasbauer, M. S. Wasbauer, and R. L. Zuparko; (11 guests) C. Barr, S. Carroll, S. A. Casari Chen, J.
E. Court, K. Graham, S. Hammond, J. Loye, J. J. McNicol, J. Robertston, A. Telang, and S. Warnock.
FIVE HUNDRED THIRD MEETING
The 503rd meeting of the Pacific Coast Entomological Society was held on 15 October 1993, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Susan B. Opp presiding.
The minutes of the 17 September 1993 meeting were read and accepted. Seven persons were proposed
and elected as new members: Mr. J. Michael Poellot as a sponsoring member; Dr. Nicholas J. Mills
and Mr. Shawn A. Steffan as regular members; and Mr. Richard J. Adams, Ms. Sha Hammond, Ms.
Jeanette J. McNicol, and Ms. Aparna Telang as student members.
Dr. Norman D. Penny introduced Dr. Derrick Blocker, Mr. Armando Equihua, and Dr. Bradley
Sinclair, who were visiting scientists at the California Academy of Sciences Entomology Department.
It was announced that the Entomology Club at the University of California, Berkeley, is selling
T-shirts for 12 dollars each.
Ms. Leslie S. Saul announced that the Center for the Arts at Yerba Buena Gardens had put in new
plants to specifically attract butterflies.
Dr. Penny announced that the Department of Entomology at the California Academy of Sciences
had two curatorial assistant positions available.
The speaker Dr. Scott Carroll of UC Davis presented a slide lecture entitled ‘““Adaptive Radiation
in Soapberry Bugs, the New Darwin’s Finches: Evolution in Action!” Dr. Carroll studied the beak
1996 PROCEEDINGS 113
lengths of soapberry bugs on balloon-vines, golden raintree and silkberrys and discussed other host
race adaptations to fitness.
The meeting adjourned at 9:30 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 43 persons were present. (30 members): T. S. Briggs, K. W. Brown, M. S. Caterino,
P. G. da Silva, S. S. Ferguson, W. E. Ferguson, N. E. Gershenz, C. E. Griswold, W. Hamersky, S.
Hammond, V. F. Lee, G. J. Mallick, T. E. Meikle, L. A. Norton, S. B. Opp, A. M. L. Penny, N. D.
Penny, J. A. Powell, A. B. Rackett, K. A. Reynolds, R. G. Robertson, L. S. Saul, W. E. Savary, J.
Schweikert, R. E. Somerby, C. Y. Takahashi, A. Telang, J. E. Tobler I, D. Ubick, and S. E. Vaughn;
(13 guests) H. D. Blocker, S. M. Carroll, A. Equihua, D. D. Giuliani, K. Graham, K. Kocurek, M.
McIntosh, S. S. Mead, M. Orr, W. E. Rauscher, J. Robertson, B. J. Sinclair, and W. Veder.
FIVE HUNDRED FOURTH MEETING
The 504th meeting of the Pacific Coast Entomological Society was held on 19 November 1993, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Susan B. Opp presiding.
The minutes of the 15 October 1993 meeting were read and accepted. One person was proposed
and elected as a student member: Ms. Tunyalee Morisawa.
Dr. Norman D. Penny announced that the Department of Entomology at the California Academy
of Sciences had filled the two full-time curatorial assistant positions.
Dr. Ronald E. Stecker presented a slide-illustrated note on the egg masses of Sialidae or alderflies
encountered in Alum Rock Park in the spring.
The speaker Dr. Deborah Gordon of Stanford University presented a slide lecture entitled ‘““The
Population Consequences of the Development of Ant Colony Behavior.” Dr. Gordon studied the
dynamic nature of ant colonies and the effect on their population as it relates to the division of labor
in seed harvesting desert ants. Dr. Gordon showed how behavioral changes occur as these ant colonies
age and grow.
The meeting adjourned at 9:40 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 48 persons were present. (38 members): L. G. Bezark, T. S. Briggs, K. W. Brown, R.
M. Brown, H. K. Court, D. K. Dabney, B. Deutsch, J. G. Edwards, S. V. Fend, C. D. Franklin II, J.
E. Hafnernik Jr., W. Hamersky, S. Hammond, C. Y. Kitayama, R. L. Langston, V. F. Lee, G. J.
Mallick, L. A. Norton, S. T. O’Keefe, S. B. Opp, A. M. L. Penny, N. D. Penny, A. B. Rackett, S.
Renkes, K. A. Reynolds, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, W. E. Savary, J. Schweikert,
H. I. Scudder, R. E. Stecker, C. Y. Takahashi, A. Telang, J. E. Tobler I, D. Ubick, S. E. Vaughn, and
S. P. Welles Jr.; (10 guests) I. Brown, S. M. Carroll, J. E. Court, L. Culp, D. Gordon, C. Hironymous,
J. Loye, W. E. Rauscher, J. Robertston, and T. Vich.
FIVE HUNDRED FIFTH MEETING
The 505th meeting of the Pacific Coast Entomological Society was held on 10 December 1993, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Susan B. Opp presiding.
The minutes of the 19 November 1993 meeting were read and accepted. Four persons were proposed
and elected as new members: Dr. James A. Bethke, Mr. John A Calderwood, and Mr. Dick Fagerlund
as regular members; and Mr. Stanley A. Wright as a student member.
Mrs. Helen K. Court gave the auditing committee report announcing that the Mr. H. Vannoy Davis,
CPA, was still reviewing the societies finances.
Mr. Curtis Y. Takahashi read the membership committee report, announcing that the Society added
23 new regular members, 2 new family members, 12 new student members, and 2 new sponsoring
members, for a total 49 new members in 1993.
President Opp gave the nominating committee report, and the following persons were proposed
and accepted as officers for 1994: Mr. Curtis Y. Takahashi as President-elect, Dr. Paul H. Arnaud,
Jr. as Treasurer, Mr. Vincent F. Lee as Managing Secretary, and Mr. Keve J. Ribardo as Recording
Secretary.
Dr. Opp passed the gavel over to incoming president Dr. Kirby W. Brown.
114 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
Mr. Vincent F. Lee announced that the fourth number of the Pan Pacific Entomologist would be
mailed shortly and reminded members to pay their dues for 1994.
Mr. D. Keith Dabney presented a slide-illustrated note on some habitats and insects of east Maui,
Hawaii.
The speaker Dr. Susan B. Opp of California State University, Hayward, presented a slide lecture
entitled “Control of Walnut Husk Fly in California: What Have We Learned from the Apple Maggot
Fly.”” Dr. Opp compared control programs of these two pest species and discussed plans for future
control of Walnut Husk Fly. Dr. Opps student, Mr. William Hamersky then took the podium to
discuss tests of Walnut Husk Fly’s attraction to volatile compounds of the walnut.
The meeting adjourned at 9:10 PM, followed by a social hour in the entomology department
conference room.—K. Ribardo, Recording Secretary.
The following 40 persons were present. (28 members): R. L. Aalbu, P. H. Amaud Jr., F. L. Blanc,
T. S. Briggs, K. W. Brown, M. S. Caterino, H. K. Court, D. K. Dabney, P. G. da Silva, E. M. Fisher,
W. Hamersky, S. Hammond, J. C. Herr, R. L. Langston, V. F. Lee, J. J. McNicol, L. A. Norton, S.
T. O’Keefe, S. B. Opp, K. A. Reynolds, R. G. Robertson, L. S. Saul, H. I. Scudder, N. J. Smith, C.
Y. Takahashi, A. Telang, D. Ubick, and S. E. Vaughn; (12 guests) M. M. Amaud, M. Baiznon, F.
Blanc, P. Coogan, J. E. Court, L. Culp, R. Derasary, K. Graham, W. E. Rauscher, J. Robertson, J. A.
Tarralba, and W. L. Yee.
PAN-PACIFIC ENTOMOLOGIST
72(2): 114-120, (1996)
PROCEEDINGS OF THE PACIFIC COAST
ENTOMOLOGICAL SOCIETY, 1994
FIVE HUNDRED SIXTH MEETING
The 506th meeting of the Pacific Coast Entomological Society was held on 21 January 1994, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Kirby W. Brown presiding.
The minutes of the 10 December 1993 meeting were read and accepted. Five persons were proposed
and elected as new members: Dr. Richard J. Bell, Mr. Walter J. Bentley, Dr. John G. O’Reilly, and
Ms. Jody McWilliams as regular members; and Mr. Derek S. Sikes as a student member.
Dr. Brown encouraged members to bring in notes and exhibits.
Two guests were introduced: Mr. Scott E. Haskins and Dr. Yu-hua Yan.
The featured speaker Dr. Rolf L. Aalbu of the California Department of Food and Agriculture
presented a slide lecture entitled “Environmental Assessment and IPM Consulting In Africa: A Maur-
itanian Adventure.’ He discussed Project Oasis and showed how crops have to contend with not only
lack of water, but insect pests, plant diseases, and sand dune encroachment. Mesquite trees are now
being grown to help in stabilizing sand dunes. Dr. Aalbu observed that scale insects are a very serious
pest and that palm scale infests all palms.
The meeting adjourned at 9:00 PM, followed by a social hourin the entomology conference room. —
K. Ribardo, Recording Secretary.
The following 37 persons were present. 29 members: F. G. Andrews, P. H. Amaud Jr., T. S. Briggs,
K. W. Brown, R. M. Brown, M. S. Caterino, J. S. Chinn, D. K. Dabney, P. G. da Silva, W. A. Doolin,
E. M. Fisher, C. E. Griswold, S. Hammond, R. L. Langston, J. D. McCarty, T. C. Meikle, L. A.
Norton, A. M. L. Penny, N. D. Penny, A. E. Rackett, K. A. Reynolds, J. M. Ribardo, K. J. Ribardo,
W. E. Savary, K. N. Schick, C. Y. Takahashi, A. Telang, D. Ubick, and S. E. Vaughn; (8 guests) M.
M. Amaud, C. B. Barr, S. E. Haskins, J. F. Parinas, W. E. Rauscher, J. Schick, Y.-h. Yan, and T.
ZavieZo.
1996 PROCEEDINGS 115
FIVE HUNDRED SEVENTH MEETING
The 507th meeting of the Pacific Coast Entomological Society was held on 18 February 1994, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Kirby W. Brown presiding.
The minutes of the 21 January 1994 meeting were read and accepted. Four persons were proposed
and elected as new members: Mr. Donald W. Darling and Ms. Dana C. Russell as regular members;
and Mr. Mark A. Isaak and Mr. Steven E. Jungers as student members.
Dr. Brown encouraged members to bring in notes and exhibits.
Dr. Susan B. Opp announced that the Pacific Coast Entomological Society is sponsoring an Ento-
mology section at the Pacific Division of AAAS. Dr. Opp especially encouraged students to give talks
or poster sessions.
The featured speaker Mr. Raymond J. Gill of the California Department of Food and Agriculture
presented a slide lecture entitled ““New Homopterans in California.’’ He noted that homopterans have
accounted for three times as many new introductions as all other insects combined. The 27 introduced
homopterans included such agriculturally important pests as the sweet potato whitefly, ash whitefly,
and silverleaf whitefly. Mr. Gill explained that pesticide spraying was not usually effective and that
problem infestations are usually turned over to biological control officers.
The meeting adjourned at 9:30 PM, followed by a social hour in the entomology conference room. —
K. Ribardo, Recording Secretary.
The following 52 persons were present. 33 members: R. L. Aalbu, P. H. Arnaud Jr., L. G. Bezark,
T. S. Briggs, K. W. Brown, R. M. Brown, M. S. Caterino, H. K. Court, D. K. Dabney, P. G. da Silva,
B. Deutsch, B. K. Eya, C. E. Griswold, S. Hammond, R. L. Langston, V. F. Lee, T. C. Meikle, L. A.
Norton, S. T. O’Keefe, S. B. Opp, A. M. L. Penny, N. D. Penny, A. E. Rackett, K. A. Reynolds, K.
J. Ribardo, R. G. Robertson, W. E. Savary, J. Schweikert, L. A. Solorzano, R. E. Somerby, C. Y.
Takahashi, D. Ubick, and S. E. Vaughn; (19 guests) M. M. Amaud, J. Brown, S. Brown, J. E. Court,
J. W. Dale, R. F. Gill, R. J. Gill, C. Hirschener, M. A. Isaak, S. Johnson, S. E. Jungers, T. Kipping,
A. L. LeMon, K. D. Levy, J. F. Parinas, J. Pretare, W. E. Rauscher, D. C. Russell, and R. Takumi.
FIVE HUNDRED EIGHTH MEETING
The 508th meeting of the Pacific Coast Entomological Society was held on 18 March 1994, at 8:00
PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Kirby W. Brown presiding.
The minutes of the 18 February 1994 meeting were read and accepted. Mr. Ricardo Ayala was
proposed and elected as a new regular member.
Mr. Steven E. Jungers exhibited an introduced sow bug and its introduced spider predator.
Dr. Brown announced that the Robert T. Wallace undergraduate research internship is being offered
by the California Academy of Sciences. Applications are being accepted until 1 April 1994.
Dr. Norman D. Penny announced that the Department of Entomology at the California Academy
of Sciences will not accept donations of specimens unless any necessary permits and accompanying
documentation are included. He also announced that the Academy will have a new exhibit this coming
summer called Monarca highlighted with live monarch butterflies.
Dr. Brown announced that the Nature Conservancy is conducting a biologicalinventory of California.
They would like help from specialists on species lists and determinations.
The featured speaker Mr. James E. Tobler I of Santa Rosa, California, presented a slide lecture
entitled ““Adventures of an Amateur Entomologist.” Mr. Tobler talked about his interest in entomology
as early as the 5th grade and reminisced about being hired by Dr. Edward S. Ross to pin insects for
the entomology department of the California Academy of Sciences during a summer vacation for
$1.50 per hour. After high school Mr. Tobler served in the United States Navy aboard an aircraft
carrier and visited 29 ports in the eastern Mediterranean and western Pacific, collecting insects when-
ever he could. When his enlistment was up, Mr. Tobler took a year-long collecting trip to the South
Pacific and shipped specimens home, later to be prepared and donated to the Academy.
The meeting adjourned at 10:00 PM, followed by a social hour in the entomology conference room .—
K. Ribardo, Recording Secretary.
The following 58 persons were present. 41 members: R. L. Aalbu, P. H. Arnaud Jr., C. M. Brandau,
116 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
T. S. Briggs, K. W. Brown, R. M. Brown, B. Buickerood, P. Buickerood, H. K. Court, D. K. Dabney,
J. A. DeBenedictis, W. A. Doolin, E. M. Fisher, N. E. Gershenz, C. E. Griswold, M. A. Isaak, S. E.
Jungers, R. L. Langston, G. J. Mallick, J. D. McCarty, J. J. McNicol, T. C. Meikle, S. T. O’Keefe, A.
M. L. Penny, N. D. Penny, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, E. S. Ross,
L. S. Saul, W. E. Savary, J. Schweikert, H. I. Scudder, C. Y. Takahashi, J. E. Tobler I, D. Ubick, S.
E. Vaughn, R. Wilson, R. J. Worth, and R. L. Zuparko; (17 guests) M. M. Amaud, V. Avery, T.
Aweeka, A. Baier, J. Baier, C. B. Barr, S. Brown, J. E. Court, K. Doi, S. E. Haskins, S. Johnson, D.
J. Leprince, J. F. Parinas, A. Prather, W. E. Rauscher, and 2 persons with illegible signatures.
FIVE HUNDRED NINTH MEETING
The 509th meeting of the Pacific Coast Entomological Society was held on 15 April 1994, at 8:00
PM, in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Dr. Kirby W. Brown presiding.
The minutes of the 18 March 1994 meeting were read and accepted. Five persons were proposed
and elected as new members: Mr. Raymond J. Gill as a regular member; and Mr. Michael R. Carbiener,
Ms. Emily J. Friess, Ms. Leslie Goldberg, and Mr. Jeffrey Honda as student members.
Mr. Darrell Ubick exhibited an adult and pupa of the acrocerid genus Turbopsebius, a family of
flies which parasitizes spiders.
Mr. Warren E. Savary presented a note on Saltonia incerta (Banks), a previously very rare spider
known from only a few specimens. Mr. Savary was able to collect approximately one hundred specimens
using pitfall traps placed in the salt flats of Soda Lake, a dry lake bed, near Baker, California.
Mr. Vincent F. Lee announced that the first issue of the Pan Pacific Entomologist for 1994 will be
mailed in a few weeks.
Dr. Brown announced that the newsletter of the Pacific Division of the American Association for
the Advancement of Science was sent out for distribution. He announced that the 8th annual Insect
Fair will be held 14 and 15 May at the Los Angeles County Arboretum.
The featured speakers Dr. James Carey, University of California, Davis, and Dr. Robert V. Dowell,
California Department of Food and Agriculture, presented a slide-illustrated discussion entitled ““The
Medfly in California: Two Perspectives.’ Dr. Dowell proposed that the medfly is not a permanent
member of the California fauna. It can be eradicated, and reinfestation occurs when infested host
material enters the state. Dr. Carey proposed that the medfly is established (at least since 1987), and
it can’t be eradicated. He believes unaffected areas should be kept free from invasion by containing
the flies where they are now.
The meeting adjourned at 9:50 PM, followed by a social hour in the entomology conference room .—
K. Ribardo, Recording Secretary.
The following 51 persons were present. 32 members: P. H. Amaud Jr., L. G. Bezark, F. L. Blanc,
T. S. Briggs, K. W. Brown, R. M. Brown, D. K. Dabney, R. V. Dowell, E. M. Fisher, C. D. Franklin
II, C. E. Griswold, W. Hamersky, S. Hammond, R. L. Langston, V. F. Lee, J. D. McCarty, T. C.
Meikle, S. B. Opp, A. M. L. Penny, N. D. Penny, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, W. E.
Savary, J. Schweikert, L. A. Solorzano, J. T. Sorensen, R. E. Stecker, C. Y. Takahashi, A. Telang, D.
Ubick, and S. E. Vaughn; (19 guests) F. Blanc, I. Brown, M. R. Carbiener, J. R. Carey, G. Flohr, M.
Gilkey, S. E. Haskins, S. L. Holt, A. L. LeMon, D. J. Leprince, S. S. Mead, J. F. Parinas, D. A.
Piechnik, W. E. Rauscher, D. C. Russell, M. J. Sawyer, K. Sorensen, K. Sorensen, and D. Vorous.
FIVE HUNDRED TENTH MEETING
The 510th meeting of the Pacific Coast Entomological Society was held on 20 May 1994, at 8:00
PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President Dr. Kirby W. Brown presiding.
The minutes of the 15 April 1994 meeting were read and accepted. Three persons were proposed
and elected as new members: Mr. Mentor F. Huebner and Dr. Stephen W. Taber as regular members;
and Ms. Gretchen Flohr as a student member.
Dr. Brown announced that the annual butterfly count will be held in mid- to late June. Areas to be
surveyed include Marin, Sonoma, Mt. Diablo, South San Francisco Bay Lands, and San Joaquin.
Mr. Vincent F. Lee announced that copies of the newsletter of the Pacific Division of the American
Association for the Advancement of Science are available in the entomology conference room.
Dr. Brown announced that the 8th annual Insect Fair was held 14 and 15 May at the Los Angeles
1996 PROCEEDINGS 117
County Arboretum. He observed that there were a lot of insects for sale, good attendance by the
public, and of special interest, an atlas of insects on stamps of the world for sale.
Mr. Keve J. Ribardo announced that the Entomology Club of San Jose State University held an
overnight collecting trip on 14-15 May at the Arroyo Seco campground in the Los Padres National
Forest. A good time was had by all, and everyone is welcome to participate in future trips.
Mr. Paul G. da Silva announced that there was a job opening for someone interested in insect pests
of fruit and nut crops in the San Joaquin Valley. Flyers would be available after the meeting.
Dr. Norman D. Penny presented a note on Nothochrysinae, a subfamily of the Chrysopidae (Neu-
roptera). California seems to be a center of endemism with two genera and six species represented in
the state. Slides of Pimachrysa nigra Adams and Nothochrysa californica Banks were shown.
The featured speaker Dr. Edward S. Ross of the California Academy of Sciences presented a slide
lecture entitled ““More Surprises Along Amazon Forest Trails,” with an update on the research facility
on the Upper Rio Napo in Ecuador. Dr. Ross showed examples of mimicry complexes of tropical
insects with many slides of both models and their mimics. Facilities and trails of Cabafias Alifahui
were featured, showing the high species diversity and endemism of this biotically rich zone.
The meeting adjourned at 9:55 PM, followed by a social hour in the entomology conference room.—
K. Ribardo, Recording Secretary.
The following 91 persons were present. 52 members: R. L. Aalbu, E. Anderson, P. H. Arnaud Jr.,
T. S. Briggs, K. W. Brown, R. M. Brown, P. Buickerood, R. Buickerood, M. R. Carbiener, J. R.
Clopton, P. G. da Silva, D. K. Dabney, B. Deutsch, J. T. Doyen, B. K. Eya, S. S. Ferguson, W. E.
Ferguson, E. M. Fisher, E. J. Friess, M. Garcia-Vidal, C. E. Griswold, W. Hamersky, A. Hom, M. A.
Isaak, B. Keh, R. L. Langston, V. F. Lee, J. J. McNicol, T. C. Meikle, W. D. Murray, A. M. L. Penny,
N. D. Penny, A. E. Rackett, K. A. Reynolds, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, E. S.
Ross, R. W. Rust, W. E. Savary, J. Schweikert, H. I. Scudder, L. A. Solorzano, R. E. Somerby, R. E.
Stecker, C. Y. Takahashi, A. Telang, D. Ubick, S. E. Vaughn, S. P. Welles Jr., T. J. Zavortink, and 1
illegible signature; (39 guests) M. M. Arnaud, T. Aweeka, R. Bandar, B. Bell, D. Bell, I. Biagi, C.
Cripps, H. V. Davis, S. Haugue, K. S. Hom, G. C. Howard, A. Jesse, D. Judd, A. Jung, P. Knykl, D.
J. Leprince, G. Y. Leung, W. A. Maffei, S. S. Mead, J. Mendeleyev, L. Merrell, F. Murray, J. F.
Parinas, J. Robertson, S. M. Ross, J. C. Rust, J. Shackelfold, K. Simms, T. E. Simms, V. Simms, D.
Truax, M. Truax, and 7 illegible signatures.
FIVE HUNDRED ELEVENTH MEETING
The 511th meeting of the Pacific Coast Entomological Society was held on 16 September 1994, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President-elect Mr. Curtis Y. Takahashi presiding.
The minutes of the 20 May 1994 meeting were read and accepted. Eight persons were proposed
and elected as new members: Mr. Robert W. Duncan, Mr. Richard E. Hill, and Mr. Daniel J. Leprince
as regular members; Mr. Luis E. Rivera-Cervantes and Ms. Edith Garcia-Real as family members;
and Mr. Matthew D. Dean, Mr. Juraj Halaj, and Mr. Michael J. Stout as student members.
Mr. Takahashi announced that Dr. Kirby W. Brown had retired after 20 years as economic ento-
mologist for San Joaquin County and was taking a part-time teaching position at Northland Pioneer
College in Holbrook, Arizona. Dr. Brown will not be able to attend the last 4 meetings of 1994, but
plans to present a lecture at the January 1995 meeting.
Mr. Jere Schweikert presented a note on Terellia fuscicornis (Loew), a tephritid fly that infests seed
heads of artichokes, artichoke thistle, and cardoon. It has been found in most counties in the Bay
Area and Central Valley. This is a new record for this fly in North America. Slides of the adults, host
plants, and damage were shown.
The featured speaker Dr. Charles E. Griswold of the California Academy of Sciences presented a
slide lecture entitled ““Rapid Assessment of Arthropod Species Richness in the Tropics.’ Dr. Griswold
described sampling protocols used to estimate biodiversity in tropical ecosystems. Three sites in
Bolivia, two in Cameroon, and two in Madagascar were sampled. Four main factors effecting sampling
were compared: site, collector, time of day, and method. Little overlap in species of arthropods between
sites was found. Dr. Griswold noted three types of information in conservation biology that systematic
biologists should be able to provide based on sampling: total species richness, degree of endemism,
and phylogenetic importance of species found. He believes that museum collecting methods can be
modified to provide statistically analyzable samples while still maximizing the collecting effort for
traditional use of specimens.
118 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
The meeting adjourned at 9:35 PM, followed by a social hour in the entomology conference room. —
K. Ribardo, Recording Secretary.
The following 51 persons were present. 33 members: P. H. Amaud Jr., J. E. Brandriff, T. S. Briggs,
J. S. Chinn, H. K. Court, P. G. da Silva, D. K. Dabney, J. G. Edwards, S. V. Fend, C. E. Griswold,
M.A. Isaak, D. H. Kavanaugh, R. L. Langston, V. F. Lee, J. J. McNicol, T. C. Meikle, L. A. Norton,
S. T. O’Keefe, A. M. L. Penny, N. D. Penny, J. A. Powell, W. J. Pulawski, A. E. Rackett, J. M.
Ribardo, K. J. Ribardo, R. G. Robertson, E. S. Ross, D. C. Russell, J. Schweikert, R. E. Stecker, C.
Y. Takahashi, S. E. Vaughn, and S. C. Williams; (18 guests) V. Ahrens-Pulawski, M. M. Amaud, T.
Aweeka, G. E. Ball, L. H. Carroll, J. E. Court, A. Elinor, D. F. Field, J. B. Fraser, M. H. Fraser, B.
R. Manchester, R. Morgan, J. F. Parinas, W. E. Rauscher, S. Renkes, A. Sebastian, L. Sebastian, and
J. Urzykowski.
FIVE HUNDRED TWELFTH MEETING
The 512th meeting of the Pacific Coast Entomological Society was held on 21 October 1994, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President-elect Mr. Curtis Y. Takahashi presiding.
The minutes of the 16 September 1994 meeting were read and accepted. Seven persons were proposed
and elected as new members: Mr. Lawrence W. Currie Jr., Dr. Manfredo Fritz, Dr. William T. Starmer,
Dr. Patrick M. Sugg, and Dr. Charles E. Turner as regular members; Ms. Johanna E. Brandriff and
Mr. Blane R. Manchester as student members.
Mr. Vincent F. Lee announced that copies of the Entomology Collection Network registration form
would be available in the entomology conference room after the meeting. Mr. Lee reminded members
that dues envelopes were included with the third issue of the Society’s journal, which was mailed a
couple of days ago, and issue number four would be out by the end of the month.
Dr. Norman D. Penny presented a note on an ant lion he collected in the Ciervo Hills of Fresno
County. This specimen is the first new species of ant lion to be discovered since the family was revised
in 1970. Dr. Penny also displayed a neuropteran with a four-inch wingspan from Honduras, which
appears to be intermediate between ithonids and rapismatids.
Mr. Larry G. Bezark presented a note on Harmonia axyridis (Pallas), a coccinellid from Asia,
released for suppression of pecan aphids. He collected two adult specimens on fennel in May, near
downtown Sacramento. This represents the first recovery of this species in California.
The featured speaker Mr. Robert L. Allen of the Natural History Museum of Los Angeles County
presented a slide lecture entitled ““Arthropods of Special Concern in California.” Mr. Allen discussed
the decline of diversity in California due to habitat destruction. Mr. Allen gave some scenarios of
how very localized populations could be wiped out when fields are disked or when sheep or goats are
put on a piece of land to mow down plants before Environmental Impact Reports are completed. He
explained that arthropods are poorly surveyed, and information is not widely available so they are
not usually included in habitat management or mitigation. There is a lack of available specialists, and
with no information and no requirements to gather any, no surveys are done.
The meeting adjourned at 9:20 PM, followed by a social hour in the entomology conference room.—
K. Ribardo, Recording Secretary.
The following 37 persons were present. 30 members: R. L. Allen, L. G. Bezark, T. S. Briggs, R. M.
Brown, D. C. Carlson, M. S. Caterino, H. K. Court, L. W. Currie Jr., B. Deutsch, W. Hamersky, A.
Hom, V. F. Lee, D. J. Leprince, B. R. Manchester, J. J. McNicol, L. A. Norton, S. T. O’Keefe, N. D.
Penny, A. E. Rackett, K. A. Reynolds, J. M. Ribardo, K. J. Ribardo, D. C. Russell, W. E. Savary, J.
Schweikert, H. I. Scudder, C. Y. Takahashi, D. Ubick, S. E. Vaughn, and S. A. Wright; (7 guests) C.
B. Barr, J. E. Court, D. D. Giuliani, W. E. Rauscher, H. Stigler, R. Takumi, and G. Yoshimura.
FIVE HUNDRED THIRTEENTH MEETING
The 513th meeting of the Pacific Coast Entomological Society was held on 18 November 1994, at
8:00 PM, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President-elect Mr. Curtis Y. Takahashi presiding.
The minutes of the 21 October 1994 meeting were read and accepted. Two persons were proposed
and elected as new regular members: Dr. William J. Roltsch and Dr. Felix A. H. Sperling.
Mr. Curtis Y. Takahashi announced that this year’s auditing committee would consist of Mr. H.
Vannoy Davis, CPA, Mrs. Helen K. Court, and Dr. Norman D. Penny; and that the nominating
1996 PROCEEDINGS 119
committee members are Mr. Larry G. Bezark, Dr. John E. Hafernik Jr., Dr. Ronald E. Stecker, and
chaired by Dr. Susan B. Opp. Mr. Takahashi announced that the 29th Annual Conference of the
Association of Applied Insect Ecologists will be held on 5—7 February 1995 in Santa Barbara, California.
He also noted that a member of the Society, Mr. Robert L. Langston, was presented the John A.
Comstock Award by the Lepidopterists’ Society on 22 October 1994.
Dr. Charles E. Griswold announced that Mrs. Stella E. Tatro, a curatorial assistant of the Department
of Entomology at the California Academy of Sciences, passed away on 12 November, and a memorial
service would be held in the Trustees Room at the Academy on Monday, 5 December at 4:00 PM.
All were welcome to attend.
Dr. J. Gordon Edwards announced that Dr. William E. Hazeltine passed away on 5 November
1994, at the age of 68. He suffered a fatal heart attack while collecting scarab beetles in the Santa Cruz
Mountains of California.
Ms. Cheryl B. Barr announced that the U. C. Berkeley insect museum was looking for a part-time
insect preparator. Interested persons were encouraged to apply.
Mr. Stanley E. Vaughn presented a note on collecting the “‘rain beetle” P/eocoma at Calero Reservoir
in Santa Clara County, California.
The featured speaker Dr. David B. Weissman presented a slide lecture entitled ““Biotaxonomy of
the Jerusalem Crickets: An Uncircumscribed Group.” Dr. Weissman discussed the status of Jerusalem
cricket genera in North America, all of which will be included in the genus Steno pelmatus. Their size
ranges from 8 to 70 mm, and they go through ten molts in the two years to reach maturity. The
crickets communicate with each other and attract mates by drumming. The cadence of the drumming
is species specific.
The meeting adjourned at 9:30 PM, followed by a social hour in the entomology conference room.—
K. Ribardo, Recording Secretary.
The following 43 persons were present. 34 members: R. L. Aalbu, P. H. Arnaud Jr., L. G. Bezark,
T. S. Briggs, M. R. Carbiener, H. K. Court, L. W. Currie Jr., B. Deutsch, W. A. Doolin, J. G. Edwards,
C. E. Griswold, A. Hom, S. E. Jungers, V. F. Lee, T. C. Meikle, S. T. O’Keefe, A. M. L. Penny, N.
D. Penny, A. E. Rackett, K. A. Reynolds, J. M. Ribardo, K. J. Ribardo, D. C. Russell, W. E. Savary,
J. S. Schweikert, H. I. Scudder, F. Sperling, R. E. Stecker, C. Y. Takahashi, D. Ubick, S. E. Vaughn,
D. B. Weissman, R. R. White, and R. L. Zuparko; (9 guests) M. M. Arnaud, C. B. Barr, J. E. Court,
B. Deutsch, J. Floh, L. Garibaldi, R. Morgan, W. E. Rauscher, and D. W. Weissman.
FIVE HUNDRED FOURTEENTH MEETING
The 514th meeting of the Pacific Coast Entomological Society was held on 9 December 1994, at
8:00 PM, in the Redwood Room of the California Academy of Sciences, Golden Gate Park, San
Francisco, with President-elect Mr. Curtis Y. Takahashi presiding.
The minutes of the 18 November 1994 meeting were read and accepted. Ms. Julieta F. Parinas was
proposed and elected as a new regular member.
Mr. Vincent F. Lee announced that Mr. George J. Mallick passed away on the 28 November 1994.
Mr. Mallick was a member of the Society for 20 years and a strong supporter of the California Academy
of Sciences. Mr. Lee reported that the Pacific Coast Entomological Society exchanged the Pan-Pacific
Entomologist for 71 publications from 53 organizations.
Dr. Norman D. Penny announced that an old alcohol case was available for the taking and that
duplicate reprints would be given away to those who wished them.
Dr. Paul H. Arnaud Jr. presented a slide-illustrated note on Mrs. Stella E. Tatro, curatorial assistant
of the Department of Entomology at the California Academy of Sciences for 30 years, who passed
away on 12 November 1994.
Mr. Paul G. da Silva presented a note on the abundance of the “‘rain beetle”? Pleocoma tularensis
Leach this year.
The featured speaker Mr. Patrick A. Luft presented a slide lecture entitled ‘““Adaptive Biology and
Behavior of Trioza eugeniae (Froggatt) (Homoptera: Triozidae) in Response to Biotic and Abiotic
Factors.”” Mr. Luft discussed the biology of psyllids and described a detailed study of T. eugeniae
including spatial distribution patterns.
The meeting adjourned at 9:20 PM, followed by a social hour in the entomology conference room. —
K. Ribardo, Recording Secretary.
The following 32 persons were present. 23 members: P. H. Arnaud Jr., L. G. Bezark, H. K. Court,
T. D. Cuneo, D. K. Dabney, P. G. daSilva, W. Hamersky, R. L. Langston, V. F. Lee, D. J. Leprince,
120 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(2)
A. M. L. Penny, N. D. Penny, A. E. Rackett, J. M. Ribardo, K. J. Ribardo, R. G. Robertson, W. E.
Savary, J. S. Schweikert, C. Y. Takahashi, S. E. Vaughn, R. R. White, S. C. Williams, and R. L.
Zuparko; (9 guests) M. M. Arnaud, M. Benoit, J. E. Court, E. Goff, P. A. Luft, W. A. Maffei, J. F.
Parinas, J. Pretare, and J. Robertson.
PAN-PACIFIC ENTOMOLOGIST
Information for Contributors
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discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English,
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volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are
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(INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author
of each species after its first mention.
Data Formats. — All specimen data must be cited in the journal’s locality data format. See volume 69(2), pages 196-198 for these
format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before
submitting manuscripts for which these formats are applicable.
Literature Cited. — Format examples are:
Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.
Blackman, R. L., PR. A. Brown & V. F Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide
some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. EK Dixon & L. Weismann (eds.). Population structure, genetics and
taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB
Academic Publishing, The Hague, The Netherlands.
Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.
Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.
Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label
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Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for
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THE PAN-PACIFIC ENTOMOLOGIST
Volume 72 April 1996 Number 2
Contents
NAJT, J. & W. M. WEINER—Geographical distribution of Brachystomellinae (Collembola:
ISSR MORON CHG Pav) eye hE ek i be Ae ge SEs Re ee 61
SIKES, D. S.—The natural history of Nicrophorus nigrita, a western Nearctic species (Cole-
Sj emg rg OTE 2s Mae BZ Ea ee ee) ee 70
RIDDICK, E. W. & N. J. MILLS—A comparison of the seasonal activity of Pterostichus beetles
(Coleoptera: Carabidae) in a commercial apple orchard in Sonoma County, California __ 82
ROGERS, D. C.—Eubranchipus bundyi Forbes (Anostraca: Crustacea), a new record from
(GRINGO Ig ere ee ee Ee a ee ee ee 89
CAMBRA, R. A. & D. QUINTERO A.—The Mexican and Central American species of Lo-
phostigma Mickel, including a new species, new distribution records, and taxonomic
netes forthe genus (hiymenoptera, Mmtllitae) 9 os 92
SCIENTIFIC NOTE
SORENSEN, J. T-—Recent California records for the sawfly Xiphydria mellipes Harris (Hy-
TMeOOpieka: Sulpyvanitciie |). Ln | Gama ee eee Oe Ree ee ee 102
Paciiie Coast Bitomo@logicalisocirety Proceedines or moo 2? «2 4 Se 104
Pacific: Coast Entomological Seciety, Proceedings for 1993. spk 109
Pacific Coast Entomological Socrety,-Proceedings for 1994.4. 2 114
The
PAN-PACIFIC
ENTOMOLOGIST
Volume 72 July 1996 Number 3
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
(ISSN 0031-0603)
The Pan-Pacific Entomologist
EDITORIAL BOARD
R. V. Dowell, Editor R. M. Bohart
R. L. Penrose, Associate Editor J. T. Doyen
R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.
Julieta E Parinas, Treasurer Warren E. Savary
Published quarterly in January, April, July, and October with Society Proceed-
ings usually appearing in the October issue. All communications regarding non-
receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary;
and financial communications should be addressed to: Julieta F Parinas, Treasurer;
at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad-
emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
Application for membership in the Society and changes of address should be
addressed to: William Hamersky, Membership Committee chair, Pacific Coast
Entomological Society, Dept. of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, CA 94118-4599.
Manuscripts, proofs, and all correspondence concerning editorial matters (but
not aspects of publication charges or costs) should be sent to: Dr. Robert V.
Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture,
1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con-
tributors, and volume 66(1): 1—8, January 1990, for more detailed information.
Information on format for taxonomic manuscripts can be found in volume 69(2):
194-198. Refer inquiries for publication charges and costs to the Treasurer.
The annual dues, paid in advance, are $25.00 for regular members of the So-
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The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes
are available; see 67(1): 80 for current prices. Make checks payable to the Pacific
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Pacific Coast Entomological Society
OFFICERS FOR 1996
Wojciech J. Pulawski, President Vincent EK Lee, Managing Secretary
Julieta E Parinas, Treasurer Stanley E. Vaughn, Recording Secretary
THE PAN-PACIFIC ENTOMOLOGIST (ISSN: 0031-0603) is published quarterly for $40.00 per
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PAN-PACIFIC ENTOMOLOGIST
72(3): 121-129, (1996)
POSSIBLE ROLE OF NONFERTILIZING SPERM AS A
NUTRIENT SOURCE FOR FEMALE DROSOPHILA
PSEUDOOBSCURA FROLOVA (DIPTERA:
DROSOPHILIDAE)
RHONDA R. SNOOK! AND THERESE A. MARKOW
Department of Zoology, Arizona State University, Tempe, Arizona 85287-1501
Abstract.—Males in some insect taxa produce chromosomally or morphologically variant sperm
types, one of which does not fertilize eggs. Hypotheses as to the functional significance of
nonfertilizing sperm in these taxa state that nonfertilizing sperm may represent nutrient resources
utilized by fertilizing sperm, the female mate or the zygote. Drosophila pseudoobscura Frolova
males produce two discrete lengths of sperm, short and long, but short sperm do not participate
in fertilization. Additionally, females of this species incorporate '*C male-derived materials into
their tissues. We tested whether these male-derived substances incorporated by D. pseudoobscura
females originate from nonfertilizing short sperm and could represent a nutrient donation by
males. We tracked the fate of '*C material from the male within female tissues and found that
females incorporated radiolabel into somatic tissues by six hours after copulation. However, short
sperm do not begin to disappear from sperm storage organs until 6 hours after copulation and
we found no association between the subsequent loss of short sperm in storage and the amount
of male-derived material consequently incorporated into female somatic tissues or oocytes. These
results suggest that short nonfertilizing sperm are not the source of '*C male-derived components
incorporated by females and we conclude that short sperm do not serve as nutrient donations to
the female mate or the zygote.
Key Words.—Insecta, nonfertilizing sperm, male-derived nutrient donations, Drosophila pseu-
doobscura, sperm polymorphism, ejaculate
Males in certain insect taxa transfer products at mating that 1) become incor-
porated into female somatic tissues and ovarian oocytes (e.g., Boggs & Gilbert
1979, Thornhill & Alcock 1983, Markow & Ankney 1988) and, 2) are associated
with increases in fitness components of the female mate or zygote (Gwynne 1981,
1988; Butlin et al. 1987; Rutowski et al. 1987). In those Drosophila species in
which females incorporate ejaculatory materials, it has been assumed that the
incorporated material is derived from male accessory gland products, but this
assumption has not been tested directly (Bownes & Partridge 1987, Markow &
Ankney 1988, Pitnick et al. 1991). Sperm, not accessory gland secretions, could
potentially function as male-derived nutrient resources. In species in which males
simultaneously produce two sperm types, one of which functions in fertilization
and one that cannot fertilize eggs, nonfertilizing sperm could be nutritive. Such
nutrients may benefit the fertilizing sperm types, the female mate or the zygote
(Hanson et al. 1952, Sivinski 1980, Healy & Jamieson 1981, Silberglied et al.
1984).
Males of species in the Drosophila obscura Fallén group produce two distinct
lengths of nucleated sperm, short and long, within each ejaculate (Beatty & Sidhu
1970). We have found that, in D. pseudoobscura Frolova, only long sperm morphs
' Current address: Department of Ecology and Evolutionary Biology, University of Tennessee,
Knoxville, Tennessee 37996-1610. Author’s page charges partially offset by a grant from the C.P.
Alexander Fund, PCES.
122 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
fertilize eggs despite each sperm type being transferred in equal proportions to
females (Snook et al. 1994). Short sperm begin to move into sperm storage organs
(SSOs) approximately two hours after copulation, but between 6 and 12 h post-
mating the number and proportion of short sperm begins to decrease in SSOs
while the number and proportion of long sperm continues to increase during these
times. These changes in the numbers of each sperm type found in SSOs do not
alter the proportion of short or long sperm found in the uterus until oviposition
begins. The mechanism of “disappearance” of short nonfertilizing sperm from
female SSOs is unknown but perhaps these sperm could be broken down and
serve as the source of male-derived '*C material incorporated into female tissue.
It is unlikely that short sperm within the uterus serve this function since the
proportion of short sperm found there does not change until oviposition begins
24 to 48 h after mating (Snook et al. 1994).
To test if male-derived material in female D. pseudoobscura is from nonfertil-
izing sperm that could then serve as a beneficial nutrient donation, we examined
the timing of incorporation of male-derived substances into female tissues. We
predicted that if short, nonfertilizing sperm function as nutrient resources, the
temporal decrease of short sperm within female SSOs beginning between 6 and
12 h after mating (Snook et al. 1994) should correspond to increasing amounts
of male-derived substances detectable in female tissues.
MATERIALS AND METHODS
Fly cultures.—Drosophila pseudoobscura was established from flies collected
on fallen citrus in Tempe, AZ in 1990 and 1991. Flies were maintained on stan-
dard cornmeal-agar-molasses food with yeast and kept at room temperature, 22—
25° C, and an approximate 12:12 L:D photoperiodic cycle. All flies used in
experiments were 5 days old and reproductively mature.
Tracking ejaculate contributions.—If short sperm function as a nutrient source,
their disappearance should precede the detection of male-derived materials in
female tissue. To track the timing of male-derived material in females, we mated
females to radiolabelled males and assayed radioactivity subsequently found in
female tissues at 6, 12, and 24 h after copulation, corresponding to prior times
examining sperm movement in SSOs (Snook et al. 1994). Fifty, first instar larvae
from culture bottles were placed into vials containing 5 g of culture media in-
oculated with 50uCi of a '*C labelled mixture of amino acids. Virgin males were
collected from this larval culture on eclosion and stored 10 per food vial with
dry yeast until they were mated to nonradiolabelled, virgin females. These females
were nutritionally stressed because previous work noted that the greatest effect
of male substances on females occurred when females were food-limited (Bownes
& Partridge 1987, Butlin et al. 1987, Markow et al. 1990, Fox 1993; but see
Chapman et al. 1994). Females were collected upon eclosion, stored in food vials
(10/vial), and nutritionally stressed by providing no surface yeast in the food vials.
Two virgin radiolabelled males were placed with one nutritionally stressed vir-
gin female in an unyeasted food vial and upon mating, the “‘unsuccessful’’ male
was aspirated from the vial without disturbing the copulating pair. Immediately
after copulation ended the successful male was removed from the vial. Nine fe-
males were examined at each time interval (6, 12 and 24 h) after mating. They
1996 SNOOK & MARKOW: ROLE OF NONFERTILIZING SPERM 123
were ether-anesthetized and processed for scintillation counting to determine the
amount of radioactivity present.
To process mated females for scintillation, flies were washed once for 30 sec
in a microcentrifuge tube containing a mixture of phosphate buffer solution (PBS)
and Triton-X to remove any external radiolabel transferred by the male during
copulation, followed by another wash in PBS after which they were decapitated
(eye pteridines contribute to a quenching effect). Female reproductive tracts were
removed intact and the ovaries detached from the common oviduct. The remainder
of the reproductive tract from each female was examined for the presence of
Sperm, and ovaries and the remaining somatic tissue were washed separately as
described above for whole flies. Ovarian and somatic tissue samples from three
females were pooled in a scintillation vial containing tissue solubilizer. Tissues
were then crushed with the end of a clean glass rod and digested for 24 h in a
50° C waterbath. At room temperature, the tissues were first neutralized with
glacial acetic acid, followed by the addition of scintillation fluid. Subsequently,
tissues were vortexed, and 24 h later samples were counted by a Beckman LS
7000 liquid scintillation system. Counts per minute were converted to disintegra-
tions per minute following a standard quench curve (Pitnick et al. 1991). Nutri-
tionally stressed females mated to nonradiolabelled males (n = 9) served as a
control and were processed as above. Scintillation vials containing no tissue (n
= 6) were used to determine background counts.
Statistical analyses.—To test the prediction that the disappearance of short
sperm within SSOs corresponds to an increase in radiolabel detected in other
tissues of females, we compared data originally reported in Snook et al. (1994),
describing the number of short sperm found in SSOs at 6, 12, and 24 h after
mating, with the amount of radiolabel found in females at these times. We gen-
erated the mean + standard error (SE) of short sperm found in SSOs at 6, 12,
and 24 h after mating, by pooling data from the ventral receptacle and sperma-
thecae of nonovipositing females. One datum collected at 24 h after mating was
an extreme outlier and was removed from analyses; the number of short sperm
found in the ventral receptacle of this sample was approximately 1000 while the
mean + SE of the remaining seven samples was 91.7 + 36.6.
Radiolabel data and sperm count data were acquired during different experi-
ments. However, reported sperm counts for each time examined in Snook et al.
(1994) were replicated 3 times over a 2 month period with the entire dataset being
collected over 10 months. Despite the temporal disparity in when replications
were collected, there were no significant differences in the numbers of short sperm
found among replicates in each sperm storage organ at 12 and 24 h after mating
(replicates were tested using Kruskal-Wallis ANOVA because values were not
normally distributed (Sokal & Rohlf 1981); 12 h (total n over all replicates =
13), ventral receptacle (VR), P = 0.34, spermathecae (SP), P = 0.38; 24 h (total
n over all replicates = 7) VR, P = 0.13, SP, P = 0.24)). At 6 h after mating,
however, there were significant differences among replicates in the number of
short sperm found in the VR (P = 0.01), but not in the SP (total n over all
replicates = 14, P = 0.183). The difference in the number of short sperm found
in replicates of the VR seemed to be due to one replicate in which there were
smaller numbers of both short and long sperm found. Subsequently, we tested if
the proportion of short sperm differed between replicates; there was no difference
124 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
(n = 14; P = 0.12; Snook et al. 1994). Therefore, the pattern of sperm storage
and the amount, either the number or proportion of short sperm, found in SSOs
are highly constant between females and throughout time, justifying the use of
these earlier storage data in testing our prediction. All statistics were performed
using Systat (version 5.03, Systat, Inc; Wilkinson 1990).
RESULTS
The proportion of short sperm in the uterus does not change from the time of
transfer until more than 24 h after copulation (F = 1.339; df = 4, 56; P = 0.267;
Fig. 1). However, comparing the number of short sperm found in SSOs between
6 to 24 h postmating indicated that short sperm “disappeared”? within SSOs (F
= 4.933; df = 2, 31; P = 0.01; Fig. 1). Tukey’s a posteriori tests indicated that
significant disappearance of short sperm occurred between 6 and 12 h (P = 0.037)
and between 6 and 24 h (P = 0.042) but not between 12 and 24 h (P > 0.05).
The proportion of short sperm found in SSOs also decreased over time (F =
6.355, df = 2, 31; P = 0.005). However, only the comparison between 6 and 24
h was significant (P = 0.005). The number of sperm found in SSOs 2 h after
mating is reported in Fig. 1 to indicate that the maximum number of short sperm
is stored at 6 h postcopulation. Comparing the number of long sperm found in
SSOs between 6 to 24 h postmating indicated that long sperm do not continue to
significantly increase (F = 2.933, df = 2, 31, P = 0.07; Fig. 1).
If short sperm serve as a nutrient contribution, then the increase in incorporation
of radiolabel in female tissues is predicted to occur subsequent to 6 h postcopu-
lation, when short sperm are maximally stored. We found that significant amounts
of radiolabel were incorporated in the somatic tissue (F = 9.284; df = 3, 8; P =
0.006; Fig. 2), but not the oocytes (F = 3.115; df = 3, 8; P = 0.09), of starved
D. pseudoobscura females. Contrary to our prediction, however, Tukey’s a pos-
teriori tests indicated that significant radiolabel incorporation occurred between O
h and 6h (P = 0.035) after mating, but there was no significant increase after 6
h (Fig. 2).
DISCUSSION
The evidence presented here indicates that short sperm do not function as a
nutrient resource for D. pseudoobscura females. If short nonfertilizing sperm in
D. pseudoobscura break down and function as a nutrient source, the surge in the
amount of radiolabel detected in female tissues should have occurred when the
short sperm begin to disappear from the SSOs, between 6 h and 12 h after cop-
ulation (Fig. 1; Snook et al. 1994). The increase in radiolabel in somatic tissues
that we observed was prior to 6 h, when the number of short sperm was the
largest within SSOs. Short sperm in the uterus are also unlikely to serve as nu-
trient donations because the proportion of short sperm does not change from the
—»
Figure 1. Mean (+SE) of the proportion of short sperm present in the uterus and the total number
of short and long sperm in the sperm storage organs (SSOs-ventral receptacle and spermathecae) of
D. pseudoobscura at various times after mating. Values at 0 h indicate sperm in the uterus immediately
after copulation and thus, represent male sperm transfer. Females do not have sperm in SSOs at 0 h
1996 SNOOK & MARKOW: ROLE OF NONFERTILIZING SPERM 125
= 1.0
i Uterus
08
LL
O 06
6
= 0.4
©
oO 02
r 0.0
400 7 sso-Short sperm b
300
Ss 200
ag
Lu
o 100
LL
O
n'a @)
im
= 2000 2
> 4800 SSO-Long sperm
x
i
O
bh
0 2 6 12 24
HOURS AFTER MATING
postmating. Sample sizes are 11 for 0 h, 15 for 2 h uterus, 13 for 2 h SSOs, 15 for 6 h, 13 for 12 h,
7 for 24 h. Values for 24 h are for females that have not oviposited. Letters a—d denote significantly
different means detected by Tukey’s a posteriori multiple comparison tests; comparisons for the uterus
(letter a), short sperm in the sperm storage organs (SSO-Short sperm) (letters b and c), and long sperm
in the sperm storage organs (SSO-Long sperm) (letter d) were tested separately.
126 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
[__ | Background counts
200 W////A Control nonradiolabelled females
MMMM 6 hrs after mating
KVV 12 hrs after mating
180 24 hrs after mating
= 160
<
S 140
a
Ae 20
2
O 100
h—
> 80
©
zageasle
a
Y 40
a)
20
0
SOMATIC OOCYTES
TISSUE
Figure 2. Mean (+SE) disintegrations per minute of radiolabel found in female somatic and ovar-
ian tissue 6, 12, and 24 h after mating to radiolabelled males compared to nonradiolabelled controls.
Numbers within bars represent the number of short sperm + SE found in sperm storage organs at
these times after copulation (from Snook et al. 1994; see also text). Letters a—c denote significantly
different means detected by Tukey’s a posteriori multiple comparison tests; comparisons for somatic
tissue (letters a and b) were tested separately from oocytes (letter c).
time of transfer until more than 24 h after copulation when oviposition begins.
The numbers of sperm in the uterus were not determined due to the large amount
of sperm transferred by males to females. Thus, our data indicate that although
nonfertilizing sperm have been suggested to perform an adaptive role as nutrient
resources to fertilizing sperm, the female mate or the zygote (Hanson et al. 1952,
Healy & Jamieson 1981, Sivinski 1980, Silberglied et al. 1984), in D. pseudoob-
scura nonfertilizing short sperm do not function in this manner. Rather than prod-
ucts from the potential breakdown of short sperm, the origin of ejaculatory con-
tributions to females are nonsperm accessory gland materials.
One assumption in arriving at the above conclusion is that active transport or
exocytosis of labelled materials occurs across the intact sperm cell. To our knowl-
edge no evidence in Drosophila exists for or against specific transport mechansims
on sperm membranes. Movement of materials across short sperm could occur
prior to our detecting their “‘disappearance”’ as a result of breaking down. How-
1996 SNOOK & MARKOW: ROLE OF NONFERTILIZING SPERM 127
ever, the mechanism contributing to the disappearance of short sperm from SSOs
in D. pseudoobscura is unknown. The decrease in short sperm found in SSOs
may not be a result of these sperm types breaking down but of being pushed out
from storage by long fertilizing sperm, a mechanism currently being tested.
The location of radiolabel incorporation that we found differs from two earlier
reports in that we found significant incorporation into somatic rather than ovarian
tissues (Bownes & Partridge 1987, Markow & Ankney 1988). The “‘ovarian’’
tissues examined by Bownes & Partridge (1987) may have included other portions
of the female reproductive tract (uterus, ventral receptacle and spermathecae) that
contained radiolabelled sperm and accessory gland secretions. This discrepancy
in protocol between studies prevents any direct comparison of the differences in
results. Markow & Ankney (1988) used the same dissection procedures as in this
report, but utilized females that were well-fed, not starved. The nutritionally
stressed females in this report had few mature oocytes in comparison to well-fed
females (Snook, personal observation). Nutritional stress may cause competition
for the same resource between reproduction and survival, resulting in trade-offs
between these functions (Stearns 1992). Females in poor condition may allocate
energy into somatic maintenance, rather than reproduction, because oogenesis
requires substantial resources (Robertson & Sang 1944, Sang & King 1961). This
tradeoff may explain the difference between prior results (Markow & Ankney
1988) and ours. We conclude that patterns of utilization of male-derived sub-
stances appear to be influenced by the physiological condition of the female.
That tradeoffs occur are supported by the observation that females regulate
Oviposition behavior based on their nutritional status with increased oviposition
rates associated with better nutritional status (Sang & King 1961, Chapman et al.
1994). Moreover, Bownes & Partridge (1987) demonstrated that nutritionally
stressed females incorporated a greater percentage of radiolabel in somatic tissues
compared to well-fed females (somatic tissues examined were not affected by
potential contamination of radiolabelled sperm or accessory gland material). Other
obscura group species also show nutrient limitations on female reproduction. Male
D. subobscura Collin present an oral drop of liquid to females as part of courtship
behavior (Brown 1956a, b; Spieth 1966, 1978) and females denied access to male
drops have reduced fecundity (Steele 1986). Males of D. pseudoobscura also
present drops to females (Steele 1986) but only if other courtship behaviors have
failed (Brown 1956a, b). The effect of these drops on the reproductive fitness of
D. pseudoobscura females is unknown. However, Turner & Anderson (1983)
found that nutritionally stressed D. pseudoobscura females given continual access
to males had a relatively greater increase in productivity compared to nutritionally
stressed females only briefly exposed to males. Males could have improved the
productivity of these females through either oral drops or ejaculatory secretions.
Whether uptake of such substances by Drosophila females increases fecundity or
survival is debated (Markow et al. 1990, Chapman et al. 1994) but is likely to
be a species specific response based on ecology and other reproductive behaviors
of a particular species. Seminal products have increasingly been found to influence
female behavior and mortality (Chen 1984, Kalb et al. 1993, Chapman et al. 1994,
1995), sperm storage (Perotti 1971), and the outcome of sperm competition
(Harshman & Prout 1994, Clark et al. 1995) suggesting their importance irre-
spective of use as nutrient contributions.
128 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
The obscura group has evolved several mechanisms of male donation to fe-
males, including ejaculatory contributions (through accessory gland secretions)
and courtship feeding. However, the production and transfer of short nonfertilizing
sperm, at least measured by “C, is not among them. The role of nonfertilizing
sperm may be to function in sperm competition by giving first or second males
a fertilization advantage (Sivinski 1980, Silberglied et al. 1984, Snook 1995).
These alternatives are currently being tested.
ACKNOWLEDGMENT
We thank Scott Pitnick for help with the radiolabel dissections and John Alcock,
Christine Boake, Kathy Church, Thomas Dowling, and Michael Moore and two
anonymous reviewers for comments on prior versions of the manuscript. Research
presented was supported by a NSF Dissertation Improvement Grant (DEB-
9224263) to R.R.S.
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Received 1 Nov 1995: Accepted 14 Feb 1996
PAN-PACIFIC ENTOMOLOGIST
72(3): 130-137, (1996)
OBSERVATIONS ON THE FORAGING PREFERENCES OF
LEIOPROCTUS (FILIGLOSSA) RAYMENT
(HYMENOPTERA: COLLETIDAE)
IN EASTERN AUSTRALIA
PETER BERNHARDT! AND-K. WALKER?
‘Department of Biology, St. Louis University, St. Louis, Missouri 63103;
2Division of Natural Sciences, Museum of. Victoria;.71 Victoria Crescent,
Abbotsford, Victoria 3067, Australia
Abstract.—Fifty three bees representing three. species in subgenus Filiglossa [(Leioproctus davisi
Maynard (16 specimens), L. filamentosa (Rayment) Michener (36 specimens) and L. hamatus
Maynard (one specimen)] were collected on nine Persoonia species (Proteaceae) distributed
through. New. South Wales and Victoria. These: records extend the foraging range of L. (Fili-
glossa) to ten Persoonia Sm. species; P: arborea F Mueller, P. asperula L. Johnson & P. Weston,
P. champaepeuce Lhotsky ex Meissner, P. isophylla K. Johnson & P. Weston, P. lanceolata
Andrews, P. mollis subsp. nectans Krauss & L. Johnson, P. pinifolia L. Johnson & P. Weston,
P. silvatica L. Johnson, P. subvellutina L. Johnson and P. virgata R. Br.. Female bees outnum-
bered males by 70%. Fifty five per cent-of female bees and 43% of males carried significant
loads of Persoonia pollen. One, female, L. filamentosa carried pollen from more than one Per-
soonia species when Persoonia species were sympatric and had overlapping flowering periods.
Pollen analyses indicated that.some females. of L. davisi and males of L. filamentosa also foraged
on coblooming Asteraceae and Myrtaceae. The poor collections of L. (Filiglossa) bees between
1946 and 1991 were. possibly due to: the failure to identify a single grain of Persoonia pollen
carried by the holotype.of L. filamentosa.
Key Words.—Insecta, Leioproctus, Colletidae, Australia :
Until 1991 the Leioproctus (Filiglossa) filamentosa (Rayment) Michener (Col-
letidae) was known from only two specimens (Rayment 1959, Maynard 1994).
One was collected in New South Wales by N. Rodd in 1947 and the second was
collected in Queensland by J. Cardale in 1967. Neither collector appears to have
kept field notes which flowers were visited. by these bees.
Rayment (1959) erected the subgenus Leioproctus (Filiglossa) for three new
species collected by Rodd (filamentosa, striatula, proxima). Rayment emphasized
the excessive length of the mouthparts of these species suggesting “‘that the bees
are associated with an equally remarkable flower.’’ Michener (1965) moved both
striatula and promixma to Leioproctus (Euryglossidia).
Rayment’s: prediction of a remarkable flower has. never come true. All Leio-
proctus (Filiglossa) spp. have subsequently been collected on flowers of Per-
soonia Sm. since 1991 (Maynard 1994, Bernhardt & Weston in press). Persoonia
remains one of the most common genera‘of shrubby Proteaceae in Australia con-
sisting of approximately 90 species distributed through most coastal habitats (Wes-
ton 1991). The genus is treated as basal to the Proteaceae as Persoonia species
lack such characters as proteioid root systems and floral protostigmas, considered
apomorphic for most of the genera in the family (Johnson .& Briggs 1975).
The extraordinary mouthparts of L. (Filiglossa) do not even appear to be evi-
dence of a mutualistic coadaptation with Persoonia flowers as L. (Filiglossa) bees
are not important pollinators of Persoonia species. To the contrary, the majority
1996 BERNHARDT & WALKER: LEIOPROCTUS FORAGING 131
of bees that pollinate Persoonia flowers belong to Leioproctus (Cladocerapis)
species (Rayment 1950, Maynard 1992, Bernhardt and Weston in press).
Leioproctus (Cladocerapis) species appear to be facultative oligoleges on Per-
soonia flowers. The Persoonia pollen carried on their bodies is most likely to
contact the stigma of the flower while the female bee rakes pollen from the anther
slits or males and females probe the base of a flower for nectar. The Persoonia
species native to eastern Australia produce radially symmetrical and tubular flow-
ers. Each tepal is hinged at its base. When bees >6 mm long probe for nectar
they depress one of the four tepals on each flower and insert their heads down
the floral tube to collect nectar secreted by each of the four glands surrounding
the stalked ovary (Rayment 1950, Bernhardt & Weston in press).
Bernhardt & Weston (in press) observed L. (Filiglossa) species collecting pol-
len from the anthers of Persoonia flowers but noted that these female bees rarely
contacted the receptive stigma surrounded by the bases of the four anthers. Leio-
proctus (Filiglossa) species are less than 6 mm long and did not or could not
depress the Persoonia tepals. These bees were observed inserting their elongated
mouthparts between the seams of interlocking tepals at the apex of the floral tube.
Walker (unpublished) observed similar feeding behaviour of L. davisi Maynard
on Persoonia arborea F. Mueller.
Males of L. (Filiglossa) species observed on P. silvatica often avoided the
tube’s apex. They would insert their mouthparts at the base of the tube gaining
direct access to the nectar chamber but avoiding the sexual organs of the flower.
These observations suggest that the mouthparts of L. (Filiglossa) bees express a
trend towards nectar robbing.
To help elucidate the zoogeography and foraging preferences of L. (Filiglossa)
we present the following information. An updated list of Persoonia species on
which L. (Filiglossa) species have been observed and captured; a cross-reference
of bee species against the identifiable pollen they carried, and a re-examination
of the literature to suggest why these insects may have been so under collected
for over forty years.
MATERIALS AND METHODS
Study Sites.—Gungulla Flat, SE of Waterfall, Royal National Park, New South
Wales (NSW), 34°09'00” S 151°00’30" E alt. 150 m; dry sclerophyll forest; un-
derstory with P. pinifolia R. Br. (Plant Voucher, R. G. Coveny 15180) (Insect
collections, 7 Feb 1991, 20 Feb 1992, 12 Mar 1992, 20 Mar 1992).
One km South of Pikes Saddle, NSW, 36°59'40” S 149°34'00” E, alt. 1280 m;
dry sclerophyll woodland; understory with P. silvatica L. Johnson, P. chamae-
peuce Lhotsky ex Meissner (Plant Voucher, P. H. Weston 1762), P. asperula L.
Johnson & P. Weston (P. H. Weston 1763), (Insect collections, 19—20 Jan 1994).
Two km South of Pikes Saddle, NSW, 36°00’10” S 149°34’00” E, alt..1280 m;
dry sclerophyll forest; open understory with P. silvatica (Insect collection, 19—
20 Jan 1994).
Approximately 0.3 km N of Banksia Street, on West Road Fire Trail, Hill Top,
NSW, 34°20'30" S 150°29’00” E, alt. 560m; dry sclerophyll forest; shrubby un-
derstory with P. mollis subsp. nectens Krauss & L. Johnson (PHW 1775), P.
lanceolata Andrews (Plant voucher, P. H. Weston 1776), (Insect collection, 2 Feb
1994, 19 Feb 1994).
132 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Table 1. The collection record of Leioproctus (Filiglossa) species on Persoonia species in Queens-
land, New South Wales and Victoria since February 1991 (includes Maynard 1994).
Filiglossa species
Persoonia species davisi filamentosa hamatus prolatus
arborea see
. as perula =
chamae peuce a
isophylla a
lanceolata =
. mollis subsp. nectens mE
pinifolia =
. silvatica =
. subvellutina =
vir gata* =
Ww DDD DD DDD
++ tT ++ et4
|
|
++ = >10 specimens captured, + = 1-6 specimens captured, — = no specimens captured.
* The reference by Maynard (1994) recording L. filamentosa collected on P. pinifolia at Hastings
Point on 7/1i/91 is corrected here. Persoonia virgata grows at the Hastings Point site not P. pinifolia.
Island Bend—Guthega road, 1.5 km WSW of Island Bend rest area, NSW,
36°20'00" S 148°27'30" E, alt. 1280 m; dry sclerophyll forest; shrubby, grass
understory with P. subvellutina L. Johnson (Plant Voucher, P. H. Weston 1764)
(Insect collection, 24 Feb 1994).
Greta Road, 1.0 km W of junction with Bumble Hill Road, Bumble Hill, NSW,
33°14'30" S 151°14'45” E, alt. 340 m; dry sclerophyll forest; shrubby understory
with P. isophylla L. Johnson & P. Weston (Plant voucher, P. H. Weston 1781)
(Insect collection, 15 Mar 1994).
About 9-15 km W of Mt. Baw Baw, Victoria, 37°50’ S 146'17” E, alt. 930 m;
wet sclerophyll forest; shrubby understory and subcanopy of P. arborea FE Muell.
(Insect collection, K. Walker 7 Feb 1996).
Collection Methods and Pollen Analyses. Bees were netted and killed in jars
with ethyl acetate fumes. Insect vouchers have been deposited in the Museum of
Victoria, Melbourne. The single specimen of L. hamatus Maynard represents the
holotype (Maynard 1994). Flowering plant vouchers were deposited in the Royal
Botanic Garden, Sydney.
Methods for the removal and identification of pollen taxa from freshly killed
bees follows Bernhardt & Walker (1984). As L. (Filiglossa) bees were killed in
the same jar as other specimens of Hymenoptera that were taken while visiting
Persoonia flowers the smaller bodies of the L. (Filiglossa) bees could have be-
come contaminated with pollen dropped by the bodies of larger bees in the same
jar. Therefore a pollen taxon was not recorded as present on the body of a bee
unless >25 individual grains could be counted in each stained sample. Calberla’s
fluid was used to stain pollen and provide a semi-permanent mount (Ogden et.
al. 1974).
RESULTS AND DISCUSSION
The flowers of 20 Persoonia species were examined for the presence of L.
(Filiglossa) species. Ten Persoonia species attracted four L. (Filiglossa) species
(Table 1). Persoonia pinifolia and P. virgata R. Br. were the only shrubs studied
1996 BERNHARDT & WALKER: LEJOPROCTUS FORAGING 133
Table 2. Pollen load analyses of Leioproctus (subgenus Filiglossa) species collected on flowers of
Persoonia species.
Pollen load
Persoonia Persoonia + Other spp.
Taxon and gender only other spp. only No pollen
L. davisi
female 1 0 5 a
male 0 0 0 5
L. hamatus
female 1 0 0 0
L. filamentosa
female 18 1 0 6
male 5 0 4
Totals 26 3 5 20
Grand Total = 54
on which more than one L. (Filiglossa) species was captured. Leioproctus fila-
mentosa (Rayment) visited the greatest number of Persoonia species and appears
to have the widest distribution within the subgenus. Leioproctus davisi appeared
to be restricted to subalpine, arborescent populations of P. arborea in Victoria
and L. hamatus Maynard is still known from a single capture on P. pinifolia in
New South Wales.
The captures of L. filamentosa on Persoonia species (Table 1) suggested that
these bees may forage selectively on some Persoonia flowers. Persoonia mollis
R. Br. is one of the most broadly distributed species in New South Wales (Weston
1991, Krauss & Johnson 1991). Bernhardt & Weston (in press) collected bees on
five subspecies (ledifolia Cunn. ex Meissner) Kruss & Johnson, leptophylla
Krauss & L. Johnson, livens Krauss & L. Johnson, nectans, revoluta Krauss &
Johnson) of P. mollis, over three years at nine sites in New South Wales. Only a
single specimen of L. flamentosa was collected on P. mollis subspecies nectens
(Table 1).
Leioproctus filamentosa was collected on different Persoonia species in flower
at the same site (e.g., on P. asperula and P. chamaepeuce at Pike’s Saddle and
on P. lanceolata and P. mollis subsp. nectans at the Hilltop site). Due to the
difference in size and grain morphology of the pollens of P. asperula and P.
chamaepeuce, it was possible to determine that one female of L. filamentosa had
visited the flowers of P. champaepeuce before it was netted on P. asperula.
Leioproctus (Filiglossa) species were collected on Persoonia flowers from 0900
h until 1500 h with peaks in floral visitation from 1000 until 1300 h. The greatest
number of bees were collected on Peroonia species in shady sites. The individual
shrubs or small trees that seemed to be most likely to be visited by L. (Filiglossa)
grew directly under the canopy (eg. P. asperula, P. chamaepeuce, P. silvatica)
or stood in light gaps adjacent to dense, shadier sections of eucalypt woodland
(eg. P. lanceolata, P. pinifolia).
Of the 53 specimens collected on Persoonia flower 54% carried significant
loads (>25 grains/specimen) of Peroonia pollen (Table 2; Fig. 1). Over 16% also
carried significant loads of pollen that did not belong to Persoonia flowers. These
134 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Figure 1. Hydrated pollen of Persoonia chamaepeuce carried by female L. filamentosa collected
on P. chamaepeuce. A, regular, triporate grain; B, irregular, tetraporate grains (<511).
Figure 2. Head of female L. filamentosa collected on P. silvatica with pollen grains clinging to
elongated palps (X40).
grains were identified most often on bees collected on P. arborea and P. subvel-
lutina and belonged primarily to coblooming Asteraceae and Myrtaceae (Table
3). The presence of grains of Asteraceae on L. davisi’suggest that the distribution
of this species may be limited by topography and altitude and not by any spe-
cialization on P. arborea for pollen or nectar.
Females (55%) carried Persoonia pollen more often than males (43%; Table
2). Males of L. filamentosa were observed to either circle the shrubs in flower or
Table 3. Pollen load analyses of L. davisi and L. filamentosa carrying Persoonia pollen mixed with
pollen of other species or no Persoonia pollen at all.
Pollen taxon
Bee taxon
and gender Ast Myrt Persoonia Irid-Type UD*
L. davisi
female 5 0 0 0 0
L. filamentosa
female 0 0) 1 1
male 0 be 2 1
* Ast = Asteraceae (Brachycome and Hypochaeris types); Myrt = Myrtaceae (Eucalyptus type);
Irid-Type (unidentified monocot, similar to Iridaceae); UD = unidentified dicot (tricolporate).
1996 BERNHARDT & WALKER: LEJOPROCTUS FORAGING 135
B
3
Figure 3. Illustrations of Persoonia-type pollen grains as drawn by T. Rayment. A = pollen grains
from Rayment (1950) showing regular, hydrated grains (arrows) outnumbered by irregular and/or
nonhydrated grains from flowers of P. lanceolata; B = pollen grain from Rayment (1959) showing
an operculum over each pore. Rayment never included a scale in either publication and drawings here
are reproduced twice the size of the originals.
hovered six to eight cm down wind of flowering branches. Males were not ob-
served to forage actively for pollen, but they did contact dehiscent, anthers when
perching on the flowers or while inserting their elongated mouthparts between the
tepals at the apex of the floral tube. Pollen washes showed that only one male
captured carried more than 90 grains of Persoonia pollen. In contrast, one female
of L. davisi, one female of L. hamatus and fifteen females of L. filamentosa carried
>100 grains of Persoonia each distributed in their scopae. Persoonia grains are
often visible clinging to the elongated mouthparts of pinned specimens (Fig. 2).
Why has this bizarre colletid evaded entomologists. for so many decades? One
reason may be that Rayment (1959) was unable to identify the pollen grain found
on the holotype of the male, L. (Filiglosa) filamentosa. In reviewing the original
paper Rayment’s drawing (Figure 3) shows a triangular-trilobate grain with three,
large, operculate, pores produced by most of the Proteaceae in Australia (Johnson
& Briggs 1975, Feuer 1986). Rayment’s stippling of the grain suggests his mi-
croscope was sufficiently powerful to detect the scabrous ornamentations on the
outer, pollen wall that are found throughout the genus, Persoonia (Feuer 1986).
Within the same paper, Rayment (1959) also suggested that large loads of pollen
found on L. (Euryglossidia) proxima had also come from a member of the Myr-
taceae (e.g., Leptospermum). Although the pollen of Leptospermum is triangular
it lacks operculate pores and a scabrous pollen wall. Rayment’s drawing of the
pollen grain found on L. (Euryglossidia) proxima is almost identical to his draw-
ing of the grain found on L. (Filiglossa) filamentosa (Fig. 3). If entomologists
ever followed Rayment’s suggestion and looked for L. filamentosa on myrtaceous
flowers they would have attempted to collect this bee on blossoms. it does not
appear to visit with any great frequency (Table 3).
Rayment’s difficulties with Persoonia pollen is most surprising considering his
strong commitment to illustrating pollen grains found on bees and/or stored in
their nests. A most self-consistent and dependable aspect of his. fieldwork was
that he identified the flower on which the bee was caught and then compared the
pollen grains in the flower to grains removed from the bee’s body and/or its pollen
loaf (Rayment 1935). In fact, toward the end of his life Rayment completed. a
manuscript on the biology of the Australian heaths (Epacridaceae) containing
136 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
dozens of meticulous illustrations of pollen morphology, wall sculpture and ger-
mination (Rayment 1961). Genera within the Epacridaceae release their pollen
grains in united packets of four known as tetrads. Rayment clearly discriminated
between four, distinct modes of tetrad configuration.
In addition, Rayment (1950) also published a paper describing the pollination
of Persoonia mollis by Leioproctus (Cladocerapis) species only nine years before
his description of Leioproctus (Filiglossa) filamentosa (Rayment 1959). In this
earlier paper Rayment emphasized the foraging behavior of Leioproctus (Clado-
cerapis) species on Persoonia flowers. As usual, he carefully drew Persoonia
grains Norman Rodd provided from flowers of Persoonia lanceolata.
The drawing of Persoonia pollen in Rayment (1950) is based on a wet mount
and shows that the author was familiar with the shape of hydrated, abortive and
irregular-tetraporate grains (fig. 3). Compared to a modern photomicrograph (Fig.
1) all of Rayment’s illustrations show the broad variation in the shape of Per-
soonia pollen.
In the case of Leioproctus (Filiglossa) species only two specimens, representing
a single species, were collected for over forty years (Maynard 1994). Perhaps
there is a lesson to be learned when sampling certain taxa that are obligate, floral
foragers. When new species are caught in random sweeps, or lack fieldnotes as
to foraging preferences, the success of future collections may be delayed. It may
be easier to relocate these insects when distinctive pollens can be removed from
the original specimens and identified using both standard palynological references
and a back up collection of mounted grains or their photomicrographs.
ACKNOWLEDGMENT
Research was funded by a Fellowship provided by the Trust of the Royal
Botanic Gardens Sydney. We thank Dr. P- H. Weston (RBG Sydney) for his iden-
tification of Persoonia species and for showing us the pertinent sites in New South
Wales and particular thanks to Ms. Natasha Baczocha and Ms. Catriona McPhee
who assisted in the collection of bees and helped record data and prepare speci-
mens over two seasons.
LITERATURE CITED
Bernhardt P. & K. Walker. 1984. Bee foraging on three species of Australian Acacia. Int. J. Entomol.,
26: 322-30.
Bernhardt, P. & P. Weston. In press. The pollination ecology of Persoonia in Eastern Australia.
Telopea.
Feuer, S. 1986. Pollen morphology and evolution in the Persoonioideae, Sphalmioideae and Carnar-
vonoioideae (Proteaceae). Pollen et Spores, 28: 123-156.
Johnson, L. A. S. & B. G. Briggs. 1975. On the Proteaceae—the evolution and classification of a
southern family. Bot. J. Linn. Soc.: 83-182.
Krauss, S. L. & L. A. S. Johnson. 1991. A revision of the complex species Persoonia mollis (Pro-
teaceae). Telopea, 4: 185-200.
Maynard, G. 1992. Revision of Leioproctus (Cladocerapis) Cockerell (Hymenoptera: Colletidae). J.
Aust. ent. Soc., 31: 1-11.
Maynard, G. 1994. Revision of Leioproctus (Filiglossa) Rayment (Hymentoptera; Colletidae). J. Aust.
ent. Soc., 33: 299-304.
Michener, C. D. 1965. A classification of the bees of the Australian and South Pacific regions. Bull.
Amer. Mus. Natur. Hist., 130: 1-362.
1996 BERNHARDT & WALKER: LEJOPROCTUS FORAGING 137
Ogden, E. C., G. S. Raynor, J. V. Haynes, D. M. Lewis & J. H. Haines. 1974. Manual of Sampling
Airborne Pollen. Hafner Press, Collier Macmillan, New York.
Rayment, T. 1935. A cluster of bees. The Endeavour Press, Australia.
Rayment T. 1950. New bees and wasps-part XV. The Victorian Naturalist, 67: 101-111.
Rayment T. 1959. A new genus of bees in the family Colletidae. Aust. Zool., 12: 324-329.
Rayment, T. 1961. Heaths of Australia; Their flower structure, pollination and evolutionary sequence
with a census of the family. Parts 1 & 2. Royal Society of Victoria’s papers. Manuscript
Collection, State Library of Victoria, Melbourne, Australia.
Weston, P. 1991. Persoonia. In, Flora of New South Wales, Vol. 2, pp. 4-19. Jn Garden, G. W. (ed.).
New South Wales U. Press, Kensington, N.S.W. Australia.
Received 18 Aug 1995: Accepted 1 Mar 1996
PAN-PACIFIC ENTOMOLOGIST
72(3): 138-141, (1996)
EFFECTS OF SEX RATIO AND FEMALE DENSITY ON
PROGENY SURVIVAL OF THE ALFALFA LEAFCUTTER
BEE (HYMENOPTERA: MEGACHILIDAE)
D. F MAYER
Department of Entomology, Irrigated Agriculture Research and Extension
Center, Washington State University, Prosser, Washington 99350
Abstract.—This paper reports the results of two 3-year studies on the effects of: 1) sex ratio in
the parental generation on percent live larvae and immature mortality in the progeny; and 2) the
density of females on percent live larvae and immature mortality in the progeny. In both studies
there were no consistent significant differences between treatments in percent live larvae or
immature mortality in the progeny.
Key Words.—Insecta, alfalfa leafcutter bee, Megachile rotundata, alfalfa seed
The alfalfa leafcutter bee, Megachile rotundata (Fabr.), is the primary pollinator
of commercial alfalfa seed in the Pacific Northwest of the United States and
southwestern Canada (Richards 1984, Mayer et al. 1990). U.S. alfalfa seed grow-
ers in the U.S. often purchase leafcutter bees in 3.79 liter units (gallon) (about
10,000 cells) from producers in Canada because, in many cases, these bees do
not reproduce well when used for alfalfa pollination in the U.S.
A major problem with the alfalfa leafcutter bees bred in the U.S., other than
chalkbrood, is the high death rate of eggs and young bee larvae (immature mor-
tality) which has reached 60% or more (Bohart 1972). Larval mortality has been
attributed to a variety of factors including insecticide residues (Waller 1969),
nutrition (Bohart 1972) and parasitism, bee senility, unidentified diseases, over-
crowding in domiciles resulting in bee confusion, type of nesting media, lack of
food resources and excessive competition for these resources (Amett 1981). Goesk
et al. (1988) found the ratio of males to females had an effect on percent females
and I thought there may be some other effects on progeny.
This paper reports the results of two 3-year studies on the effect of the sex
ratio in the parental generation and female density on percent live larvae and
immature mortality.
METHODS AND MATERIALS
New sterilized nest blocks were prepared annually by taping laminate wood
pieces (1 cm X 13 cm X 12 cm) together with strapping tape to form small blocks
with 104 nesting tunnels, and covering the back of each block with aluminum
foil. Tunnels were 5 mm in diameter and 12 cm deep. One nest block was placed
in each cage.
Loose bee cells were obtained annually from Mr. Pollination Services in Canada
during the winter and stored at 3° C for about 36 weeks. These bees contained
no chalkbrood. In the spring, the cells were removed from storage and incubated
at 28-29° C. Adults that emerged after about 19-21 days were allowed to fly in
the laboratory and males and females were counted and collected into separate
vials. The adults were then released into the cages containing blooming alfalfa
and the nest blocks.
1996 MAYER: MEGACHILE LARVAL SURVIVAL 139
Table 1. Effect of number of male leafcutting bees on percent live larvae (LL), immature mortality
(IM), dead mature larvae (DL) and pollen masses (PM). Prosser, WA.
1990 1991 1992
¢iales eS ee ee — — al
to males LL IM DL PM LL IM DL PM LL IM DL PM
1:6 89a 6a Sa Oa 59a 26ab 3a 12a 65a 25a 2a 8a
1:3 T7a 15a 7a la 66a 23a la 10a 60a 28ab 4a 8a
1:2 86a 8a 4a Oa 56ab 3lab Sa 8a 58a 35b 3a 4a
521 86a 8a 6a Oa 47b 42b 5a 6a 6la 32ab la 6a
Means within a column followed by the same letter are not significantly different at the P = 0.05
level, Newman-Keuls studentized range test.
For the sex ratio study, 16 cages (6 X 6 X 1.8 m) and for the female density
study, 12 similar cages were erected over different plots of blooming alfalfa at
Prosser, WA. For sex ratio studies, 80 females were put in each cage and then
males were added to obtain a 6:1 male to female ratio in each of 4 cages, 3:1
ratio in each of 4 cages, 2:1 ratio in each of 4 cages, and 1:5 ratio in each of 4
cages. This method led to differences in male density between treatments. How-
ever, males feed only on nectar and nectar is constantly produced by alfalfa flow-
ers until the flower is tripped. For female density studies 52 females were put in
each of 4 cages (1 female:2 tunnels), 104 females were put in each of 4 cages
(1 female: 1 tunnel) and 208 females in each of 4 cages (2 females: 1 tunnel).
Males were put in the cages at the same time and in equal number to females (1:
1). Thus both nesting density effects and provisioning resources were limitations.
Bees were put in the cages on 30 Jul 1990; 5 Jul 1991; and 15 Jul 1992.
The bees foraged and constructed cells in the nest blocks during each season.
At the end of the nesting season (August) all the bee cells were extracted from
the laminate boards and put into cold storage at 3° C. During each winter the
cells were cut open, inspected and the number of live prepupae, dead eggs or
young larvae (instars 1-3), dead older larvae and pollen masses (no visible egg
or larva) recorded. I examined all the cells produced each year.
The data were analyzed as a randomized complete block design after transfor-
mation by analysis of variance, with Newman-Keuls studentized range test for
mean separations (Lund 1989).
RESULTS AND DISCUSSION
The total number of cells produced for the sex ratio studies were 1849 (6:1
ratio), 2008 (3:1 ratio), 2080 (2:1 ratio), and 2061 (1:5 ratio). The total number
of cells produced for the female density studies were 1492 (1 female: 2 tunnels),
1282 (1 female: 1 tunnel) and 767 (2 females: 1 tunnel).
There were no consistent significant differences among sex ratio treatments in
the percent live larvae, mortality of immature stages, dead mature larvae or pollen
masses (Table 1). In 1990, there were no significant differences between treat-
ments in live larvae (F = 1.37, P = 0.314, SE = 3.64), immature mortality (F
= 1.85, P = 0.195, SE = 1.86) dead mature larvae (F = 0.47, P = 0.709, SE =
2.54) or pollen masses (F = 2.08), P = 0.173, SE = 0.364). In 1991, there were
no significant differences between treatments in immature mortality (F = 2.45, P
= 0.131, SE = 5.27), dead larvae (F = 2.35, P = 0.141, SE = 1.84) or pollen
140 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Table 2. Effect of number of female leafcutting bees per tunnel on percent live larvae (LL), im-
mature mortality (IM), dead mature larvae (DL) and pollen masses (PM). Prosser, WA.
Females 1990 1991 1992
tunnel LL IM DL PM LL IM DL PM LL IM DL PM
1:2 82a Ta 8a 3a 54a 33a 7a 6a 60a 25a 7a 7a
1:1 82a 9a 6a 3a 50a 37a 4a 9a 54a 32a Sa 9a
2:1 87a 6a 5a 2a 52a 32a 6a 9a 56a 36a Sa 3a
masses (F = 1.61, P = 0.254, SE = 1.91). However, there were significantly
fewer live larvae (F = 0.49, P = 0.700, SE = 3.5) in the 1:5 sex ratio as compared
to the other treatments. Though not significant there appeared to be more im-
mature mortality in this treatment. In 1992, there were no significant differences
between treatments in live larvae (F = 1.98, P = 0.296, SE = 4.32), dead mature
larvae (F = 2.04, P = 0.149, SE = 0.48) or pollen masses (F = 2.12, P = 0.135,
SE = 1.85). However, there were significant differences between treatments in
immature mortality (F = 12.5, P = 0.018, SE = 4.73) although there were no
apparent correlations with the other two years of the study. It appears that im-
mature mortality is the major variable factor and dead mature larvae and pollen
masses are of lesser importance. Mortality of immature stages appeared to be
somewhat higher in 1991 and 1992 as compared to 1990. The sex ratio in a
population appears to have little or no effect on mortality of the immature stages
of their progeny.
Female density had no significant effect among treatments in 1990 in the per-
cent live larvae (F = 0.81, P = 0.818, SE = 2.74), immature mortality (F = 2.46,
P = 0.432, SE = 1.26), dead mature larvae (F = 3.03, P = 0.123, SE = 1.24)
or pollen masses (F = 0.47, P = 0.646, SE = 0.76) (Table 2). Female density
had no significant effect among treatments in 1991 in the percent live larvae (F
= 0.21, P = 0.818, SE = 3.65), immature mortality (F = 0.09, P = 0.911, SE
= 3.55), dead mature larvae (F = 1.45, P = 0.307, SE = 1.25), or pollen masses
(F = 3.47, P = 0.10, SE = 1.36). Female density had no significant effect among
treatments in 1992 in the percent live larvae (F = 0.62, P = 0.561, SE = 2.85),
immature mortality (F = 3.05, P = 0.185, SE = 3.21), dead mature larvae (F =
1.26, P = 0.285, SE = 1.020, or pollen masses (F = 2.48, P = 0.408, SE =
3.4).
Mortality of immature stages does not appear to be affected by different sex
ratios or the female density and the results were fairly consistent over three years.
ACKNOWLEDGMENT
I thank J. D. Lunden and M. R. Jasso for their help and the Washington Alfalfa
Seed Commission and the USDA/ARS for partial funding of this research.
LITERATURE CITED
Amett, W. H. 1981. Factors influencing egg and early larval leafcutting bee mortality (pollen mass).
Proc. Ann. Interstate Alfalfa Seed Growers School, 12: 1-3.
Bohart, G. E. 1972. Management of wild bees for the pollination of crops. Ann. Rev. Entomol., 17:
287-312.
Gosek, J., A. Ruszkowsi, M. Bilinski & K. Kaczmarska. 1988. The percentage of females in the
1996 MAYER: MEGACHILE LARVAL SURVIVAL 141
progeny of the lucerne leafcutter bee (Megachile rotundata) in relation to the percentage of
females in the parent generation and to the breeding line and population density. Pszczelnicze
Zeszty Naukowe, 32: 99-110. (In Polish.)
Lund, R. E. 1989. MSUSTAT statistical analysis package 4.12. Montana State University, Bozeman,
Montana.
Mayer, D. F, J. D. Lunden & E. R. Miliczky. 1990. Effects of fungicides on chalkbrood disease of
alfalfa leafcutting bee. Appl. Agr. Res., 5: 223-226.
Richards, K. W. 1984. Alfalfa leafcutter bee management in Western Canada. Agr. Canada Publ.
1495/E. 53 pp.
Waller, G. D. 1969. Susceptibility of an alfalfa leafcutting bee to residues of insecticides on foliage.
J. Econ. Entomol., 62: 189-92.
Received 18 Aug 1995: Accepted 14 Feb 1996
PAN-PACIFIC ENTOMOLOGIST
72(3): 142-144, (1996)
A NEW MICROBEMBEX ENDEMIC TO THE ALGODONES
DUNES, CALIFORNIA (HYMENOPTERA: SPHECIDAE)
TERRY L. GRISWOLD
USDA-ARS Bee Biology & Systematics Lab,
Utah State University,
Logan, Utah 84322-5310
Abstract—Microbembex elegans, NEW SPECIES, a sand wasp endemic to the Algodones
Dunes, California, is described. This species appears to be restricted to areas of the dunes with
a combination of active slip faces and perennial vegetation. Known populations are in areas
subject to off-road vehicle activity, which may endanger the future of this wasp.
Key Words.—Insecta, Hymenoptera, Sphecidae, endemic, endangered, sand wasp, sand dunes
Microbembex Patton is a small genus of New World bembicine wasps which
frequent sandy environments, making shallow nests in the soil which are provi-
sioned with dead and moribund arthropods (Evans 1966). In contrast to South
American Microbembex where there are four distinct species groups (Bohart &
Willink 1989), known North American species are frustratingly uniform morpho-
logically, with species discrimination often dependent on rather variable color and
punctation. Bohart & Horning (1971) revised the genus for the continental United
States. Study of over 2000 specimens demonstrated that all but one of the seven
included species are widespread. (The only exception is M. rufiventris Bohart,
known only from two sites in the southern San Joaquin Valley.) Collections were
from throughout the deserts of California. The presence of a new species of
Microbembex endemic to the Algodones Dunes (or Glamis Dunes) is, therefore,
surprising. This new species, Microbembex elegans, appears to be restricted even
within the dune system to areas with active slip faces. The wasp was found only
around the bases of shrubs where detritus collects. Microbembex elegans is like-
wise remarkable for the distinctive morphological characters separating it from
all other species.
Microbembex elegans may be threatened by off-road vehicle activity. The two
known populations were in areas open to such activity but not currently receiving
a high level of disturbance. At the time the type material was collected search of
stabilized dunes and areas of high vehicular use devoid of vegetation failed to
yield M. elegans.
In the description which follows, terga are numbered T1, T2, ..., sterna, S1,
S2, etc.
Microbembex elegans Griswold, NEW SPECIES
(Figs. 1-5)
Types.—Holotype, male; CALIFORNIA. IMPERIAL Co.: Glamis Dunes, 1.6
km (1 mi) W of Glamis, 9 Oct 1988, T. Griswold; deposited: USDA Bee Biology
and Systematics Laboratory, Logan, Utah. Paratypes: same data as holotype, 3
males, 4 females; same except 7 Oct 1988, 1 male; same except 29 Sep 1987, 1
male; 6.4 km (4 mi) S Ogilby, 10 Oct 1988, T. Griswold, 1 male; deposited:
USDA Bee Biology and Systematics Laboratory, Logan, Utah.
1996 GRISWOLD: NEW MICROBEMBEX SPECIES 143
Figures 1-6. Microbembex elegans. Figure 1. Male genitalia, dorsal view. Figure 2. Abdomen in
ventral view, S2—8. Figure 3. Male S7. Figure 4. Male S8, lateral view. Figure 5. Male midleg, posterior
view. Figure 6. Male midcoxa, posterior view.
Male.—Length 12, mm; forewing length 8 mm. Head and thorax black marked with pale yellow on
scape, pedicel, first flagellar segment, narrow line around eye interrupted dorsally, most of pronotum,
lateral margin of scutum, scutellum except for anterior triangle, metanotum, transverse band on pos-
terior face of propodeum and most of lateral face, dorsal mesopleural spot, legs except bases of coxae;
mandible except apically, clypeus, labrum white. Abdomen yellow with amber basal marks on terga
and sterna except S2. Wings clear, veins in basal half of wing except radial vein white, apical veins
brown. Pubescence silvery, dense, appressed, obscuring sculpture on pronotal lobe, dorsum of meso-
soma, mesopleuron. Punctation of entire body dense. Flagellar segments 6—11 with shiny raised inner
welts; length of segment 11 less than 2 X width. Forecoxa with small apical spine. Foretarsi slender,
unmodified, basitarsus with 5 rake setae. Midcoxa with inner longitudinal carina ending in distinct
spine (Fig. 6). Ventral surface of midfemur concave basally, thickened and ridged apically (Fig. 5).
Midbasitarsus with carinate ventral lobe (Fig. 5). Hindfemur concave basoventrally. Hindbasitarsus
with carinate ventral lobe. T7 with apical margin broadly rounded, not emarginate medially. S2 without
projection; S3 with pronounced rounded lateral lobe; S4 with similar but smaller lobe (Fig. 2). Apical
margin of S7 emarginate laterally, fringed with long hair (Fig. 3). S8 with apical spine bearing very
long hair, with slender dorsal basal spine (Fig. 4). Genitalia as in Fig. 1.
Female.—Length 9-10 mm; forewing length 7—7.5 mm. Markings as in male except mesosomal
markings more nearly white; amber markings of sterna present only on S4—5; wide apical white bands
on T1—5. Foretarsal rake setae pale. Clypeus evenly arched, not strongly projecting medially. Labrum
densely punctate. Mouthparts not exceeding labrum. Forecoxa with projecting obtuse apical angle.
Midcoxa with weak inner longitudinal carina. T6 densely punctate throughout, apical notch weak,
very narrow.
Diagnosis.—Microbembex elegans differs from all other North American Mi-
crobembex by the carina on the midcoxa which, though strongest in the male, is
present in both sexes. Males are unique among Microbembex in the modified mid-
and hind legs, the lateral humps of S3—4, the shape of S7, and the dorsal spine
of S8. Females differ from all other North American species by the densely punc-
tate clypeus, the shape of the clypeus, the presence of an apical process on the
144 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
forecoxa, and the densely punctate and narrowly notched T6. They further differ
from those in the continental United States by the all white rake setae on the
foretarsi.
Discussion.—Microbembex elegans differs from the generic diagnosis of Mi-
crobembex (Bohart & Menke 1976) in several characters. The clypeus is evenly
convex, the male midleg is modified, the apical spine of male S8 is not down-
curved and a basal dorsal spine is present. Despite these differences, M. elegans
clearly belongs in Microbembex. The midocellus is reduced to a transverse scar,
the marginal cell is bent away from the margin of the forewing, the propodeum
is not armed with lateral angles, and male T7 lacks spiracular lobes.
Variation.—A small male (8 mm long) has the mesosomal markings reduced,
the amber abdominal markings replaced by more extensive dark brown markings
and the light abdominal markings pale yellow.
Material Examined—See Types.
ACKNOWLEDGMENT
Thanks to Marianne Cha Filbert and Greg Frehner for producing the illustra-
tions. R. M. Bohart and B. A. Alexander kindly reviewed the manuscript. This is
a contribution from Utah Agricultural Experiment Station, Utah State University,
Logan, UT 84322-4810, Journal Paper No. 4545, and USDA-ARS Bee Biology
and Systematics Laboratory, USU, Logan, UT 84322-5310.
LITERATURE CITED
Bohart, R. M. & D. S. Horning, Jr. 1971. California Bembicine sand wasps. Bull. Calif. Insect Surv.,
13: 1-49.
Bohart, R. M. & A. S. Menke. 1976. Sphecid wasps of the world, a generic revision. Univ. Calif.
Press, Berkeley.
Bohart, R. M. & A. Willink. 1989. The genus Microbembex in Argentina. Trans. Amer. Entomol.
Soc., 114: 295-310.
Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps. Harvard Univ. Press,
Cambridge.
PAN-PACIFIC ENTOMOLOGIST
72(3): 145-151, (1996)
TAXONOMIC CHANGES IN CHINESE LOPHOPIDAE
WITH A CHECK LIST OF CHINESE SPECIES
(HOMOPTERA: FULGOROIDEA)
AI-PING LIANG!
Department of Entomology, American Museum of Natural History,
New York, New York 10024-5192
Abstract—The monotypic genus Boresinia Chou is removed from Lophopidae and synonymized
with Asiraca Latreille of the subfamily Asiracinae of Delphacidae, and Boresinia choui Yuan &
Wang becomes Asiraca choui (Yuan & Wang), NEW COMBINATION. Lacusa yunnanensis
Chou & Huang is synonymized with L. fuscofasciata (Stal). Bisma elasmoscelis Jacobi is trans-
ferred to Serida and new distributional data for this species is given. The sugarcane and rice
pest Pyrilla perpusilla (Walker) and the species Pitambara dawnana Distant are reported for the
first time in southern China. A check list of the Chinese species of Lophopidae is provided.
Key Words.—Insecta, Fulgoroidea, Lophopidae, Delphacidae, China, new synonymies, new com-
binations, new records, check list
The Lophopidae is a very small family of Fulgoroidea with 44 genera and 137
species known. The world fauna is confined to the tropics and warm temperate
regions. Four genera and four species were reported from China (Chou et al.
1985). The family can be recognized by the following combination of characters:
head narrower than pronotum; vertex much narrower than its length, sometimes
protuberant, lateral margins strongly elevated; frons longer than wide, with carinae
in addition to angulate margins; clypeus with lateral carinae; rostrum short, stout,
with the apical joint short; pronotum and mesonotum short and broad, tricarinate;
tegulae large; forewings with apical margin narrower, much shorter than anal
margin, clavus not extending to apical part of forewing; fore and middle tibiae
usually compressed and expanded, second hind tarsomere very small and without
apical row of spines; male aedeagus robust with a complicated set of apical spines;
and female genitalia incomplete.
During the preparation of a checklist of Fulgoroidea found in China and a
review of the genera of Chinese Delphacidae, it became evident that several tax-
onomic changes in the Lophopidae were necessary. This paper indicates these
changes. These include one generic synonym, one new specific synonym, one
species transferred to the correct genus and another species to Delphacidae. There
are also additions of one newly detected sugarcane and rice pest and another
species. A check list summarizing the nomenclatural and distributional knowledge
about Chinese species of Lophopidae is also provided.
Depositories.—Specimens studied here are deposited in the following institu-
tions: American Museum of Natural History, New York, USA (AMNH); Institute
of Zoology, Academia Sinica, Beijing, China (IZAS); Department of Entomology
Insect Collection, North Carolina State University, Raleigh, North Carolina, USA
(NCSU); Entomological Museum, Northwestern Agricultural University, Yang-
' Present address: Department of Entomology, Institute of Zoology, Academia Sinica, 19 Zhong-
guancun Lu, Beijing 100080, People’s Republic of China.
146 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
ling, Shaanxi, China (NWAU); and Shanghai Institute of Entomology, Academia
Sinica, Shanghai, China (SIE).
NEw GENERIC SYNONYMY, AND SPECIES
EXCLUDED FROM LOPHOPIDAE
Asiraca Latreille
Cercopis Latreille 1796: 91. Type species: Cicada clavicornis Fabr. 1794: 41, by
subsequent designation of Latreille 1810: 434. [Homonym of Cercopis Fabr.
1775.]
Asiraca Latreille 1796: 12, 202. [Replacement name for Cercopis Latreille.]
Manchookhonia Kato 1933: 10. Type species: Manchookhonia granulipennis
Kato 1933: 11, fig. 13, by original designation and monotypy. [Synonymized
by Asche 1985: 116, 374.]
Boresinia Chou in Chou et al. 1983: 60, 66. [Replacement name for Manchoo-
khonia Kato.) [Synonymized with Manchookhonia Kato, 1933: 10 by Dessart
1983: 318.] NEW SYNONYMY.
Discussion.—The generic name Boresinia was proposed by Chou (See Chou et
al. 1983) to replace Manchookhonia, which was described by Kato (1933) in
Lophopidae for the Manchurian species M. granulipennis Kato (see Metcalf
1955). Dessart (1983: 318) refused Chou’s (1983) action and treated Boresinia as
a new synonymy of Manchookhonia Kato. Asche (1985: 116, 374) synonymized
Manchookhonia Kato with the delphacid genus Asiraca Latreille. Thus, Boresinia
Chou is a junior subjective synonym of Asiraca Latreille.
Asiraca choui (Yuan & Wang), NEW COMBINATION
Boresinia choui Yuan & Wang 1992: 179 (Chinese), 182 (English), fig. 1-A-D.
Holotype, male, China ‘Shaanxi’ (NWAU) [examined]. Transferred to Asiraca
Latreille of the Asiracinae of Delphacidae.
Discussion.—Boresinia choui was described by Yuan & Wang (1992) in Lo-
phopidae on the basis of four specimens, two males and two females, from
Shaanxi Province, China. I have examined the holotype of Boresinia choui and
believe that it is not a lophopid, and that it is clearly a member of Asiraca Latreille
of the Asiracinae of Delphacidae.
Asiraca, with two known species, occurs in the Palaearctic region. The transfer
of B. choui to Asiraca brings the total number of known species in the genus to
three and represents the far southern range of this primitive delphacid genus (Asche
1985, 1990).
Distribution.—China (Shaanxi Province).
Type Material Examined—Holotype, male, CHINA. SHAANXI: Mt. Qinling, 8 May 1980, L.-C.
Xiang & N. Ma (NWAU). Paratypes: CHINA. SHAANXI: same data as holotype, 1 male, 2 females
(NWAU).
NEw SPECIFIC SYNONYMY AND NEW COMBINATION
Lacusa fuscofasciata (Stal)
Elasmoscelis fuscofasciata Stal 1854: 248.
Cixius eminens Walker 1858: 42. [Synonymized by Atkinson, 1886: 42.]
1996 LIANG: CHINESE LOPHOPIDAE 147
Lacusa fuscofasciata (Stal); Stal 1862: 309.
Lacusa yunnanensis Chou & Huang in Chou et al. 1985: 128 (Chinese), 137
(English), fig. 119a, b. Holotype, female, China ‘Yunnan’ (NWAU) [examined].
NEW SYNONYMY.
Discussion.—Lacusa fuscofasciata (Stal) is common and widely distributed in
northeastern India, northern Myanmar (Burma) and southwestern China. Chou &
Huang (see Chou et al., 1985) described Lacusa yunnanensis on the basis of a
single female from Yunnan Province in southwestern China. They noted that L.
yunnanensis was closely related to L. fuscofasciata, but that it could be distin-
guished from it by the fuscous transverse band near outer margin on forewing
branched and the fuscous meso- and metathorax. My examination of specimens
of L. fuscofasciata from Yunnan, Guizhou, Guangdong, and Hainan Provinces in
southern and southwestern China, and Chou & Huang’s female holotype of L.
yunnanensis shows that the latter represents part of a range of variation in color
morph within L. fuscofasciata (Stal). I, therefore, propose L. yunnanensis as a
new synonym of L. fuscofasciata.
Distribution.—India, Myanmar (Burma), China (Guangdong, Guizhou, Hainan,
and Yunnan Provinces).
Type Material Examined.—Holotype, female (of Lacusa yunnanensis Chou &
Huang 1985), CHINA. YUNNAN: Xishuangbanna, Menglun, 21—30 Apr 1974, I.
Chou, E Yuan & Y.-Y. Hu (NWAUD).
Other Specimens Examined.—CHINA. E. GUANGDONG [KWANTUNG]: Yim
Na San, 17 Jun 1936, J. L. Gressitt, 1 female (NCSU). GUIZHOU: Huangguoshu,
24 Jul 1958, D.-Y. Bi & Ren, 2 males (SIE). HAINAN: Ta Hau, 7 Jul 1935, J.
L. Gressitt, 1 male (NCSU); Yinggen, 200 m, 8 Jul 1960, C.-Q. Li, 1 female
(ZAS). YUNNAN: Cheli, 620 m, 8 Apr 1957, L.-C. Zang, 1 male; Hekou, Xiaon-
anxi, 200 m, 8 Jun 1956, K.-R. Huang et al., 1 male; 765 km S of Kunluo hwy,
1000 m, 26 Apr 1957, E-J. Pu, 1 male; same loc., but 1050 m, 26 Apr 1957, Q.-
Z. Liang, 1 female; Mangshi City, 900 m, 18 May 1955, V. Popov, 1 male;
Xiaomengyang, 850 m, 2 Apr & 4 May 1957, S.-Y. Wang, 1 male, 1 female;
same loc., but 3 Apr 1957, L.-C. Zang, 2 males; Xishuangbanna, Menga, 1050—
1080 m, 17, 20 Aug 1958, E-J. Pu, 2 males; same loc., but 7 Aug 1958, S.-Y.
Wang, 1 female; Xishuangbanna, Menghun, 750 m, 9 Jun 1958, Y.-R. Zhang, 1
male; Xishuangbanna, Yunjinghong, 850 m, 26 Jun 1958, L.-Y. Zheng, 1 male
(all in IZAS). PROVINCE UNKNOWN: Yen-ping, 13 Sep 1917, Ac. 5148, 1
female (AMNH).
Serida elasmoscelis (Jacobi), NEW COMBINATION
Bisma elasmoscelis Jacobi 1944: 17; Metcalf 1955: 48.
Discussion.—Jacobi (1944) described elasmoscelis in Bisma from one male and
two females collected at Kwangtseh of Fujian Province in southeastern China. I
am here transferring E. elasmoscelis into Serida on the basis of the shape of head,
veins of forewings and hindwings, metatibial spines and the structure of male
genitalia.
_ Distribution.—China (Fujian, Hainan, and Jiangxi Provinces). This is the first
report of S. elasmoscelis in Hainan and Jiangxi Provinces.
148 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Specimens Examined.—CHINA. HAINAN: Ta Han, 23 Jun 1935, J. L. Gressitt, 2 females (NCSU).
SE JIANGXI [KIANGSI]: Hong San, 22, 27, 28, 29, 30 Jun 1936, J. L. Gressitt, 5 males, 3 females
(NCSU; IZAS).
SPECIFIC ADDITIONS
Pyrilla perpusilla (Walker)
Pyrops perpusilla Walker 1851: 269.
Zamila lycoides Walker 1862: 305, pl. 15, fig. 3. [Synonymized by Fennah 1963:
720.]
Pyrilla perpusilla (Walker); Distant 1907: 220.
Pyrilla pusana Distant 1914: 326. [Synonymized by Fennah 1963: 720.]
Discussion.—This species has been reported from India, Sri Lanka, and Thai-
land. It is a major pest of sugarcane and rice in India (Rahman & Nath 1940,
Brar & Bains 1979). I recently discovered 42 examples (17 males and 25 females)
of this species from southern China among unidentified lophopid material in the
IZAS, SIE, and AMNH. This species may become a significant pest of sugarcane
in southern China, especially where the crop is grown for sugar production. This
is also the first report of the genus Pyrilla in China.
Distribution.—China (Fujian, Guangxi, Hainan, Jiangxi, and Yunnan Prov-
inces).
At present, P. perpusilla (Walker) has been found in five southern Chinese
Provinces, from Yunnan Province in the west through to Fujian Province in the
east; however, its apparent absence in several other southern Chinese Provinces
(e.g., Sichuan, Guizhou, Guangdong, and Hunan Provinces) may be an artifact of
lack of collecting. The distribution of this species likely will expand because
sugarcane and rice are cultivated widely across southern China.
Host Plants.—This species not only attacks sugarcane and rice, but also may
attack maize, wheat, barley, oats, guinea grass and other gramineous crops (Rah-
man & Nath 1940, Kalode 1983, O’Brien et al. 1987, Wilson & O’Brien 1987).
Wilson & Claridge (1985) noted that P. perpusilla only attacks rice under favor-
able conditions.
Specimens Examined—CHINA. FUJIAN: Chongan, Chengguan, 250-300 m, 8 Jun 1962, G.-T. Jin
& Y.-M. Lin, 1 female; Chongan, Xingcun, 230-250 m, 1, 4 Jun 1960, G-T. Jin & Y-M. Lin, 6
males, 1 female (all in SIE); same loc., but 210 m, 6 Jun & 13 Sep 1960, Y.-R. Zhang, 2 females;
Chongan, Xingcun, Longdu, 580 m, 27 Jun 1960, Y. Zuo, 1 female; same loc., but 580-640 m, 19
Jun 1960, Y.-R. Zhang, 1 female (all in IZAS); Jianning, 28, 31 May & 5, 6 Jun 1959, G.-T. Jin &
Y.-M. Lin, 5 females; Jianning, Mt. Jinraoshan, 14 Jun 1959, G.-T. Jin & Y.-M. Lin, 1 male (head
missing); Jianyang, Aotou, 970 m, 2 Jul 1960, G.-T. Jin, 1 male; Jianyang, Huangkeng, 350 m, 5 Jul
1960, G.-T, Jin & Y.-M. Lin, 1 female (all in SIE); Jianyang, Huangkeng, Changba, 340-440 m, 22
Aug 1960, Y. Zuo, 1 male; Jianyang, Huangkeng, Tangtou, 310-350 m, 24 Aug 1960, Y. Zuo, 4
males, 3 females (all in IZAS); Ninghua, 19 May 1959, G.-T. Jin & Y.-M. Lin, 1 female (SIE); Wuyi,
30 Aug 1953, 1 male (IZAS); Tongmuguan, 970 m, 3 Jun 1960, G.-T. Jin & Y.-M. Lin, 1 male;
Yongan, Xiyang, 25 Apr 1962, G-T. Jin, 1 female (both in SIE). GUANGXI: Jinxiu, Zhongliang,
Linzucun, 600 m, 21 Nov 1981, G-T. Jin & F-L. Li, 1 female (SIE). HAINAN: Qiongzhong, 6 Mar
1959, G.-T. Jin & Y.-M. Lin, 1 female (SIE); Shuiman, 640 m, 25 May 1960, X.-Z. Zhang, 1 male;
Tongshi, 340 m, 27 May 1960, C.-Q. Li, 1 female; Yinggen, 200 m, 6 Jul 1960, C.-Q. Li, 2 females
(all in IZAS). JIANGXI: Mt. Jiulian, Hualu, 16 Sep 1986, P-Y. Zheng & G.-P. Gan, 1 female (SIE).
YUNNAN: Xishuangbanna, Ganmanta, 580 m, 22 Apr 1957, E-J. Pu, 1 female; Jinping, Changpotou,
700 m, 24 May 1956, K.-C. Huang et al., 1 female (both in IZAS). PROVINCE UNKNOWN. Yen-
ping, 8 Feb 1917, Ac. 5148, 1 male (AMNH).
1996 LIANG: CHINESE LOPHOPIDAE 149
Pitambara dawnana Distant
Pitambara dawnana Distant 1912: 189; 1916: 83, fig. 63.
Discussion.—Distant (1912) described P. dawnana from a single specimen col-
lected in Dawna Hills, Burma and later (1916) redescribed and illustrated the
holotype. No host has been reported. Among undetermined Lophopidae material
(IZAS), I discovered five male examples of P. dawnana Distant from Yunnan
Province, southwestern China; this is the first record of the genus and species in
China.
Distribution.—Lower Myanmar (Burma), China (Yunnan Province).
Specimens Examined—CHINA. YUNNAN: Cheli to Menghai, 720 m, 23 Apr 1957, D.-H. Liu, 1
male; Menghai, Mt. Nannuoshan, 1300 m, 24 Apr 1957, D.-H. Liu, 1 male; Menglong, Banna, Meng-
song, 1600 m, 24 Apr 1958, F-J. Pu, 1 male; Xishuangbanna, Menghun, 1200-1400 m, 19 May 1958,
Y.-R. Zhang, 1 male; Xishuangbanna, Yunjinghong, 850-2040 m, 30 Jun 1958, Y.-R. Zhang, 1 male
(all in IZAS).
CHECK LIST OF CHINESE SPECIES OF LOPHOPIDAE
Fennah (1956) provided a key to five genera and briefly discussed three species
of Lophopidae from China. Chou et al. (1985) recorded four genera and four
species. I here list seven genera and eight species and expect further collecting
will undoubtedly reveal more species.
Elasmoscelis perforata (Walker 1862: 309).—Guangdong and Hainan Provinces,
Taiwan.
Lacusa fuscofasciata (Stal 1854: 248). (= L. yunnanensis Chou & Huang in Chou
et al. 1985: 128, 137, fig. 119a, b. NEW SYNONYMY.—Guizhou, Guangdong,
Hainan, and Yunnan Provinces.
Lophops carinata (Kirby 1891: 140, pl. 5, fig. 9)—Hainan Province, Taiwan.
Paracorethrura iocnemis (Jacobi 1905: 437, pl. 21, figs. 6, 6a)—-Guangxi Prov-
ince.
Pitambara dawnana Distant 1912: 189.—Yunnan Province. New Record.
Pyrilla perpusilla (Walker 1851: 269).—Fujian, Guangxi, Hainan, Jiangxi, and
Yunnan Provinces. New Record.
Serida elasmoscelis (Jacobi 1944: 17), NEW COMBINATION.—Fujian, Hainan,
and Jiangxi Provinces.
Serida latens Walker 1857: 158.—Fujian Province.
ACKNOWLEDGMENT
I express my appreciation to Robert L. Blinn and Lewis L. Deitz (NCSU),
Hong-Xing Li and Hong-Guo Sun (IZAS), Zhi-Yi Luo and Biao Jin (SIE), and
Jing-Ruo Zhou (NWAU) for the loan of specimens, Lois B. O’Brien of the De-
partment of Entomology, Florida A & M University, Tallahassee, Florida, Norman
D. Penny of the Department of Entomology, California Academy of Sciences,
San Francisco, and A. EK Emeljanov and I. M. Kerzhner of the Zoological Institute,
Russian Academy of Sciences, St. Petersburg, Russia, for reviewing the manu-
Script and for their valuable comments in improving the contents of the paper.
The work on which this paper is based was supported by the Theodore Roosevelt
Memorial Fund, Postdoctoral Fellowship Program, American Museum of Natural
History, with Randall T: Schuh.
150 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
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PAN-PACIFIC ENTOMOLOGIST
72(3): 152-159, (1996)
CAVE-ASSOCIATED TROPICAL AMERICAN
NEODITOMYIA (DIPTERA: MYCETOPHILIDAE)
EDWARD I. COHER
Division of Natural Sciences,
Southampton College, Long Island University,
Southampton, New York 11968
Abstract.—With the addition of the adults of two new species of Neoditomyia, as well as the
larva of one, seven species are now known for this cave-associated genus. Flies of this genus
are Neotropical in distribution. The first detailed figures of the male terminalia of N. aeropiscator
(Jackson) and the larval head of N. farri NEW SPECIES are given.
Key Words.—Insecta, Diptera, Mycetophilidae, Caves, Tropical America
The genus Neoditomyia Lane & Sturm, 1958 was proposed for two Neotropical
species of mycetophilids whose larvae spin webs bearing droplets, possibly of
oxalic acid (Buston 1933), to catch and kill prey. The web-spinning habit is well
known for other genera of keroplatine fungus gnats with predaceous larvae, such
as Macrocera (Matile 1975) and Arachnocampa (Hudson 1950). Since that time
three Circumcaribbean species have been described, one each from Honduras,
Trinidad and Cuba. In this study, the adults of two new species are described,
one from Jamaica, West Indies and one from Brazil, the latter known only from
adult males taken in a Malaise trap, but with no data on whether in or outside of
a cave.
In January, 1961, I visited the Institute of Jamaica, West Indies, and Thomas
Farr of the Institute and I climbed the Blue Mts. and visited the Windsor Great
Cave. Entrance into the cave with nothing but flashlights was a bit overly bold.
However, we persisted on penetrating the inner cave, our only mishap being a
‘‘Nantucket sleighride’’ down a large slanting boulder covered with bat guano.
At its bottom, in a large chamber, we were soon enveloped in clouds of cerato-
pogonids that were attracted to the lights. The most striking thing about the upper
chamber walls of the cave was the presence of a covering of webs, sticky to the
touch, with dangling ‘fishing lines’. Slim, worm-like, cream-colored larvae with
a prominent head capsule were resting in the webs. On the lines were tiny drop-
lets. A brief search for pupae and adults of the larva, recognized to be a kero-
platine, was of no avail. Some years later Thomas Farr sent me males and females
of this fly taken in Windsor Great Cave. They proved to be a new species of
Neoditomyia, a known Neotropical cave-dwelling orfeliine genus.
Neoditomyia was originally described as a ditomyiine; it encompassed two new
cave inhabiting species from Colombia with N. colombiana designated as the
generotype. Coher (1963) suggested that they were platyurines and more properly
belonged in Platyura (Taulyrpa) Edwards, 1929. Nicholas (1968) reported the
presence of luminescent dipterous larvae in two caves in Guatemala. The larval
web that he illustrated is similar to that of Neoditomyia, but it would be the only
luminescent species of that genus if that is its affinity. Sturm (1973) reviewed the
biology of the two Colombian species in detail.
Orfelia aeropiscator Jackson, 1974 was described from adults reared from pu-
1996 COHER: CAVE-ASSOCIATED MYCETOPHILIDS 153
pae taken in a Honduran cave. Larvae that he found outdoors living in webs on
the underside of leaves in Costa Rica were ascribed to O. aeropiscator; confir-
mation of this observation is needed. Jackson made valuable observations on the
biology of these larvae, but apparently did not rear any.
Matile (1975) described adults and larvae of a new species N. troglophila from
caves in Cuba, and correctly placed O. aeropiscator in Neoditomyia. In 1982, he
described the male and female of N. spinosa and reviewed the bionomics and
phylogeny of the genus. A slight modification is discussed below.
A report on cave and roadside webs by Cook (1913) is rather limited. The
webs that he and Barber (in Cook 1913) noted could mean that his fly and that
of Nicholas represent a rather widespread species of as yet uncertain affinities.
Neoditomyia Lane & Sturm, 1958
Neoditomyia Lane & Sturm 1958:199.
Platyura (Taulyrpa): Coher 1963:23.
Orfelia (Neoditomyia): Matile 1972:113.
Neoditomyia: Matile 1975:369.
Type Species.—N. colombiana Lane & Sturm, 1958:199 (original designation).
Description—Head: antenna compressed, 2+ 14 segmented, length subequal to mesonotum, scape
with apical ring of short setae; flagellar segments broader than long, penultimate segment nearly
square, apical segment longer than wide, apex rounded; palpus with 4 segments, basal segment greatly
reduced, segment 2 slightly inflated, segments 2 and 3 strongly setiferous, length of apical segment
equal to combined length of segments 2 and 3; posterior clypeus setifereous, frons bare; vertex and
occiput with abundant, short setae, occipital area with shortened seta ventrally, a tuft of long setae at
lower posterior corner of compound eye; ocellar prominence black with 2 large ocelli separated by
less than their diameter, separated from eye margin by more than ocellar diameter. Thorax: with patch
of precoxal setae; mesonotum with variable pattern, heavily clothed with short, dark setae and longer
lateral setae, either with short posteromedian stripe that continues onto scutellum, or with distinct V-
shaped stripe with apex at mid-posterior margin continued onto scutellum, arms of “V’ broadly con-
nected to a wide, heavily setose anterolateral area, a narrow median stripe also present or appearing
as broad median stripe that is wider anteriorly; humeri lightly pigmented; area of median pattern
appearing somewhat raised from remainder of the mesonotum; anterior and posterior margin of both
spiracles bearing row of strong, dark setae; anterior and posterior pronotum setose; proepisternum
with patch of setae; anterodorsal anepisternite with wide, linear patch of strong, dark setae; katepi-
sternite at an angle to anepisternite; mesopleuron bare; pleurotergite heavily setose; scutellum and
median postnotum densely setose; metepimeron bare. Wing 6.0 mm long; costa ending well beyond
apex of R,, Sc short, ending free; Rs, Sc, M, Cu and their branches bare, M;3,, not quite reaching
wing margin, Cu, reaching or not; halter with knob setose, base dark, widened apically with bare
membranous area occupying most of apex. Legs: forecoxa with heavy anterior setae, group of 2—7
large posterolateral apical setae; femora setose; tibiae and tarsi with setae in rows; tibial spur formula
1-1-1, ratio of tibial spurs 1/2/4, apex of midtibia with an internal comb, apex of hind tibia expanded,
combs on both internal and external surfaces. Abdomen: unicolorous, heavily setose. Terminalia: male;
all dististyles with a median internal compressed process (= median plate).
Larva.—Apneustic. Head with typical elongate, forward-projecting, subtriangular maxillae of ker-
oplatine larvae (Fig. 1); mandibles subrectangular with apical edge toothed in a roughly double row
(Fig. 1); stemma large, below posterior antenna (Fig. 1); anterior head capsule with epicranial plates
separated by membranous area; V-shaped submentum heavily sclerotized and pigmented, connected
to premental sclerite; with scimitar-shaped submaxillary sclerites; abdomen with 6 pigmented creeping
welts, widest medially, first and last as single row, others in double row (Fig. 1), all welts under high
magnification appearing as rows of fine hairs; terminally with 2 soft lateral papilla as in Macrocera
(Mansbridge 1933).
154 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Neoditomyia Larva
CREE
HEEL
welt mentum-submentum
maxilla
mandible
head capsule-ventral
Figure 1. Neoditomyia larvae. Maxilla, welt, mentum—submentum, mandible, ventral view of
head capsule.
Neoditomyia farri Coher, NEW SPECIES
Types.—Holotype male: JAMAICA. TRELAWNY: Windsor Cave, 25 Aug
1955, T. H. Farr; deposited: California Academy of Sciences, San Francisco. Al-
lotype: With the same data as the holotype; deposited: California Academy of
Sciences, San Francisco. Paratypes: Same date as holotype, 2 males deposited:
Institute of Jamaica and Coher collection; same data as holotype, but 14 May
1956, T. H. Farr (deposited: Institute of Jamaica), females; Dromilly Cave, 4 Nov
1992, reared from pupa collected 24 Oct 1992, I. Stringer (deposited: Institute of
Jamaica). 17 Jan 1961, E. I. Coher and T. H. Farr 1 larva (deposited: California
Academy of Sciences).
Male.—Head: antenna with scape and pedicel slightly yellow, flagellar segments brown or yellow-
brown; palpus yellow; clypeus and frons slightly yellow; vertex and occiput appearing brown due to
presence of abundant, short setae, occipital area appearing slightly yellow laterally; ocellar prominence
black, attenuated posteriorly; eyes sparsely setiferous, narrowed dorsally above level of antennal base.
Thorax: mesonotum (Windsor form) with variable pattern, heavily clothed with short, dark setae and
longer lateral setae, either yellow-brown with short posteromedian stripe that continues onto scutellum,
or with distinct pattern of V-shaped brown stripe with apex at mid-posterior margin continued onto
scutellum and arms of *‘V’ broadly connected to wide, heavily setose anterolateral brown area, narrow
median brown stripe also present or appearing as broad median brown stripe that is wider anteriorly;
humeri lightly pigmented; area of median pattern appearing somewhat raised from remainder of meso-
notum. Alternatively mesonotum (Dromilly form) lighter with distinct pattern of V-shaped brown stripe
with apex at mid-posterior margin continued onto scutellum, apical arms of ‘V’ with distinct broad
patch of setae; pleura nearly yellow. Wing 5.0 mm long; suffused, darker apically in cell Rs, slightly
lighter in anal cells; R, slightly sinuous or not. Legs: coxae and tibiae yellow, forecoxa with heavy
1996 COHER: CAVE-ASSOCIATED MYCETOPHILIDS 155
anterior setae and a group of 2—7 large posterolateral apical setae; tibiae and tarsi brown. Abdomen:
yellow-brown. Terminalia: (Fig. 2). Setation less (Dromilly male) to strongly developed (Windsor
males).
Female.—Similar to male; mesonotum (Windsor form) with pattern of broad median stripe that is
wider anteriorly or may be obsolete along its middle so that there are pair of brown spots anteriorly
and single median one posteriorly. Alternatively, mesonotum (Dromilly female) similar to Dromilly
male; wing 5.0—6.5 mm long.
Larva.—(Fig. 1). With characteristics given for the genus.
Diagnosis.—The males of N. farri, and its sibling N. spinosa Matile, are easily
differentiated on the basis of the shape of the dististyle, which is strongly trian-
gulate in N. spinosa and bluntly triangulate in N. farri. The setal tuft at the base
of the dististyle of N. spinosa is attenuated basally and is broad in N. farri.
Discussion.—The flies from Dromilly Cave in Jamaica were furnished by I. A.
N. Stringer. He is involved in studies of the bionomics of this fly, which he has
succeeded in rearing. This is undoubtedly the species noted by Matile (1982) as
having been collected by A. S. Peck (1975); I have not been able to determine
where those specimens have been deposited. A comparative study of the anatomy
and bionomics of the larvae of these apparently troglobitic sibling species is
needed.
Entomology.—This species is named for Thomas H. Farr whose contribution
to this study was invaluable.
Material Examined.—See types.
Neoditomyia para Coher, NEW SPECIES
Types.—Holotype male: BRAZIL. PARA: Santo Antonio de Taua, April 1980,
M. Boulard, Malaise trap; deposited: Museum National d’ Histoire naturelle, Paris.
Paratypes: Males (4). Same data as the holotype but, October 1979 (1 male),
February 1980 (1 male), March 1980 (2 males), M. Boulard, all taken in a Malaise
trap (1 deposited: Museum National d’ Histoire naturelle, Paris; others retained by
Coher).
Male.—Head: antenna with scape and pedicel slightly yellow; flagellar segments brown or yellow-
brown, apical 5 segments variably darkened or not; palpus yellow or yellow-brown; clypeus and frons
light yellow-brown; vertex and occiput appearing light brown due to presence of abundant, short setae,
occipital area light yellow-brown laterally. Thorax: mesonotum heavily clothed with short, dark setae,
longer lateral setae, nearly yellow-brown with distinct V-shaped brown stripe with apex at mid pos-
terior margin, arms of “V’ broadly connected to wide, heavily setose anterolateral brown area, narrow
median brown stripe of variable length present; pleura light yellow-brown. Wing 4.5—5.0 mm long;
entirely suffused, or cells R;, Ry, R; and M, darker (1979 paratype); costa ending well beyond apex
of R;, Sc short, Sc, M and its branches and Cu, bare, M;,, not reaching wing margin, Cu, reaching
or not; halter with brown stem, knob setose, slightly white, base dark. Legs: coxae and femora yellow,
forecoxa with heavy anterior setae and group of 5-7 large posterolateral apical setae; tibiae, tarsi
brown, setae arranged in rows. Abdomen: yellow-brown. Terminalia: (Fig. 2).
Female and Larva.—Not known.
Diagnosis.—N. para is most closely related to N. farri. The Brazilian species
has a distinctive bifid median internal compressed process.
Discussion.—The presence of N. para in Brazil extends the range of the Neo-
ditomyia from several Caribbean Islands, Central America and northwestern South
America to the eastern Neotropical Region. Roughly the range of the genus is
now between 16° N and 4° S latitude. This will correct data presented by Matile
156 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Neoditomyia
aeropiscator
para
tr Hise
DAA tara ee
it Hattie
HARA
i} 3
Wie
farri
Figure 2. Terminalia. N. farri NEW SPECIES, N. para NEW SPECIES, N. aeropiscator. Ilustra-
tions are drawn to scale, with dorsal aspect with tergal portion to right.
1996 COHER: CAVE-ASSOCIATED MYCETOPHILIDS 157
(1982) because my examination of the species reported by him from southern
Brazil as Neoditomyia is not that, but actually represents a new genus. There are
no data on the collection labels to indicate whether the adults were taken in or
near a cave.
Neoditomyia aeropiscator (Jackson), 1974
Orfelia aeropiscator Jackson 1974: 240.
Neoditomyia aeropiscator: Matile 1975: 369.
Neoditomyia aeropiscator: Matile 1982: 229.
Remarks.—British Honduras, Cayo District. The original drawings of the male
terminalia and larva of this species were not in detail; the male terminalia are re-
drawn here (Fig. 2). There are no further records of this species. Although Pa-
pavero (1978) indicated that he was erecting a new combination for this species,
it had already been done by Matile (1975).
Neoditomyia andina Lane & Sturm, 1958
Neoditomyia andina Lane & Sturm 1958: 201.
Platyura (Taulyrpa): Coher 1963: 23.
Neoditomyia andina: Sturm 1973: 61.
Neoditomyia andina: Matile 1982: 229.
Remarks.—Colombia, Sierra de la Macarena, 400 to 500 m. No further collec-
tion data. The larva and pupa and their bionomics were well described in the
original study and in further studies by Sturm (1973).
Neoditomyia colombiana Lane & Sturm, 1958
Neoditomyia colombiana Lane & Sturm 1958: 204.
Platyura (Taulyrpa): Coher 1963: 23.
Neoditomyia colombiana: Sturm 1973: 61.
Neoditomyia colombiana: Matile 1982: 229.
Remarks.—Colombia, Resina, 1600 to 2000 m altitude. No further collection
data. The larva and pupa are described by Lane & Sturm (1958) with amplification
by Sturm (1973). I have borrowed the holotype of N. colombiana. The specimen
is intact, therefore the original drawings must have been made from a paratype.
Neoditomyia spinosa Matile, 1982
Neoditomyia spinosa Matile 1982: 205, 229.
Remarks.—Dominica, in several locales, no further collection data. Only the
holotype male was taken in a cave, the allotype and a paratype male are not noted
as being found in caves. The larva and pupa are unknown.
Neoditomyia troglophila Matile, 1975
Neoditomyia troglophila Matile 1975: 369.
Neoditomyia troglophila: Matile 1982: 229.
Remarks.—Cuba, but no further collection data. Described from both sexes and
larvae taken in seven caves.
158 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
BIONOMICS
The presence of eyes in both the adult and larva and of pigmentation in adult
Neoditomyia indicates that cave habitation is probably not a geologically ancient
development for the genus. The vicariant distribution in caves supports Matile’s
(1982) contention that the cave habitat resulted after loss of a more widespread
habitat. Although I consider that several species may be troglobites, the retention
of eyes and pigment may indicate that there are extra-cave populations that in-
terbreed with the cave-inhabiting forms and thus maintain anatomical and color-
ation characteristics expected from populations outside the cave.
The terminalia of male Neoditomyia vary around modifications of the shape of
the dististyle and its median plate, the shape of the setal tuft at the base of the
dististyle and the shape of the central gonostyle. Except for genitalia, I can discern
no sexual dimorphism of any species. Adult coloration characteristics await a
larger sampling before they can be considered to be useful. Larval differences
apparently center around the shape of the mentum-submentum. Comparison of
the structure of the larval webs would be of value, but cannot be evaluated based
on data that is available.
Larval and/or pupal collections indicate that N. aeropiscator, N. farri, N. spi-
nosa and N. troglophila may be troglobitic, but N. andina and N. colombiana are
troglophilic at best. The larvae of N. andina, and N. colombiana were found in
humid conditions on rocks or hanging from vegetation (leaves, fallen tree trunks).
The immatures of N. para are unknown and the exact provenance of the adults
is not known.
As yet there is no detailed anatomical description of N. aeropiscator larvae.
Larvae from Costa Rica ascribed to N. aeropiscator by Jackson (1974) have not
been reared, and because adults and larvae from Honduras were taken only in
caves and the larvae from Costa Rica were found outside on foliage, their rela-
tionship to N. aeropiscator needs further examination.
The larval web of Neoditomyia farri is similar to that described by Lane &
Sturm (1958), Sturm (1973) and Jackson (1974). It provides a larval habitat and
functions with its sticky droplets to capture or kill prey Buston (1933). In Jamaica,
I found numerous ceratopogonids entangled on the ‘fishing lines’. These flies,
easily captured in large numbers, have been identified by W. Wirth as a yet un-
described species of Dasyhelea (Diptera: Ceratopogonidae). They have been de-
posited in the collection of the California Academy of Sciences. In addition, a
single small empid was taken.
In Windsor Great Cave, webs on the wall surface appeared as threads of silk
with larvae apparently enclosed in their own mucus and stretched out on hori-
zontal lines. The long, slim larvae were able to slide back and forth on these
lines. It could not be determined at that time whether the larvae could glide
backwards or whether they performed a ‘U-turn’ to reverse direction. Sturm
(1973) reported that they can perform either function. Other orfeliines such as
Platyura fultoni move in their webs in a similar manner (Fulton 1941). Vertical
fishing lines were noted to be roughly up to 12 cm long with droplets of a sticky
substance. The distribution of these droplets was not noted. A sample of the
larvae, which occurred isolated from each other, was taken. The form of the web
1996 COHER: CAVE-ASSOCIATED MYCETOPHILIDS 159
almost certainly seems to have evolved in caves and rock shelters where air
turbulence would not interfere with development and dessication.
Unfortunately, Lane & Sturm (1958) give a dorsal view of the head capsule of
N. colombiana and a ventral view of the head capsule of N. andina, so that they
cannot be fully compared. The submentum in N. farri is labelled hypopharynx
by Lane & Sturm (1958). I have not been able to examine larvae other than those
of N. farri.
ACKNOWLEDGMENT
I thank Thomas Farr (Institute of Jamaica) for helping to obtain material of the
species dedicated to him; Ian Stringer (Massey University, New Zealand), who
kindly allowed me to study his flies from the Dromilly Cave in Jamaica; Loic
Matile, who has allowed me to include a new Brazilian Neoditomyia sent to him
for study; R. J. Gagné (U.S. National Museum of Natural History, Washington,
D.C.), who arranged a loan of N. aeropiscator paratypes gave valuable recom-
mendations on the development of this study; and Annette Vollers, for her extreme
patience and good humor in joining me to develop all the rough pencil and final
inked drawings.
LITERATURE CITED
Bertrem, J. G. 1928. Monographie der Indo-australischen Scoliidae (Hym. Acul.) mit zoogeogra-
phischen Betrachtungen. Treubia 9 (Suppl.): 3.
Buston, H. W. 1933. Note on the chemical nature of the fluid from the webs of larvae of Platyura
and Ceroplatus. Trans. R. Soc. London, 81: 90-92.
Coher, E. I. 1963. Asian Macrocera Meigen, 1803 (Diptera: Mycetophilidae), with some remarks on
the status of the genus and related genera. Bull. Brooklyn Entomol. Soc., 58: 23-36.
Cook, O. E 1913. Web-spinning fly larvae in Guatemalan caves. J. Washington Acad. Sci., 3: 190-—
193.
Fulton, B. B. 1941. A luminous fly larva with spider traits. Ann. Entomol. Soc. Am., 34: 289-302.
Hudson, G. V. 1950. The natural history of the New Zealand glow-worm. pp. 15-37. Fragments of
New Zealand entomology. Ferguson and Osborn Ltd., Wellington, New Zealand.
Jackson, J. E 1974. Goldschmidt’s Dilemma resolved: notes on the larval behavior of a new Neo-
tropical web-spinning mycetophilid (Diptera). American Midland Nat., 92: 240-245, figs. 1-4.
Lane, J. & H. Sturm. 1958. A new genus of “Ditomyiinae’’. Description of two new species with
bionomical notes (Diptera, Mycetophilidae). Rev. Brasil. Biol., 18: 199-207.
Mansbridge, G. H. 1933. On the biology of some Ceroplatinae and Macrocerinae (Diptera, Myceto-
philidae). Trans. R. Entomol. Soc. London, 80: 75-90.
Matile, L. 1972. Keroplatinae des lles del’Océan Indien Occidental (Diptera, Mycetophilidae). Cah.
ORSTOM, ser. Biol. no. 16: 105-123.
Matile, L. 1975. Un Keroplatinae cavernicole nouveau de Cuba (Diptera: Mycetophilidae). Jn T.
Orghidau et al. Resultats des expéditions biospéologiques cubana-roumaines a Cuba. Acad.
Rep. Soc. Roum. Bucharest: 369-371.
Matile, L. 1982. Systématique, phylogénie et biogéographie des Diptéres Keroplatidae des Petites
Antilles et de Trinidad. Bull. Mus. natn. Hist. nat., Paris. 4 sér., sect. A, 4(1—2): 189-235.
Nicholas, G. 1968. Fauna in Mayan caves. Explorers J., 46: 168-171.
Papavero, N. 1978. A catalogue of the Diptera of the Americas, south of the United States. Family
Keroplatidae, 19C: 1-22. Sao Paulo, Brazil.
Peck, S. B. 1975. The invertebrate fauna of Tropical American caves, Part III: Jamaica, an introduc-
tion. J. Speleol., 7: 303-326.
Sturm, H. 1973. Fanggespinste und Verhalten der Larven von Neoditomyia andina and N. colombiana
Lane (Diptera, Mycetophilidae). Zool. Anzeiger, 191: 61-85, figs. 1-13.
PAN-PACIFIC ENTOMOLOGIST
72(3): 160-161, (1996)
Scientific Note
A RANGE EXTENSION OF HOMALODISCA COAGULATA
(SAY) (HEMIPTERA: CLYPEORRHYNCHA:
CICADELLIDAE) TO SOUTHERN CALIFORNIA
Although there are 13 species of Homalodisca in the western hemisphere
(Young, D. A. 1968. U.S. National Museum Bull., 261), there are only four in
the U.S.: H. coagulata (Say) [eastern, especially southeastern, U.S.], H. lacerta
(Fowler) [Arizona, California], H. insolita (Walker) [southeastern U.S.], and H.
elongata Ball [Texas, Arizona] (Young, D.A. 1958. Bull. Brooklyn Entomol. Soc.,
53: 7-13). Of these, H. coagulata has been previously referred to under the syn-
onym H. triquetra Fabr. by many American authors (Young 1958), and H. lacerta
similarly has been referred to under the synonym H. liturata Ball (Nielson, M.W.
1968. USDA Tech. Bull., 1382); see Nielson (1968) or Young (1968) for the
synonymies of these species.
Homalodisca coagulata has become established recently in southern California,
making it the second Homalodisca sp., along with the native H. lacerta, to occur
in the state. Specimens of H. coagulata have been taken in Ventura, Los Angeles,
Orange, Riverside and San Bernardino Counties, on Citrus, Pinus, Eucalyptus,
Platanus, Prunus, and Ficus with collections dating back to 1990. We suspect
that this leafhopper probably entered California in nursery stock, as eggs, which
are difficult to detect but are frequently intercepted during agricultural quarantine
inspections.
Homalodisca coagulata is a vector of phony peach disease (Turner W. E & H.
N. Pollard. 1955. J. Econ. Entomol., 48: 771-772; Young 1958; Nielson 1968)
and Pierce’s disease of grape (Nielson 1968) in the southern U.S. It feeds broadly,
and has been reported from 73 plant species in 35 families (Turner W. EK & H.
N. Pollard. 1959. USDA, Tech. Bull., 57; Nielson 1968). In southern California,
new circumstances suggest that it may be implicated as a vector of a bacterium
(MLO) causing dieback to Oleanders planted along freeways (A. Purcell, personal
communication). Eggs are laid in clusters in the lower epidermis of leaves of,
usually herbaceous, plants (Turner & Pollard 1959, Nielson 1968). First and sec-
ond instar nymphs, which do not survive well on woody plants, prefer herbaceous
hosts, especially corn, cotton, cowpeas, hollyhock, okra, lambsquarter, and sun-
flower (Turner & Pollard 1959, Nielson 1968). After summer hosts become se-
nescent, older nymphs and adults move to the stems and twigs of woody plants,
and feed preferentially on ash, crapemyrtle, oak, peach, and silktree (Turner &
Pollard 1959, Nielson 1968). Adults usually feed solitarily, but can sometimes
congregate in large numbers on single plants (Turner & Pollard 1959, Nielson
1968). In southern California, they have been observed to line up along terminal
twigs of leafless, dormant peaches (N. Nisson, personal communication). In Geor-
gia, overwintering adults on oak have been known to drop to the ground during
cold (freezing) nights and return to the tree when temperatures rose during the
day (Pollard, H. N. & G. H. Kaloostian. 1961. J. Econ. Entomol., 54: 810-811).
Young (1958) provides a key to the Homalodisca of the U.S. Homalodisca
1996 SCIENTIFIC NOTE 161
Table 1. Diagnostic traits for Homalodisca sp. in California.’
Character H. coagulata H. lacerta
Male size 10.8—13.0 mm 8.5-10.2 mm
Female size> 12.3-13.8 mm 10.0-11.2 mm
Head markings? discontinuous continuous
white/creme white/creme
flecks sinuous lines
Aedeagal processes? 2 pair® 1 pair?
well developed greatly reduced
4U.S. species with transparent anterior area of forewing.
> Young (1958: couplet 2).
¢ Young (1958: fig. 1 c—g).
4-Young (1958: fig. 2 a-g).
coagulata and H. lacerta, as the only California Homalodisca, together, separate
from H. insolita and H. elongata, by having an anterior area on their forewing
that is transparent (vs. forewing completely opaque) (Young 1958: key couplet
1). Homalodisca coagulata can be separated from H. lacerta by its slightly larger
size, the development of the markings on its cephalic dorsum, and the number
and development of aedeagal processes (Table 1).
Acknowled gment.—Jerry Davidson, Nick Nisson, Rosser Garrison and Eldon
Reeves (Departments of Agriculture in Santa Barbara, Orange, Los Angeles and
Riverside Counties, respectively) were instrumental in delimiting the occurrence
and distribution in California. Information on possible involvement of H. co-
agulata as a vector associated with Oleander dieback was provided by A. H.
Purcell (U.C. Berkeley), who is researching the problem.
Records—CALIFORNIA. LOS ANGELES Co.: Rosemead, 12 Sep 1995, D. Papilli, Ficus benja-
mana. ORANGE Co.: Irvine, 19 Jan 1990, S. Quintana, Citrus sp., Pinus sp. RIVERSIDE Co.: Corona,
1 Sep 1994, E. Reeves. SAN BERNARDINO Co.: Fontana, 28 Feb 1994, S. Langford. VENTURA Co.:
Ventura, 7 Mar 1994, P. Phillips, Eucalyptus sp.
John T. Sorensen and Raymond J. Gill, Plant Pest Diagnostics Center, Cali-
fornia Dept. of Food & Agriculture, 3294 Meadowview Road, Sacramento, Cal-
ifornia 95832-1448.
PAN-PACIFIC ENTOMOLOGIST
72(3): 162-163, (1996)
Scientific Note
NEW GENERIC ASSIGNMENT, NEW SYNONYMY, AND
NEW DISTRIBUTION RECORDS FOR THE
NEOPTROPICAL GENUS ICTERALARIA RAZOWSKI
(LEPIDOPTERA: TORTRICIDAE)
Razowski (1991. Misc. Zool., 14: 105-114) proposed the genus Icteralaria
(Tortricidae: Tortricinae: Euliini) to accomodate two new species, J. idiochroma
Razowski and I. ichnobursa Razowski. The description of J. idiochroma was
based on a single male from Costa Rica; that of 1. ichnobursa on a single female
from Colombia. Through the examination of material in the collections of the
University of California, Berkeley (UCB), United States National Museum of
Natural History, Washington, D.C. (USNM), Instituto Nacional de Biodiversidad,
San Jose, Costa Rica (INBio), Los Angeles County Museum of Natural History
(LACM), and The Natural History Museum, London (BMNH), I discovered ad-
ditional representatives of Icteralaria idiochroma, including both sexes. This pa-
per provides additional data on the morphology and distribution of Jcteralaria,
rectifies a spelling error in the original description, and proposes a new synonymy
and new combination in the genus.
Icteralaria belongs to a group of genera within the Euliini characterized by a
long, slender, strongly curved uncus, long-scaled socii, and a transtilla that bears
spines or teeth (e.g., Paraptila). These male genitalic characters typically are
associated with Sparganothini; assignment of these genera to Euliini is based on
the possession of a foreleg hairpencil in the male (Brown, J. W. 1990. Entomol.
News, 101: 109-116) and/or other plesiomorphic characteristics of the male and
female genitalia, such as the distally joined gnathos arms in the male (Brown, J.
& J. Powell 1991. Univ. Calif. Publ. Entomol., 111: 1—87).
In his description of Icteralaria, Razowski (1991) designated Icteralaria
‘““diochroma”’ as the type species. However, this is certainly a misspelling of the
species name idiochroma, which is used in three other places in the paper, in-
cluding the abstract.
Icteralaria idiochroma is a small moth with a narrow, yellow-gold forewing
that bears a broad, rhomboidal, gray-brown patch extending from the costa to the
hind margin across the middle of the wing. The male genitalia are essentially as
described and illustrated by Razowski (1991). The female genitalia are charac-
terized by a simple sterigma; narrow papillae anales; long apophyses; a slender
ductus bursa; a broad, round-ovoid corpus bursa with a large, curved signum; and
a frail membranous accessory bursa that originates from near the middle of the
corpus bursa. They are similar to those illustrated by Razowski (1991) for I.
ichnobursa.
Icteralaria idiochroma appears to be restricted to Central America; it has been
recorded from Honduras and several localities in Costa Rica. Adults have been
collected in May, July, August, and September.
Specimens Examined——COSTA RICA. ALAJUELA PROVINCE: 10.4 rd km (8 rd mi) N of Vera
1996 SCIENTIFIC NOTE 163
Blanca, Volcan Poas, 1500 m, holotype male, 26 May 1985 (J. Powell, UCB); Mount Poas, no date,
1 male (USNM). CARTAGO PROVINCE: Orosi, no date, 1 male (Coll. Wm. Schaus, USNM). CAR-
TEGENA PROVINCE: Ref. Fauna Silv. Tapanti, 1250 m, Aug 1991, 2 males, 1 female (G. Mora,
INBio). GUANACASTE PROVINCE: Est. Pitilla, 9 km S of Santa Cecilia, 700 m, Sep 1990, 1 male
(C. Moraga & P. Rice, INBio), 4-14 Sep 1991, 1 male (A. Guadamuz, INBio). SAN JOSE PROVINCE:
Estac. Carrillo, Parque Nacional Braulio Carillo, 600 m, 28—30 Jul 1990, at lights, 1 female (J. Powell,
UCB). HONDURAS: DEPT. YORO: 45 km S of Tela, 800 m, 25—27 May 1978, 1 male (E. Giesbert,
LACM).
Icteralaria ichnobursa was described from a single female that is superficially
identical to the holotype male of Cnephasia incusa Meyrick, 1917. The type
locality of I. ichnobursa is 4 km NW of San Antonio, Colombia; that of C. incusa
is La Crumbre, Colombia. Based on the similarity in facies and the geographic
proximity of the type localities, 1. ichnobursa, NEW SYNONYM, is synonymized
with I. incusa, NEW COMBINATION. The species incusa was transferred to
Eulia by Clarke (1958, Cat. Microlepid. Brit. Mus. Descr. Meyrick, Volume 3:
131), who illustrated the adult and male genitalia; the female genitalia are illus-
trated (as ichnobursa) by Razowski (1991).
Specimens Examined——COLOMBIA: La Crumbre, 2130 m (6600"), holotype male (incusa), May
1914 (Parish, BMNH); Villa de Cauca, 4 km NW of San Antonio, 2000 m (6500’), holotype female
(ichnobursa), 19 Sep 1958 (A. H. Miller, UCB).
Adults of /. incusa are superficially indistinguishable from I. idiochroma. The
male genitalia of J. incusa lack the narrow, spine-like sacculus of J. idiochroma,
and have the dense patch of fine setae of the aedeagus much reduced. The female
genitalia of J. incusa are similar to I. idiochroma, but the ductus bursa is broader
and the signum is apparently more extensive.
Acknowled gment.—I thank J. A. Powell, Essig Museum of Entomology, Uni-
versity of California, Berkeley; R. W. Hodges, U.S. Department of Agriculture,
Systematic Entomology Laboratory, % National Museum of Natural History,
Washington, D.C.; E. Phillips R., Instituto Nacional de Biodiversidad, San Jose,
Costa Rica; J. P Donahue, Los Angeles County Museum of Natural History, Los
Angeles, California; and K. Tuck, The Natural History Museum, London, for
allowing me to examine material in their care.
John W. Brown, Department of Entomology, San Diego Natural History Mu-
seum, San Diego, California 92112.
PAN-PACIFIC ENTOMOLOGIST
72(3): 164-167, (1996)
Scientific Note
LEAF AGE PREFERENCE FOR OVIPOSITION BY THE
MONOPHAGOUS WHITEFLY, ALEUROTITHIUS
TIMBERLAKE!I (HOMOPTERA: ALEYRODIDAE)
Virtually all studies of whitefly-host plant interactions have utilized polypha-
gous species of whitefly (Homoptera: Aleyrodidae), primarily because the major
economic pests in the family Aleyrodidae are polyphagous species. Although this
focus on polyphagous species is understandable from an economic perspective, it
may provide a very skewed perspective of the family Aleyrodidae from an evo-
lutionary viewpoint because the majority of whitefly species appear to be mo-
nophagous or oligophagous (Mound & Halsey 1978. Whitefly of the world, Wiley
& Sons, New York). Of the approximately 1000 named species that have host
plant records listed in Mound & Halsey’s (1978) catalogue, approximately 70%
have been recorded from only a single plant family and approximately 62% have
been recorded from only a single plant genus.
Determination of a whitefly’s leaf age preference for oviposition is an important
step in understanding its host selection. Immature whiteflies are sessile except for
the early first instar “‘crawler stage.’’ However, even though whitefly crawlers are
mobile, their mobility is very limited and they rarely disperse from the leaf on
which they hatched (Lloyd 1922. Ann. Appl. Biol., 9: 1-32, Dowell et al. 1978.
J. New York Entomol. Soc., 86: 121-122, Mound & Halsey 1978). Thus, the
Ovipositing female not only determines the host plant individual that her offspring
will have to contend with, but she also determines the particular leaf on which
her offspring will be forced to live (or die). The age of the leaf on which immature
whiteflies occur can be a critical factor in their survival. For example, survival
of newly hatched crawlers of bayberry whitefly, Parabemisia myricae (Kuwana)
was 49% on young leaves in contrast to 0% on mature leaves of lemon, Citrus
limon (L.) (Walker & Aitken 1985. Environ. Entomol., 14: 254-257). Conse-
quently, a female whitefly’s choice of leaf age for oviposition can have a profound
effect on her fitness.
Leaf age selection for oviposition by whiteflies has been noted for only a few
species (Aleurocanthus woglumi Ashby, Aleurothrixus floccosus [Maskell], Aleu-
rotrachelus jelinekii [Frauenfeld], Bemisia argentifolii Bellows & Perring, Be-
misia tabaci [Gennadius], Dialeurodes citri [Ashmead], Parabemisia myricae
[Kuwana], Trialeurodes rara Singh, and Trialeurodes vaporariorum [West-
wood]), and all but A. jelinekii are very polyphagous (Hargreaves 1915. Ann.
Appl. Biol., 1: 303-334, Husain & Trehan 1933. Indian J. Agric. Sci., 3: 701—
753, Avidov 1956. Ktavim, 7: 25—41, Khalifa & El-Khidir 1964. Bull. Soc. En-
tomol. Egypte, 48: 115—129, Mound 1965. Empire Cotton Growing Review, 42:
33—40, Gameel 1974. Rev. Zool. Afr., 88: 784-788, Southwood & Reader 1976.
J. Anim. Ecol., 45: 313-325, Yamada et al. 1979. Bull. Veg. & Ornamental Crop
Res. Sta. Series A, 5: 191-199, Ohnesorge et al. 1980. Z. Ang. Entomol., 90:
226—232, Swirski et al. 1980. Alon Ha-notea, 34: 627—635, Xu et al. 1984. Z.
Ang. Entomol., 97: 305-313, Walker & Aitken 1985. Environ. Entomol., 14: 254—-
1996 SCIENTIFIC NOTE 165
257, Noldus et al. 1985. Z. Ang. Entomol., 100: 494-503, Noldus et al. 1986. J.
Appl. Entomol., 101: 492-507, Southwood et al. 1989. J. Anim. Ecol., 58: 921-—
942, Dowell 1990. Pan-Pacif. Entomol., 66: 212—216, Walker & Zareh 1990.
Entomol. Exp. appl., 56: 31-45, Tonhasca et al. 1994. Environ. Entomol., 23:
949-954). In the present study, the leaf age preference for oviposition by the
monophagous whitefly, Aleurotithius timberlakei Quaintance & Baker was ex-
amined. The known geographic range of A. timberlakei is restricted to California
(USA), and it is known to occur only on plants in the genus Eriodictyon (Hydro-
phyllaceae) (Mound & Halsey 1978).
The site of this study was an area of natural vegetation adjacent to California
Highway 243, about 1.7 km N (following the highway) of the Poppet Flats road
junction. The plants used were E. crassifolium Bentham that were occurring nat-
urally and in abundance at the study site. Feral A. timberlakei adults collected
from the study site were used in the tests. Adult whiteflies that were alighted on
the foliage were collected by capturing them in transparent drinking straws (5
mm inside diameter, ca. 3.5—-5 cm long) which were then sealed with corks at
both ends. One whitefly was captured per straw. Whiteflies were collected in this
manner from both young and old leaves. This collection method is much gentler
than aspiration where the whiteflies collide at great speed with the sides of the
aspirator vial.
Within 2 h of capturing the whitefly adults in drinking straws, the whiteflies
were placed in preference test cages (one whitefly per cage) that provided them
access to the abaxial surface of a young and an old E. crassifolium leaf. Leaves
were classified as young if they were on new apical or lateral shoots that were
clearly separated by a distinct node from the older stem from which they arose.
The new shoots were easy to distinguish from the older stems because they were
densely hirsute, whereas the older stems were not. The new shoots were obviously
recent growth, as they had at their apex either an actively growing apical meri-
stem, Or very young unexpanded leaves, or a new flower cluster. Leaves on the
older stems were classified as old leaves. Young leaves were softer and more
pliable than old leaves. At the time of year that the tests were conducted (late
May-—mid June), the distinction between young and old leaves at this site was
obvious. The preference test cages were transparent plastic cylinders (12 mm
inside diameter, 12.5 mm length) that were secured between two leaves (one
young and one old), with the abaxial surface of one leaf covering one of the open
ends of the cylinder and the abaxial surface of the other leaf covering the other
open end of the cylinder. Thus, whiteflies placed in a cage had access to the
abaxial surface of either leaf. The test cage was held between the two leaves using
a hair clip, and a foam plastic gasket at each end of the cylinder made the seal
between the leaf surfaces and the open ends of the cylinder escape-proof. An
illustration and a more detailed description of the preference test cages were given
by Walker & Zareh (1990). The two leaves that were connected together by a
preference test cage were left intact on the plant and were positioned so that the
plane of each leaf was vertical (i.e., neither leaf was “‘above” or “‘below’’ the
other, thus minimizing geotactic or phototactic cues that the whiteflies might use
in selecting one leaf over the other).
A single whitefly adult was placed in each preference test cage by removing
the corks from both ends of the drinking straw in which the whitefly was origi-
166 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Table 1. Numbers of eggs present on young and old E. crassifolium leaves in areas enclosed by
the preference test cages.
Paired t-test Fisher’s distribution-free test
Mean number of eggs wins?
Leaf age a SiB: N t Prob. (young > old) Prob.®
Young 21.05 + 3.40 37
<O.
Old 035 +012 37 6.10 <0.0001 35/36 0.0002
4 The ratio in the column “wins” is the number of replicates where more eggs were laid on the
young leaf than on the old leaf over the total number of replicates (excluding one replicate where an
equal number of eggs was laid on each leaf) (Hollander & Wolfe 1973).
> Two-tailed probability.
nally captured, placing one end of the straw in the cage’s entry hole (see Walker
& Zareh 1990), and gently blowing through the other end of the straw until the
whitefly entered the cage. The entry hole of the cage then was sealed with a cork.
Preference tests were set up in this manner on three dates, 31 May, 7 Jun, and
13 Jun, 1991. Over five different individual FE. crassifolium plants were used in
the tests. Six to seven days after the preference tests were set up in the field, the
test leaves were excised from the plants and were brought back to the laboratory
for examination with a stereomicroscope. The numbers of whitefly eggs present
within the area enclosed by the preference test cage on each leaf was recorded.
For the purpose of statistical analysis, the young and old leaf connected by the
same preference test cage were considered a pair. The number of eggs present on
the examined areas of the young and old leaves were compared using a paired t-
test and the conservative non-parametric Fisher Distribution-Free Sign Test (Hol-
lander & Wolfe 1973. Nonparametric statistical methods, Wiley & Sons, New
York). The Fisher test compares the observed proportion of replicates where more
eggs were laid on the young versus the old leaf to the expected proportion of
replicates where more eggs were laid on the young leaf if neither leaf age was
preferred (i.e., 0.50). Pairs of leaves where no eggs were laid on either leaf were
excluded prior to analysis.
The results from the three dates were similar; therefore, the data were pooled
over all dates. The results clearly indicate that A. timberlakei has a very strong
Oviposition preference for young leaves over old leaves of E. crassifolium (Table
1). Over 98% of the eggs were laid on the young leaves. The preference for
young leaves occurred regardless of whether the adults in the tests were collected
from young or old leaves.
A review of the literature (cited earlier) on leave age preference for whitefly
Oviposition indicates that all of the other whitefly species that have been studied
(all but one of which are very polyphagous with =14 host plant families recorded
by Mound & Halsey 1978) generally prefer the younger leaves of their host plants,
although this preference can be influenced by the age of the plant, the season, or
other factors (Husain & Trehan 1933, Mound 1965, Ohnesorge et al. 1980). In
some cases, the very young, unexpanded leaves are less preferred than slightly
older, but still young, fully expanded leaves, although in these cases, the young,
fully expanded leaves still are preferred over fully expanded mature leaves (Dow-
ell 1990, Walker & Zareh 1990). Nonetheless, by preferring young leaves, the
1996 SCIENTIFIC NOTE 167
monophagous A. timberlakei has a leaf age preference similar to those of its
polyphagous counterparts.
G. P. Walker, Department of Entomology, University of California, Riverside,
California 92521.
PAN-PACIFIC ENTOMOLOGIST
72(3): 168-170, (1996)
Scientific Note
DESCRIPTION OF THE MALE OF CERANISUS
AMERICENSIS (GIRAULT) (HYMENOPTERA:
EULOPHIDAE)
Members of the genus Ceranisus Walker (Eulophidae, subfamily Entedoninae)
are solitary, internal parasitoids of immature stages of thrips (Thysanoptera). The
Nearctic species of Ceranisus have been recently revised (Triapitsyn, S. V. & D.
H. Headrick. 1995. Trans. Am. Entomol. Soc., 121(4): 227—248). However, this
revision lacks a description of the male of Ceranisus americensis (Girault), a
common parasitoid of flower thrips, Frankliniella spp., in North America. The
importance of C. americensis as a biological control agent against western flower
thrips, Frankliniella occidentalis (Pergande) (Greene, I. & M. Parrella. Green-
house Grower, Dec 1992: 69-72, as Ceranisus sp.), warrants complete taxonomic
knowledge of this parasitoid species.
Ceranisus americensis, originally described as a Thripoctenus (Girault, A. A.
1917. Proc. U.S. Nat. Mus., 53(2213): 445-450), was known only from females
until recently, when Robert Zuparko sent several Ceranisus specimens to us for
a possible species identification. This material, collected in California in. 1964,
contained females of C. americensis along with two males that we consider as
being conspecific. Despite sexual dimorphism in some antennal structures com-
mon among Ceranisus spp. (i.e., a swollen scape and a 3-segmented club in males
and a slender scape and a 2-segmented club in females), both female and male
C. americensis share similar funicular characters: Fl is small and usually lacking
sensilla whereas F2 is much larger than Fl and bears several sensilla (Fig. 1).
Figures 1-3. Ceranisus americensis (Girault), male. 1. Antenna; 2. Forewing; 3. Genitalia.
1996 SCIENTIFIC NOTE 169
ef
4b 6ES Were
Figures 1-3. Continued.
170 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
This combination of characters appears to be unique relative to other known males
of this genus. A description of the male of C. americensis follows.
Male—(n = 2) Length: 0.73—0.81 mm. Color: Body coloration black; antennal flagellum, legs, and
wing venation dark brown or brown. Head: About as wide as mesosoma, smooth, except occiput,
frontovertex and malar space with very fine sculpturing. Antenna (Fig. 1) inserted above level of
middle of eye; radicle small and slightly transverse; scape swollen, about 2.0X as long as wide, with
reticulate sculpturing (Fig. 1); pedicel markedly strigate and relatively narrow, about 2X longer than
wide, as wide as F2; FI slightly elongate or otherwise globular, without sensilla; F2 oval, much larger
than F1, with 2 sensory ridges which extend beyond apex of segment; each of three segments of club
with several large sensory ridges, distal segment with distinct apical spicula. Scape and pedicel sparse-
ly setose, flagellum more densely covered by longer setae. Mesosoma: Forewing (Fig. 2) slightly
narrower than in female. Metasoma: Gaster slightly longer and markedly narrower than mesosoma.
Genitalia as shown in Fig. 3. Length: 0.15—0.16 mm.
Material examined [slide mounted except those indicated] USA. CALIFORNIA. VENTURA CO.:
10 mi N of Ojai, Sespe Gorge, 26 Jun 1964, E. FE Riek and G. I. Stage, 18 females and 2 males
(including 13 females on points); 6 mi N of Ojai, Wheeler Gorge, same date and collectors, 5 females
(including 2 on points) [all in Essig Museum, University of California, Berkeley (CISC)].
Acknowledgment.—We thank Robert L. Zuparko (University of California,
Berkeley) for the loan of material and John T. Huber (Biological Resources Di-
vision, CLBRR, Agriculture Canada) for the initial identification of the specimens
to genus.
Serguei V. Triapitsyn and David H. Headrick, Department of Entomology, Uni-
versity of California, Riverside, California 92521-0314.
Received 1 Dec 1995: Accepted 7 Mar 1996.
PAN-PACIFIC ENTOMOLOGIST
72(3): 171-172, (1996)
Scientific Note
THE FIRST NORTH AMERICAN RECORD FOR THE ANT
PHEIDOLE FERVENS FR. SMITH (HYMENOPTERA:
FORMICIDAE)
Pheidole fervens (Fr. Smith) is a small myrmicine ant whose distribution is:
Japan (southern part of Kyushu, Ryukyus), Sri Lanka, Burma, Singapore, China,
Java, Borneo, Taiwan, and Oceania (Ogata, K. 1982. Kontyu, 50: 189-197.) It is
spottily distributed in the Pacific region, including Hawaii where it has apparently
been spread by human commerce (Westview Press, Reimer, N. et al 1990. Chap.
5 in Applied myrmecology: A world perspective). On 24 Aug 1995, I discovered
several colonies of these ants nesting outdoors in downtown Los Angeles (The
Bradbury Building 304 N. Broadway, northwest corner of Third and Broadway
south to 364 N. Broadway and also around the corner on third street to the
northeast side of Spring Street). On a subsequent return trip to the site on 27 Dec
1995 the ants were also nesting on the northeast side of Broadway from Third
and Broadway to 315 N. Broadway. At these sites the nests were established in
the sidewalk and the pavement in cracks and along the sides of buildings. The
workers forage in columns from nest to nest indicating that they are possibly
polydomous (occupy multiple nests). My observations represent the first intro-
duction of this ant in North America (E.O. Wilson pers. comm.)
From my observations, it appears that P. fervens is primarily a scavenger be-
cause the workers were observed carrying the remains of dead insects and scraps
from human foodstuffs. They probably will prey on any live insects they can
overcome, and they possibly also feed on seeds.
Like our native Pheidole (Gregg, R. E. 1958. J.N.Y. Entomol. Soc., 66: 7—48.)
this species has a dimorphic worker caste; a light brown minor worker 2—2.25
mm in length, and a larger, darker major, or soldier caste about 3.5 mm long that
is more robust with a disproportionately larger and more massive head.
The pest potential of this ant is probably slight, although it appears to be
opportunistic and adaptable to disturbed habitats. I suspect competition from the
two most common ants in the urban areas of the Los Angeles basin, the Argentine
ant, Linepithema humile (Mayr) and the native Southern fire ant Solenopsis xyloni
(McCook), will most likely hinder the spread of P. fervens. I suspect the infes-
tation of this new exotic ant is fairly recent because their populations are only in
a two-block area.
Material examined—USA. CALIFORNIA LOS ANGELES CO.: Downtown
Los Angeles: on 24 Aug 1995 and 27 Dec 1995. M. J. Martinez.
Acknowled gment.—\ thank my wife Charlean for her support; Dr. Rosser W.
Garrison, Los Angeles County Entomologist, for his assistance; my fellow myr-
mecologist Robert J. Hamton for identifying this ant as Pheidole fervens and for
reviewing the manuscript; Roy R. Snelling, Los Angeles Co. Museum of Natural
History and Stefan Cover of Harvard University for confirming the identification
of this ant; and my nephew Eric for typing the manuscript.
172 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(3)
Michael J. Martinez, City of Long Beach, Dept. of Parks, Recreation and Ma-
rine, 2760 Studebaker Road, Long Beach, California 90815.
Received 15 Nov 1995: Accepted 14 Feb 1996.
PAN-PACIFIC ENTOMOLOGIST
72(3): 173, (1996)
Book Review
Emmel, T. C., M. C. Minno, and B. A. Drummond. 1992. Florissant Butterflies:
A Guide to the Fossil and Present-Day Species of Central Colorado. Stanford
University Press. 118 pp. + I-IX.
This book has two sections. The first describes the 12 fossil butterfly species
discovered in the Florissant fossil beds including a full citation of the original
description and speculation on the relationship of the fossil butterfly to extant
species. The second covers the region’s extant butterflies and lists adult size, larval
hosts, overwintering stage, and an evaluation of their commonness in the region.
Nine color plates show pinned specimens of the region’s butterflies.
Fifty-three black and white figures cover habitat types and selected larvae,
pupae, and adult butterflies. A butterfly checklist, glossary, and indices of host
plants and general terms are provided. The butterflies are numbered as they are
presented in the text. The numbers are used on the plates, black and white pictures,
and checklist to provide continuity. There are color photographs of 16 live but-
terflies, four habitats and two fossil butterflies.
The text is easily read but suffers from numerous inconsistencies and poor
editing. There is inconsistent conversion of English measurements into metric
equivalents. Some pages have both, others have some (pgs. 5, 15) and others lack
them completely (pgs. 13, 57, 67, 76, 77, 78, 81). English measurements are used
for temperatures, elevation, etc., but only metric measurements are given for adult
butterfly sizes. On page 5, 80° F is 26.7° C, not the 18° C listed.
There are numerous incorrect text references to color figures on the plates. For
butterfly 23 it should be Figs. 23a—-d not 23a-c; butterfly 26—Fig. 26a—b not Fig.
a—b; butterfly 70—Fig. 70a-c not 70c-; butterfly 97—Fig. 97a not 97a—b; and
butterfly 29 is on Plate III not Plate II.
Plate IX seems to have been added late. The text for butterfly 27 doesn’t refer
to Plate IX Figs. 27a—b although they are much better than the black and white
Fig. 33. There are no text references to the color photographs of live butterflies,
habitats, or fossil butterflies.
The paper quality gives poor reproduction of black and white photographs
which appear dull, flat, and lifeless. Many are so poor that they are useless (Figs.
4, 9, 10,21, 27, 33, 37, 40, 47).
The authors extensively cite sources for most of their information yet conclude
that 100 butterfly species existed in the region 35 million years ago, about the
same number as today, without any indication of how 12 fossil species suddenly
increased eight-fold.
These and other problems, including inconsistent citations, detract from the
book which is a must for anyone interested in fossil butterflies, the Florissant
region, or Colorado butterflies but a marginal reference otherwise.
Robert V. Dowell, California Department of Food and Agriculture, 1220 N
Street, Sacramento, California, 95814.
PAN-PACIFIC ENTOMOLOGIST
72(3): 174-175, (1996)
PACIFIC COAST ENTOMOLOGICAL SOCIETY
NOTES TO THE FINANCIAL STATEMENTS
YEAR ENDED SEPTEMBER 30, 1992
SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES
Accounting Method
Income and expenses are recorded by using the cash basis of accounting.
Note from the Treasurer
The Pan Pacific Entomologist, the journal of the Pacific Coast Entomological
Society, 1s published quarterly. However, due to editorial delays, the issues are
often not published and charged to the Society on schedule. This explains the
abnormal fluctuation in publishing costs seen between FY 90-91 and FY 91-92.
In FY 90-91 six issues were paid for at a cost of $41,561. In FY 91-92 only 3
issues were paid for at a cost of $23,366. Estimated cost for publishing Vol. 68,
Nos. 3 and 4, which should have been published and charged in FY 91-92 but
have been delayed until FY 92-93, will be about $11,000.
Capital Expenditures
Annual capital expenditures of $5,000 or less are charged to expense.
Marketable Securities
American Telephone & Telegraph Co. and Pacific Telesis Group common stocks
are carried at market value. Increases and decreases in value are reflected in
income.
Income Tax
The Society is exempt from Federal Income and California franchise tax.
As Chairman of the Auditing Committee, and in accordance with the Society’s
bylaws, I have reviewed the financial records of the Society but have not made
an audit of them.
During the course of this review, nothing was noted which indicated any material
inaccuracy in the financial statements.
H. Vannoy Davis
Chairman of the Auditing Committee
1996 1991 & 1992 FINANCIAL STATEMENT 175
PACIFIC COAST ENTOMOLOGICAL SOCIETY
STATEMENT OF INCOME, EXPENDITURES AND
CHANGES IN FUND BALANCES
YEARS ENDED SEPTEMBER 30, 1992 AND 1991
2 £19925 ae +1991
Income
Dues and subscriptions .......... 0.0... cece cee cee cence eee aes $ 12,585 § 17,375
Reprints and miscellaneous .......... 0.0.0... cece cece cece eee 10,894 18,159
TEGLES YF TOA SEB, ark Be Re OR Oe Pn da. bole eR ei PP 3,657 5,407
DORIC HGS Fret ek) Merny cn ns Bee ta cn ee tN, RONEN aL Ae Bas Niet 676 722
Increase (Decrease) in value of capital stock:
American Telephone & Telegraph Company ..................... 490 530
Pacific “Telesis Group-< 2 fn. Bock tee dares enh sie cae ened 726 (627)
ictal Teaneck out ee BS ee cL cen oD neal $_ 29,028 $ 41,566
Expenditures
Publication costs—Pan-Pacific Entomologist ....................00. $ 23,366 $ 41,561
Postage, newsletter and miscellaneous expenses .................04- 830 1,462
‘Total -Exnenditures:). &- 994-4. rags ee a, oe rome ee en Poa $ 24,196 $ 43,023
Increase (Decrease) in fund balances .............. 0.0.0 cece eee eee $ 4,832 $ (1,457)
Fund balances October 1, 1991 and 1990 ..............0.. 0... cece 110,544 112,001
Fund balances September 30, 1992 and 1991 ......................... $115,376 $110,544
STATEMENT OF ASSETS
AS OF
SEPTEMBER 30, 1992 AND 1991
1992 1991
Cash in bank
G@ommercial Accousite or... es ied oP hk Pet head | pie ete 2 $ 5,823 $ 5,144
Undeposited dividend checks ........... 0.0... e cece cee ees 183
Certificates of Deposit and Money Fund:
General Fund— Wells Fargo Bank .............. 00... cc cece neces 9,538 9,219
C. P. Alexander Fund—Capital Preservation Fund
Fall Memoir Fund— Wells Fargo Bank ...................0000 00 35,497 34,313
"Fotal-Cash in Danita se tok S UE We neal eBook POR eel gs $100,105 $ 96,489
Capital Stock (at market value)
American Telephone & Telegraph Co., 80 shs. .................005. 3,490 3,000
Pacific Telesis Group, 264 shs.
et ENS etd oo aneys 49,247 47,630
ered cay eae tre ne Meret ne kites eee ee 11,781 11,055
15,271 14,055
$115,376 —$110,544
See accompanying notes to the financial statements
PAN-PACIFIC ENTOMOLOGIST
72(3): 176-177, (1996)
PACIFIC COAST ENTOMOLOGICAL SOCIETY
NOTES TO THE FINANCIAL STATEMENTS
YEAR ENDED SEPTEMBER 30, 1993
SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES
Accounting Method
Income and expenses are recorded by using the cash basis of accounting.
Capital Expenditures
Annual capital expenditures of $5000 or less are charged to expense.
Marketable Securities
American Telephone & Telegraph Co. and Pacific Telesis Group common stocks
are carried at market value. Increases and decreases in value are reflected in
income.
Income Tax
The Society is exempt from Federal Income and California franchise tax.
As Chairman of the Auditing Committee, and in accordance with the Society’s
bylaws, I have reviewed the financial records of the Society but have not made
an audit of them.
During the course of this review, nothing was noted which indicated any material
inaccuracy in the financial statements.
H. Vannoy Davis
Chairman of the Auditing Committee
PACIFIC COAST ENTOMOLOGICAL SOCIETY
STATEMENT OF INCOME, EXPENDITURES AND
CHANGES IN FUND BALANCES
YEARS ENDED SEPTEMBER 30, 1993 AND 1992
__ 1993
Income
ues And =suPscriptrons™ 85. sets.) Sat Bik coh. sha aee siete he es ome $ 14,682
Reprints and miscellaneous ............. 0.0 ccc cece eee eee eee ees 15,322
FTES Ye ND ca RE an, li Ai on Rl tet an Oe Oe 2 Ae 2,308
IDIViGENndS bento a ete gk eet Peas oth Oe ween SNe AE ot yO, Oe 681
Increase (Decrease) in value of capital stock:
American Telephone & Telegraph Company 1,200
Pacifie Telesis Groups <2 4 iain oe ee en i al ee ae a 2,178
otal line onic secs forse SE tlt ee Lee ad OR Oe Ler ae LS $ 36,371
1992
$ 12,585
10,894
3,657
676
490
726
$ 29,028
1996 1992 & 1993 FINANCIAL STATEMENT 177
Expenditures
Publication costs—Pan-Pacific Entomologist ................... eee $ 28,076 $ 23,366
RRCOTIT COSTSSE Ne, sof ok So, Petey Rie RRL Tels Seeks Shoe, eaten ee RE, 2,050
Postage, newsletter and miscellaneous expenses .................05: 2,457 830
Bota xpemditures: 2-26 fjccat cg te gente dees Macias een te pee de mentees De $ 32,583 $ 24,196
Increase (Decrease) in fund balances ............. 0.00. c eee cece ees $ 3,788 $ 4,832
Fund balances October 1, 1991 and 1990 ........... 0.0.0.0... ce eee, 115,376 110,544
Fund balances September 30, 1993 and 1992 ......................... $119,164 $115,376
STATEMENT OF ASSETS
AS OF
SEPTEMBER 30, 1993 AND 1992
1993 1992
Cash in bank
Commercial secounte ese A ie toce ocean oe ee erie eet $ 9,011 $ 5,823
Undeposited dividend checks .............. ccc cece cee cence eee eees
Certificates of Deposit and Money Fund:
General Fund— Wells Fargo Bank ................ 000 ccc ccc eee 4,731 9,538
C. P. Alexander Fund—Capital Preservation Fund ................ 50,426 49,247
Fall Memoir Fund— Wells Fargo Bank .................. 0000000 36,347 35,497
Tetal cash insbank’.. 5... 6.50) -<ds 66 colts Rio pete tes BO eed eh dee hee $100,515 $100,105
Capital Stock (at market value)
American Telephone & Telegraph Co., 80 shs. ..............0.0 0005 4,690 3,490
Pacific Telesis Group, 264 shs. ........ 0... cece cee cece eee cece
13,959 11,781
18,649 S271
$119,164 ($115,376
See accompanying notes to the financial statements
PAN-PACIFIC ENTOMOLOGIST
72(3): 178-179, (1996)
PACIFIC COAST ENTOMOLOGICAL SOCIETY
NOTES TO THE FINANCIAL STATEMENTS
YEAR ENDED SEPTEMBER 30, 1994
SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES
Accounting Method
Income and expenses are recorded by using the cash basis of accounting.
Note from the Treasurer
The Pan Pacific Entomologist, the journal of the Pacific Coast Entomological
Society, is published quarterly. However, due to editorial delays, the issues are
often not published and charged to the Society on schedule. This explains the
abnormal fluctuation in publishing costs.
Capital Expenditures
Annual capital expenditures of $5000 or less are charged to expense.
Marketable Securities
American Telephone & Telegraph Co., Pacific Telesis Group and Air Touch
Communications common stocks are carried at market value. Increases and de-
creases in value are reflected in income. Air Touch Communications was spun
off from Pacific Telesis Group during the year.
Income Tax
The Society is exempt from Federal Income and California franchise tax.
As Chairman of the Accounting and Tax Committee, and in accordance with the
Society’s bylaws, I have reviewed the financial records of the Society but have
not made an audit of them.
During the course of this review, nothing was noted which indicated any material
inaccuracy in the financial statements.
H. Vannoy Davis
Chairman of the Accounting and Tax Committee
1996
PACIFIC COAST ENTOMOLOGICAL SOCIETY
1993 & 1994 FINANCIAL STATEMENT
179
STATEMENT OF INCOME, EXPENDITURES AND
CHANGES IN FUND BALANCES
YEARS ENDED SEPTEMBER 30, 1994 AND 1993
Income
Dues and subscriptions: 0). 0. occa. oe ie vests cele ew tne sw eren eee eee
Reprints and miscellaneous
TNIGCT ESE pees: he ee Ey oe 8 TERE Berk Bl bee he Deas Mk ate oie & eT. BT bea hoe
DIVIG ERS oe aes WR Rees ee ke VSR Ae OT Ah, DO ae
Increase in value of capital stock:
American Telephone & Telegraph Company .....................
Pacific Telesis Group and Air Touch Communications (see note) ...
Total Income
Expenditures
Publication costs—Pan-Pacific Entomologist
FRE PIINE-COStS oe) IE een Se ee Se eT ee Se Sed Oe el
Postage, newsletter and miscellaneous expenses
Total Expenditures
Increase (Decrease) in fund balances .............. 000.0 cece eee
Fund balances October 1, 1993 and 1992 ........... 0... ccc cee
Fund balances September 30, 1994 and 1993
STATEMENT OF ASSETS
AS OF
SEPTEMBER 30, 1994 AND 1993
Cash in bank
Commercial AecOumnt eke ec F Us ts fed Sin she Pere Gg hteine fon Ap Geis She 2
Certificates of Deposit and Money Fund:
General Fund—Wells Fargo Bank .............. 0.0. c cece eee
C. P. Alexander Fund—Capital Preservation Fund
Fall Memoir Fund— Wells Fargo Bank
Total cash in bank
Capital Stock (at market value)
American Telephone & Telegraph Co., 80 shs. ..................005
Pacific Telesis Group, 264 shs. ..........0... ccc cece cece vec eeeveves
Air Touch Communications, 264 shs. ......... 0.0... c cece eee
See accompanying notes to the financial statements
1994 _ 1993
$ 17,504 $§$ 14,682
11,010 15,322
2,604 2,308
681 681
(370) 1,200
1,716
: 2,178
$ 33,145 $ 36,371
$ 19,448 §$ 28,076
2,138 2,050
a5 2,457
$ 22,939 $ 32,583
$ 10,206 $ 3,788
119,164 115,376
$129,370 —_$119,164
1994 1993
$ 15,386 $ 9,011
4,832 4,731
52,015 50,426
37,142 36,347
$109,375 _$100,515
4,320 4,690
8,118 13,959
7,557
19,995 18,649
$129,370 $119,164
PAN-PACIFIC ENTOMOLOGIST
72(3): 180, (1996)
ANNOUNCEMENT
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submitting manuscripts for which these formats are applicable.
Literature Cited. — Format examples are:
Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.
Blackman, R. L., PR A. Brown & V. F Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide
some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. F Dixon & L. Weismann (eds.). Population structure, genetics and
taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB
Academic Publishing, The Hague, The Netherlands.
Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.
Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.
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THE PAN-PACIFIC ENTOMOLOGIST
Volume 72 July 1996 Number 3
Contents
SNOOK, R. R. & T. A. MARKOW—Possible role of nonfertilizing sperm as a nutrient source
for female Drosophila pseudoobscura Frolova (Diptera: Drosophilidae) ____--_-_---- sve!
BERNHARDT, P. & K. WALKER—Observations on the foraging preferences of Leioproctus
(Filiglossa) Rayment (Hymenoptera: Colletidae) in eastern Australia _. 130
MAYER, D. E—Effects of sex ratio and female density on progeny survival of the alfalfa
leatcubter beac ChivyincnopichamyrecacmiMidic ets yes Sete ee ee 138
GRISWOLD, T. L.—A new Microbembex endemic to the Algodones Dunes, California (Hy-
Wile @ vera Rein Seta ae) ess ie ee es 142
LIANG, A.-P—Taxonomic changes in Chinese Lophopidae with a check list of Chinese species
CHOmipinte lar soMal deny; &. hla 80. al Wi kk ee ea We Ee BF 145
COHER, E. I—Cave-associated tropical American Neoditomyia (Diptera: Mycetophilidae) -.. 152
SCIENTIFIC NOTES
SORENSEN, J. T. & R. J. GILL—A range extension of Homalodisca coagulata (Say) (He-
miptera: Clypeorrhyncha: Cicadellidae) to southern California 160
BROWN, J. W.—New generic assignment, new synonymy, and new distribution records for
the neotropical genus /cteralaria Razowski (Lepidoptera: Tortricidae) ___---_--_- 162
WALKER, G. P.—Leaf age preference for oviposition by the monophagous whitefly, Aleuro-
titthus timberlake (Homoptera: Aleyrodidae) oe eee 164
TRIAPITSYN, S. V. & D. H. HEADRICK—Description of the male of Ceranisus americensis
(Giratlt <(rlyimencpierd: Eile pin dae or: al. Enea Be! we Ra iu ed ae ne 168
MARTINEZ, M. J.—The first North American record for the ant Pheidole fervens Fr. Smith
CEL YIN GHGOp ler sr OPIN Cle), Bee S = = etme nce UP Re en ee ed ns aR ae ee 171
BOOK REVIEW
DOWELL, R. V.—Emmel, T. C., M. C. Minno & B. A. Drummond. 1992. Florissant Butter-
flies: A Guide to the Fossil and Present-Day Species of Central Colorado. Stanford
UiniWverpibyer esse IE isis cat ok ony ae Oe ely oss ae Pewee a 173
Pacific Coast Entomological Society, financial statement for 1991, 1992 __ 174
Pacific Coast Entomological Society, financial statement for 1992, 1993 ___ 176
Pacific Coast Entomological Society, financial statement for 1993, 1994
Announcement—publications of the Pacific Coast Entomological Society 180
The
PAN-PACIFIC
ENTOMOLOGIST
Volume 72 October 1996 Number 4
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
(ISSN 0031-0603)
The Pan-Pacific Entomologist
EDITORIAL BOARD
R. V. Dowell, Editor R. M. Bohart
R. L. Penrose, Associate Editor J. T. Doyen
R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.
Julieta E Parinas, Treasurer Warren E. Savary
Published quarterly in January, April, July, and October with Society Proceed-
ings usually appearing in the October issue. All communications regarding non-
receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary;
and financial communications should be addressed to: Julieta EK Parinas, Treasurer;
at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad-
emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.
Application for membership in the Society and changes of address should be
addressed to: William Hamersky, Membership Committee chair, Pacific Coast
Entomological Society, Dept. of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, CA 94118-4599.
Manuscripts, proofs, and all correspondence concerning editorial matters (but
not aspects of publication charges or costs) should be sent to: Dr. Robert V.
Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture,
1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con-
tributors, and volume 66(1): 1—8, January 1990, for more detailed information.
Information on format for taxonomic manuscripts can be found in volume 69(2):
194-198. Refer inquiries for publication charges and costs to the Treasurer.
The annual dues, paid in advance, are $25.00 for regular members of the So-
ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for
institutional subscriptions or sponsoring members. Members of the Society receive
The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes
are available; see 67(1): 80 for current prices. Make checks payable to the Pacific
Coast Entomological Society.
Pacific Coast Entomological Society
OFFICERS FOR 1996
Wojciech J. Pulawski, President Vincent FE Lee, Managing Secretary
Julieta EK Parinas, Treasurer Stanley E. Vaughn, Recording Secretary
THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per
year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate
Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and additional
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This issue mailed 18 September 1996
The Pan-Pacific Entomologist (ISSN 0031-0603)
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PAN-PACIFIC ENTOMOLOGIST
72(4): 181-192, (1996)
NEW CYNIPID WASPS FROM THE SOUTHWESTERN
UNITED STATES (HYMENOPTERA: CYNIPIDAE)
ROBERT J. LYON
2120 Bristow Dr., La Canada, California 91011
Abstract.—Seven new cynipid species are described from Arizona and western Texas: Ceropteres
snellingi, Xanthoteras pungens, X. tuckeri, Antron daileyi, A. franklinensis, A. madera, and An-
dricus flocculentus. Appropriate morphological features of the species are illustrated. Included
is a key to the described United States species of Antron.
Key Words.—Insecta, Cynipidae, inquiline, unisexual female, monothalamous gall, Quercus ar-
izonica, Q. pungens, Q. toumeyi, southwestern U.S. (Arizona and western Texas).
The phytophagous cynipid fauna of the southwestern United States (Arizona,
New Mexico, and western Texas) and adjacent Mexico needs additional study.
Weld (1960) listed 130 species of Cynipidae from this area but in addition illus-
trated 117 galls from which no gall wasps were ever reared, thus there is still
collecting and rearing to be done. No life cycle, i.e., no linking of alternating
unisexual and bisexual generations of any species, has ever been achieved through
experimental rearings in the region. Knowledge of such alternating generations,
which are frequently morphologically very dissimilar, would prove useful in un-
derstanding the taxonomic status of these wasps at both specific and generic lev-
els.
Of particular interest are the cynipids that induce galls on the shrub oak, Quer-
cus pungens Liebmann, which until recently (Lyon 1993) was not known to be
a cynipid host. Since it is a white oak (Subgenus Lepidobalanus) it was found to
be “‘galled’’ by several described cynipids that form galls on other white oaks in
this area. The following are New Host Records for species that form galls on Q.
pungens: Neuroterus howertoni Bassett, Disholcaspis rubens Gillette, Atrusca
brevipennata (Gillette), Callirhytis juvenca Weld, C. frequens (Gillette). The two
new Xanthoteras species described below show some most unusual morphological
variations, particularly in the reduction of the wings and their peculiar venation.
Other undescribed galls were collected during this study, but I was unable to rear
adult wasps.
ANDRICUS FLOCCULENTUS LYON, NEW SPECIES
(Figs. 18, 19)
Holotype female; TEXAS. EL PASO CO.: Franklin Mountains, El Paso, 15
Feb 1969, R. J. Lyon, from galls on Quercus pungens Liebmann; deposited: U.S.
National Museum of Natural History, Washington, D.C. Paratypes: same data as
holotype, except collected 16 Feb—16 Mar 1962, 15 Feb 1969; deposited: U-S.
National Museum of Natural History, Washington, D.C. (3), California Academy
of Sciences, San Francisco, CA (3), Natural History Museum of Los Angeles
County, Los Angeles, CA (3), R. J. Lyon collection (5).
Description.—Female. Black, except legs, antennal segments and ventral spine of metasoma light-
brown. Length: 1.5-1.75 mm (* = 1.6 mm, n = 17). Head transverse in dorsal view, as broad as
mesosoma, occiput concave; gena not broadened or visible behind eye in frontal view; interocular
182 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
area 2X as broad as high; malar space striate, 0.20 eye length (EL); ocellar area, frons and other
parts of head, coriaceous. Antenna 13-segmented, segments 3 and 4 equal in length, terminal segment
2X as long as 12. Scutum coriaceous, marginate, subcircular and only slightly longer than broad, bare;
notauli incomplete. Scutellum round behind, sparsely pubescent; surface entirely rugose or rugose only
posteriorly; foveal pits replaced by a transverse groove with smooth, shining bottom. Propodeal carinae
arcuate, enclosure smooth and shining. Mesopleuron aciculate and protuberant. Legs pubescent; tarsal
claws toothed. Forewing hyaline and pubescent, margins ciliate; veins brown; areolet well-developed,
reaching one-fifth distance to Basal Vein (Rs+M); M, not reaching Basal Vein; Rs, angled; Rs, arcuate;
Radial Cell 3X as long as broad. Metasoma higher than long, three terga visible along dorsal curvature;
terga smooth and shining; tergite 2 with basal pubescent patch; ovipositor sheaths protruding, tip of
Ovipositor curved. Ventral spine sparsely pubescent, length 4X width.
Diagnosis.—Distinguished from other members of the genus by its striate malar
area, completely aciculate mesopleuron, and nearly circular mesoscutum.
Gall.—(Figs. 18, 19).—Clustered, individual cells forming a woolly mass on
the underside of the leaf midrib. Similar galls occur on other white oaks in the
area, but no adults of A. flocculentus were reared from them.
Host.—Quercus pungens
Material Examined.—See types.
CEROPTERES SNELLINGI LYON, NEW SPECIES
Holotype female; TEXAS. El PASO CO.: Franklin Mountains, El Paso, 15 Feb
1969, R. J. Lyon, from galls of Andricus flocculentus Lyon on Quercus pungens;
deposited: U.S. National Museum of Natural History, Washington, D.C. Para-
types: same data as holotype; deposited: U.S. National Museum of Natural His-
tory, Washington, D.C. (3), California Academy of Sciences, San Francisco (3),
Natural History Museum of Los Angeles County, Los Angeles (3), R. J. Lyon
collection (8).
Description.—Female. Head and body red-brown; legs, antennae and ventral spine amber, with
darker infuscations. Length 1.25-1.75 mm (« = 1.47 mm; n = 15). Head transverse in dorsal view,
microcoriaceous; occiput concave; vertex flattened, median ocellar area depressed; interocular area
wider than high; malar space with prominent, divergent striae; gena not broadened behind eye; a
parallel ridge on each side extends between antennal socket and clypeus. Antenna 12-segmented,
segment 3 equal to 4 in length, segment 12 longer than 11. Scutum slightly broader than long,
somewhat truncate anteriorly, microcoriaceous and with numerous white bristles; notauli incomplete,
extending one-half the distance to pronotum; lateral lines faint and median line represented by a short
notch. Mesopleuron smooth, shining, not striate. Scutellum rounded behind, rugose, with long white
bristles; foveal pits small, oval, separated, almost hiden by coarse sculpture in some specimens. Tarsal
claws edentate. Forewing hyaline, pubescent, margins ciliate; veins pale; areolet well-developed, ex-
tending one-fourth distance to Basal Vein; M, not reaching Basal Vein; radial cell closed, about 2X
as long as broad; Rs, arcuate; Rs, almost straight. Metasoma slightly higher than long, petiole ridged
as in Synergus; tergite 2 bristly, saddle-shaped, 0.4 as long as 3 and separated from it by a connate
suture; ovipositor sheaths long, projecting almost vertically above terga. Ventral spine short.
Male—Slightly smaller than female, length 1.0-1.5 mm; color similar. Antenna 14-segmented;
notauli extending three-fourths distance to pronotum; tergite 2 sparsely pubescent.
Diagnosis.—This species can be separated from other members of the genus
by the following combination of characters: small, separated foveal pits; promi-
nent, long, vertically-projecting ovipositor sheaths; incomplete notauli; female an-
tenna 12-segmented.
Host.—This species is an inquiline in the galls of Andricus flocculentus on
Quercus pungens.
Etymology.—This species is named for Roy R. Snelling, Natural History Mu-
1996 LYON: NEW SOUTHWESTERN CYNIPID WASPS 183
Figures 1-6. Xanthoteras tuckeri, Lyon, NEW SPECIES. Figure 1. Metasoma, lateral view show-
ing shape of terga. Figure 2. Head, dorsal view (massive: more than 0.5X as long as broad). Figure
3. Head, frontal view, showing malar furrows. Figure 4. Mesosoma, dorsal view, showing configuration
and unusual wing venation. Figure 5. Monothalamous gall showing position of larval cell. Figure 6.
Leaf of Q. pungens showing gall on margin of upper surface.
seum of Los Angeles County, who has been of great assistance and encourage-
ment in the writing of this paper.
Material Examined.—See types.
XANTHOTERAS TUCKERI LYON, NEW SPECIES
(Figs. 1-6)
Holotype female; TEXAS. EL PASO CO.: Franklin Mountains, El Paso, 15
Feb 1969, R. J. Lyon, from galls on Quercus pungens; deposited: U.S. National
Museum of Natural History, Washington, D.C. Paratypes: same data as holotype,
deposited: U.S. National Museum of Natural History, Washington, D.C. (2), Cal-
184 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
ifornia Academy of Sciences, San Francisco (2), Natural History Museum of Los
Angeles County, Los Angeles (2), R. J. Lyon collection (9).
Description—Unisexual female. Uniformly red-brown, except metasomal segments dark brown
apically. Length 1.2-1.9 mm (x = 1.4 mm; n = 16). Head more than 0.5X as long as broad, massive
in dorsal view, broader than mesosoma; surface granulose; gena not broadened behind eye in frontal
view; ocellar area depressed; malar space almost as long as eye, malar furrow present; interocular
area nearly 2X as broad as high. Antenna 14-segmented, moniliform, segment 3 2X as long as 4,
segment 14 longer than 13. Scutum as wide as long, smooth and shining; notauli complete in some
specimens, replaced by a row of pits in others. Mesopleuron smooth and shining, slightly bulging in
center. Forewing greatly reduced, venation modified from that of typical, phytophagous cynipids (Fig.
4): Sc+R well-developed, heavy; M, heavy, extending toward base of wing; R, claviform; Rs, very
short; M, short, heavy, terminating in a rounded club; other veins, including areolet, absent. Legs
pubescent; tarsal claws dentate. Metasoma longer than high, all terga visible along dorsal curvature;
terga smooth, except punctate last segment; ovipositor sheaths punctate. Ventral spine bristly, 2.5X as
long as broad in lateral view.
Diagnosis.—Separable from other species in this genus by the massive head
and the reduced wing with unique venation described above (Fig. 4).
Gall.—(Figs. 5, 6)—Monothalamous, tiny (2.5 mm diameter), spherical, on
upper leaf surface; larval cell occupying lower portion of gall; appears in late
summer, maturing in late December through January and February.
Host.—Quercus pungens.
Etymology.—Named for Professor John Tucker, University of California, Davis,
who located the stands of Q. pungens used in this study.
Material Examined.—See types.
XANTHOTERAS PUNGENS LYON, NEW SPECIES
(Figs. 7-12)
Holotype female; TEXAS. El PASO CO.: Franklin Mountains, El Paso, 13 Jan
1969, R. J. Lyon, from galls on Quercus pungens deposited: U.S. National Mu-
seum of Natural History, Washington, D.C. Paratypes: same data as holotype
except collected 15-30 Dec 1972; deposited: U.S. National Museum of Natural
History, Washington, D.C. (2), California Academy of Sciences, San Francisco
(2), Natural History Museum of Los Angeles County, Los Angeles (2), R. J. Lyon
collection (9).
Description—Unisexual female. Uniformly red-brown, posterior metasomal terga darker. Length
2.3-2.7 mm (x = 2.5 mm; n = 20). Head transverse, broader than mesosoma; surface coriaceous;
mandibles with prominent teeth; gena not broadened behind eyes in frontal view; malar space less
than .75X eye length, furrow present; interocular area 2X as wide as high; ocelli amber, each ocellus
in surrounding depression. Antenna stoutly moniliform, 13-segmented, segment 3 longer than 4, seg-
ment 13 nearly 2X as long as 12. Pronotal side smooth, lightly striate along borders. Scutum smooth
and polished; notauli visible only posteriorly, but extending forward as a row of punctures and a few
bristles. Scutellum longer than broad, margined and longitudinally ridged; fovae represented by an
arcuate groove. Propodeum with arcuate carinae, enclosure rugose. Mesopleuron smooth and polished,
with striate area below wing. Legs pubescent, claws dentate. Forewing (Fig. 7) short, not quite reaching
tip of mesosoma, pubescent, short-ciliate along posterior margin. Veins thick and heavy; areolet absent;
Radial Cell short; Rs, swollen and curved upward; Cubital Cell 3 with dark, irregular patch extending
into radial cell; slight darkening in Discoidal Cell and along M,, Rs+M and Median Veins. Metasomal
terga smooth and polished, tergum 2 with pubescent patch. Ventral spine slightly more than 2X as
long as thick in lateral view, tapering to blunt point.
Diagnosis.—This species can be separated from the other species in the genus
1996 LYON: NEW SOUTHWESTERN CYNIPID WASPS 185
Figures 7-12. Xanthoteras pungens, Lyon, NEW SPECIES. Figure 7. Forewing. Figure 8. Me-
sosomal configuration, dorsal view. Figure 9. Metasoma, lateral view. Figure 10. Monothalamous gall,
sagittal section, showing structure and position of larval cell. Figure 11. Leaf of Q. pungens, showing
gall on margin of upper leaf surface. Figure 12. Head, frontal view.
by its shortened forewing with distinctive venation, stout, moniliform antenna and
by the posterior incomplete notauli, represented by a row of punctures in the
anterior region of scutum.
Gall.—Monothalamous and dome-shaped (Figs. 10, 11), projecting 3-5 mm
from upper leaf surface; larval cells basal, with radiating filaments to wall. Adults
mature and emerge from late December to mid-January.
Host.—Quercus pungens.
186 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Etymology.—This species is named for its host.
Material Examined.—See types.
ANTRON MADERA LYON, NEW SPECIES
(Figs. 13, 14)
Holotype female; ARIZONA. PIMA CO.: Santa Rita Moutains, Tucson, 1 Apr
1972, R. J. Lyon, from galls on Quercus arizonica Sargent; deposited: U.S. Na-
tional Museum of Natural History, Washington, D.C. Paratypes: same data as
holotype except collected 10—15 Apr 1972 on Q. toumeyi Sargent; deposited: U.S.
National Museum of Natural History, Washington, D.C. (1), R. J. Lyon collection
(3).
Description.—Unisexual female. Light yellow-brown; eyes, ocelli, tips of tarsi, terminal antennal
segments and ovipositor sheaths darker brown. Length 2.5-2.7 mm (* = 2.65 mm; n = 5). Head
transverse, as broad as mesosoma; gena slightly broadened behind eyes; interocular space wider than
high; malar space less than one-third as long as eye, groove absent; antenna 13-segmented, segment
3 longer than 4 and succeeding segments, segment 13 longer than 12. Scutum longer than broad,
surface coriaceous, pubescent laterally; notauli complete, curving laterally. Scutellum elongate, longer
than broad, rugose along margins, rounded behind; pits represented by a shallow, arcuate groove.
Mesopleuron slightly bulging, smooth, shining, with white setae along anterior and ventral edges.
Propodeal carinae arcuate, enclosure smooth. Legs pubescent; claw dentate. Forewing pubescent; veins
brown; margin ciliate; M, reaching Basal Vein; areolet small; Radial Cell 3.5X as long as broad; faint,
darkened, translucent patches in Radial, Cubital, and Discoidal Cells. Metasoma as long as high,
smooth and shining; three terga visible along dorsal curvature; tergum 2 with small pubescent patch.
Ventral spine stout, bristly, 3X as long as broad in lateral view.
Diagnosis.—Separable from other species in this genus by the following com-
bination of characteristics: tergite 2 not foliiform, antenna 13 segmented, foveal
pits replaced by a shallow, arcuate groove, and faint, translucent, dark patches in
the Radial, Cubital and Discoidal Cells.
Gall.—Monothalamous, spherical (Figs. 13, 14), small (3 mm diameter); at-
tached to midrib on undersurface of leaf, flattened at point of attachment. Mature
galls are slightly wrinkled and have scattered red and white microstellate hairs.
The larval cell is centrally located and surrounded by thick nutritive layer that
becomes spongy in older galls.
Hosts.—Quercus arizonica and Q. toumeyi.
Etymology.—This species is named for its site of occurrence; Madera Canyon,
Santa Rita Mts., Arizona.
Material Examined.—See types.
ANTRON DAILEYI LYON, NEW SPECIES
(Fig. 17)
Holotype female; TEXAS. EL PASO CO.: Franklin Mountains, El Paso, 23
Mar 1970, R. J. Lyon, from galls on Quercus pungens deposited: U.S. National
Museum of Natural History, Washington, D.C. Paratypes: same data as holotype
except collected 23—26 Mar 1970; deposited: U.S. National Museum of Natural
History, Washington, D.C. (8), California Academy of Sciences, San Francisco
(8), Bohart Museum of Entomology, University of California, Davis (6), Natural
History Museum of Los Angeles County, Los Angeles (6), R. J. Lyon collection
(30).
Description.—Unisexual female. Black, with red tinges; basal five antennal segments, tibiae and
1996 LYON: NEW SOUTHWESTERN CYNIPID WASPS 187
18
Figures 13-19. Galls. Figures 13-14. Antron madera, Lyon, NEW SPECIES. Figure 13. Gall
attached to upper surface midrib. Figure 14. sagittal section of gall showing detail of larval cell and
thickened cell wall. Figures 15-16. Antron franklinensis, Lyon, NEW SPECIES. Figure 15. Position
of twig gall on Q. turbinella. Figure 16. Sagittal section of gall showing detail of larval cell and
thickened cell wall. Figure 17. Antron daileyi, Lyon, NEW SPECIES. Galls on twigs of Q. pungens.
Figures 18—19. Andricus flocculentus, Lyon, NEW SPECIES. Figure 18. Cluster of woolly galls on
midrib of Q. pungens. Figure 19. Sagittal sectons of galls showing detail of individual galls.
188 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
tarsi light brown. Length 1.7—2.6 mm (x = 2.2 mm; n = 54). Head transverse, as broad as mesosoma;
slightly broadened behind eyes; occiput concave; median ridge extending from median ocellus nearly
to level of antennal sockets; interocular space almost 3X as broad as high; malar space one-third eye
length, striate; ocellar area and frons rugose, frons with white bristles. Antenna 14-segmented, segment
3 longer than 4, segment 14 2X as long as 13. Scutum slightly broader than long, prominently high-
arched in side view, coriaceous, shining and with white bristles near posterior and lateral margins;
notauli complete; anterior lines appear as a pair of smooth, shining, parallel streaks extending poste-
riorly from the anterior margin of the scutum, some specimens with very short median line. Scutellum
marginate, coarsely rugose and with white bristles; foveal pits deep, separated, with smooth, shining
bottoms. Propodeum slightly rugose, but without distinct carinae. Mesopleuron bulging, partly acic-
ulate. Legs pubescent; tarsal claws prominently dentate. Forewing hyaline, veins dark brown, pubes-
cent, and with ciliate margins; areolet well-developed, extending one-fourth distance to Rs+M; M,
not reaching Basal Vein, Rs, angled; Rs, not enlarged at wing margin; Rs+M with distinct “knobs”
in some specimens; Radial Cell 3X as long as broad. Metasoma longer than high, only three terga
visible along dorsal curvature; tergum 2 smooth and shining, remaining terga punctate; ovipositor
sheaths protruding, punctate and bristly. Ventral spine bristly, 3X as long as broad in lateral view,
triangular in ventral view (Fig. 28).
Diagnosis.—This species can be separated from the other species in the genus
by the following combination of characteristics: gena, in frontal view, broadened
behind eyes; tergite 2 not foliiform; antenna with 14 segments; scutum high-
arched in side view.
Gall.—(Fig. 17) - Monothalamous, thin-walled blisters that burst from cracks
in the bark of twigs; galls are smooth, tan, 5-7 mm long. Adult females emerged
23-26 Mar and oviposited in unopened, dormant buds.
Host.—Quercus pungens. Galls of this type were also seen on Q. arizonica, Q.
grisea Liebmann and Q. oblongifolia Torrey but no wasps were reared from them.
Etymology.—This species is named for Charles Dailey, Sierra College, Rocklin,
California, who has made many contributions to our knowledge of the Cynipidae.
Material Examined.—See types.
ANTRON FRANKLINENSIS LYON, NEW SPECIES
(Figs. 15, 16)
Holotype female; TEXAS. EL PASO CO.: Franklin Mountains, El Paso, 29
Dec 1962, R. J. Lyon, from galls on Quercus turbinella Greene; deposited: U.S.
National Museum of Natural History, Washington, D.C. Paratypes: same data as
holotype; deposited: U.S. National Museum of Natural History, Washington, D.C.
(1), R. J. Lyon collection (3).
Description—Unisexual female. Uniformly light brown; terminal antennal segments and tarsi darker
brown. Length 2.9-3.1 mm (x = 3.0 mm; n = 5). Head transverse, as broad as mesosoma; gena
broadened behind eyes; occiput strongly concave; interocular space 2X as wide as high; malar space
shorter than eye length and without groove. Antenna 13-segmented, segment 3 longer than 4, segment
13 equal to 12 in length. Scutum coriaceous, with white bristles, strongly convex in profile, slightly
broader than long; notauli incomplete, lateral area smooth and shining; anterior lines barely visible as
smooth streaks. Scutellum with microscopic, longitudinal ridges partially obscured by long, white
bristles; foveal pits replaced by an arcuate groove. Mesopleuron smooth and polished, slightly pro-
tuberant beneath wing base. Propodeal carinae short, arcuate. Legs pubescent; tarsal claws with long
tooth. Forewing hyaline and pubescent, margins ciliate; veins dark chocolate-brown; areolet small,
extending one-fifth of distance to Basal Vein; M, reaching Rs+M; Rs+M with distinct spur at junction
with M+Cu; Rs, arcuate, terminating in a flattened club; Radial Cell 2X as long as broad. Metasoma
longer than high, three terga visible along dorsal curvature, tergite 2 folitiform; tergites impunctate.
Ventral spine bristly, broad and tapering to point, 3X as long as wide in profile.
1996 LYON: NEW SOUTHWESTERN CYNIPID WASPS 189
Diagnosis.—This species can be separated from the other species in the genus
by a combination of the following characteristics: tergite 2 foliiform; notauli in-
complete; distinctive wing venation; Rs, arcuate, terminating in a flattened club;
gena distinctly broadened behind eyes.
Gall.—(Figs. 15, 16).—A small monothalamous gall, 7 mm long, thorn-shaped;
appears on twigs in late fall; mature adults collected in December.
Host.—Quercus turbinella.
Etymology.—This species is named for the Franklin Mountains where the host
species of oak occurs.
Material Examined.—See types.
KEY TO THE DESCRIBED SPECIES OF ANTRON KINSEY IN THE UNITED STATES
1. Tergites 2-7 abundantly pubescent; tergite 2 foliiform (Fig. 20) .... 2
he Tergite 2 with pubescent patch only, other terga bare; tergite 2 fo-
hifonmronmoes (Eig 2 Bye hereto ie es te re ld eek ben 3
2(1). Head, in frontal view, with gena visible and broadened behind eyes
(Fig. 22); Cubital Cell clear or with very faint darkened patches
in some specimens; apex of ventral spine rounded in ventral view
(Fig. 26). Light red-brown species from rounded, lead-colored
galls on Quercus arizonica, Q. toumeyi, and Q. oblongifolia in
PNT Z- OTA Fe ts fe OMe Te Pee Ny EL RIE eh A. plumbeum (Weld)
page Head, in frontal view, with gena not visible or broadened behind
eyes (Fig. 23); Cubital Cell with two distinct dark, translucent
patches sometimes fused; apex of ventral spine acute in ventral
view (Fig. 28). Dark brown or red-brown species from red, woolly
clusters on midribs beneath leaves of Q. arizonica, Q. toumeyi,
and Q. oblongifolia in Arizona ......... A. quercusnubila (Bassett)
3(1’). Notauli incomplete; tergite 2 foliiform; forewing clear, with small,
dark patch around Rs,. Light brown species from small, thorn-
shaped twig galls on Q. turbinella in western Texas (Fig. 15)....
Cee ee, eee, ae. nes ia ers Ste One A. franklinensis Lyon
che Notauli complete; tergite 2 foliiform or not ...................4.0. 4
4(3'). Gena in frontal view visible and broadened behind eyes ........... 5
4’. Gena in frontal view not visible or broadened behind eyes ....... 10
5(4). lercite 2-nottokirorms 24%... 82 28), Sener een coe ee 6
ae Feri fe sero litt OLN aa Ak ge ee eile en PhD cite tet sc. gL ater 7
6(5). Antenna 14-segmented; all terga beyond second punctate; scutellum
rugose, foveal pits deep, separated, with shining bottoms; scutellar
tip broadly rounded; wing hyaline, clear, M, not reaching Basal
Vein; ventral spine bristly with triangular tip (Fig. 28); dark spe-
cies from thin-walled blisters that burst from cracks on the twigs
of Q. pungens in Arizona, New Mexico and western Texas......
ats Meal PON Ree teen ans SES Coe oO sD APNE Me ep eee ae A. daileyi Lyon
6’. Antenna 13-segmented; terga smooth, impunctate; scutellum rugose
only on the side; foveal pits replaced by a shallow, arcuate groove;
scutellar tip sloping to a narrow but rounded point; wing with faint
darkened patches in the Radial, Cubital and Discoidal Cells; M,
reaching Basal Vein; ventral spine bristly with rounded, notched
190 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
20 21
| {
26 27 28 29
Figures 20-29. Key characters in Antron taxonomy. Figure 20. Foliiform metasomal tergum 2.
Figure 21. Non-foliiform metasomal tergum 2. Figure 22. Frontal view of head to show gena broadened
behind eyes. Figure 23. Frontal view of head to show gena not broadened. Figure 24. Oval-shaped
head in frontal view. Figure 25. Deltoid-shaped head in frontal view. Figure 26. Ventral spine of
metasoma, ventral view, showing rounded apex. Figure 27. Ventral spine, curving to blunt point.
Figure 28. Ventral spine with triangular tip. Figure 29. Ventral spine, with blunt, notched tip.
1996
9(7').
9".
10(4’).
10’.
11(10’).
11’.
LYON: NEW SOUTHWESTERN CYNIPID WASPS 191
tip (Fig. 29). Light yellow-brown species from small, spherical,
thick-walled galls on midrib of the undersurfaces of leaves of Q.
arizonica, Q. oblongifolia and Q. toumeyi in Arizona (Fig. 13)
1s Ae Se Oy Vag) ars ae a ee Pe A A. madera Lyon
Atitenna P3-seemenmted® 1. er Sesion 5 a oo ence sae 8
Antenna 14-segmented
Head oval in frontal view (height about 0.66 X< width, (Fig. 24);
scutum micropunctate and somewhat roughened; scutellum finely
rugose. Dark, almost black species from spiny spring galls on mid-
rib of undersurface of leaves of Q. gambelii, QO. toumeyi, and Q.
undulata Torr. in Arizona ...............4-- A. acras piformis Weld
Head deltoid in frontal view (almost as high as wide) (Fig. 25);
scutum smooth, shining; scutellum microscopically coriaceous.
Brown species from globular (3 mm diameter), thin-walled galls
on undersurfaces of leaves of Q. turbinella in Arizona.....
ON OR a ahs eT Eee ee ere ec ere ert aire A. magdalenae Weld
Arcuate foveal furrow, at base of scutellum, narrow, shining and
almost smooth at bottom; scutellum margined, ridges extending
onto sides of disc. Red-brown species from, distinctive, spiny,
club-shaped leaf galls on Q. dumosa Nutt. and Q. lobata Née in
Calstornid. yo 5.2 pee. ae oe Lemuel erer atm Welers) a0 A. douglasii (Ashmead)
Foveal furrow broad, ridged on bottom; scutellum margined on sides,
but ridges not extending onto disc. Darkly infuscated species from
spiny leaf galls resembling miniature ‘‘sea urchins’? on Q. doug-
lasii Hook. & Am. in California ............ A. echinus (Ashmead)
Antenna 15-segmented; wings hyaline, clear; scutum smooth, shin-
ing. Amber yellow species from terminal, cap-shaped bud galls on
Q. turbinella in New Mexico ..................25: A. pileus Weld
Antenna 14-segmented; wings with dark patches in Cubital Cell;
SCULMIM-COMAGCEOMS, Tin) aie ot Be |, Leet ios retard meee alt 11
Disc of scutellum longer than broad and coarsely rugose; Rs, of
forewing only slightly enlarged at wing margin; outline of head
deltoid in frontal view (Fig. 25). Dark species from globular bud
galls on Q. dumosa and Q. douglasii in California (possible bi-
sexual generation of A. echinus) ................. A. ribes (Kinsey)
Disc of scutellum as broad as long and finely rugose; Rs, forming a
distinct club at wing margin; outline of head, oval in frontal view
(Fig. 25). Light brown species from thin-walled bud galls on Q.
douglasii and Q. lobata in California (possible bisexual generation
of A. douglasii) .............005. A. lobata (McCracken & Egbert)
ACKNOWLEDGMENT
I thank John M. Tucker (University of California, Davis) for identifying spec-
imens of Quercus pungens; Arnold Menke (U.S. National Museum of Natural
History, Washington D.C.) for assistance with the Antron key; Roy Snelling (Los
192 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Angeles County Museum of Natural History, Los Angeles) for his continual en-
couragement and constructive comments during the writing of this paper.
LITERATURE CITED
Lyon, R. J. 1993. Synonymy of two genera of cynipid gall wasps and description of a new genus
(Hymenoptera: Cynipidae). Pan-Pacific Entomol. 69(2): 133-140.
Weld, L. H. 1960. Cynipid galls of the Southwest. Edwards Bros., Ann Arbor, Michigan (51 pp.)
Received 12 Mar 1995; Accepted 7 Mar 1996.
PAN-PACIFIC ENTOMOLOGIST
72(4): 193-201, (1996)
BIOGEOGRAPHICAL ASPECTS OF THE ANT FAUNA OF
CORSICA (HYMENOPTERA: FORMICIDAE)
JANINE CASEVITZ-WEULERSSE
E.P. 90 du CNRS and Laboratoire d’Entomologie, MNHN, 45 rue Buffon,
F-75005 Paris, France
Abstract——Corsican myrmecofauna (26 genera, 83 species) is mostly European in characteristics
and shows very low endemism (only 3 endemic species). A comparison with 15 neighboring
countries shows its particular similarity with the faunas of Sardinia and Provence. Of the 83
species, 26 are widely distributed in most of these 15 countries. The origins of Corsican ant
populations are discussed. The development of a modern fauna in Corsica before the island
became isolated from the continent, human activity, distinctive conquering capacities of ants,
their strong ability to settle in many habitats, interspecific competition and the great diversity of
landscapes of the island explain the present composition of the myrmecofauna of Corsica.
Key Words.—Insecta, Corsica, Formicidae, biogeography, neighboring countries, competition.
Corsica is an island located in the western part of the Mediterranean, not far
from the coast of Provence (160 km) and Italy (82 km), about 50 km from the
island of Elba, only 12 km from Sardinia, and more than 450 km from North
Africa and Spain. It is the smallest, but the most mountainous of the three Tyr-
rhenian islands and it has been inhabited by Man for at least 10,000 years. Fau-
nistic and systematic data on ants of Corsica obtained in the last twenty years
(Casevitz-Weulersse 1974, 1986a, b, 1990a, b) demonstrate the large diversity and
wealth of the Corsican myrmecofauna, which is composed of 26 genera and 83
species. An ecological study of these 83 species and their distribution in the island
had been presented in a previous paper (Casevitz-Weulersse 1990c). We present
here the principal results of the biogeographical analysis of the Corsican ant fauna
and compare it to neighboring countries.
GENERAL CHARACTERS OF THE CORSICAN MYRMECOFAUNA
Holldobler & Wilson (1990) list eleven and Bolton, more recently (1994), six-
teen extant subfamilies in the family Formicidae. The four with the greatest world-
wide distribution (Ponerinae, Myrmicinae, Dolichoderinae, Formicinae) are pres-
ent in Corsica, together with a fifth subfamily (Leptanillinae) (Table 1).
Most Corsican species generally belong to the European Region and their dis-
tribution is either cosmopolitan (2.41%), holoarctic (8.43%) or palaearctic
(21.69%), or European (36.15%) only. The strictly Mediterranean species are less
common (27.71%). Endemism is very low in the Corsican myrmecofauna; only
three of the 83 species present on the island can be qualified as endemic today,
with the caveat that endemism is a very relative and variable concept. For ex-
ample, several species which had long been considered as strictly cyrno-sardinian
were shown by enlarged faunistic surveys and taxonomic revisions to be much
more widely distributed, e.g., Leptanilla revelierei Emery, Messor wasmanni
Krausse, (Casevitz-Weulersse 1992) or Epimyrma corsica (Emery) (Buschinger
& Winters 1985). Two of the three endemic Corsican species have been recently
described: Leptothorax melas by Espadaler et al. (1984b) and Stenamma orousseti
by Casevitz-Weulersse (1990a); the third species, Tetramorium sp., is under study.
194 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Table 1. Formicidae of Corsica: subfamilies, genera and number of species for each genus.
Sub-families Genera Number of species
(26) (83
Leptanillinae Leptanilla
Ponerinae Cry ptopone
Hypoponera
Ponera
Myrmicinae Smithistruma
Epitritus
Myrmica
Stenamma
Aphaenogaster
Messor
Pheidole
Crematogaster
Solenopsis
Monomorium
Myrmecina
Le ptothorax
Epimyrma
Tetramorium
Strong ylognathus
Dolichoderinae Tapinoma
Bothriomyrmex
Line pithema
Formicinae Plagiolepis
Lasius
Camponotus
Formica
NANION RK RK WR NW NHR KB NR KP HhRPN PRK NKN HR eH
COMPARISON WITH OTHER REGIONS
French Mainland.—The island shares 69 of its 83 species with continental
France, which includes 44 genera and about 180 species (Bernard 1968). How-
ever, 14 Corsican species, including Aphaenogaster spinosa Emery and Messor
minor (André) which are among the most common ants on the island, are not
found on mainland France. However, a dozen genera included in the mainland
fauna are absent in Corsica. In particular, Manica (Myrmicinae), Dolichoderus
(Dolichoderinae), Cataglyphis, Proformica and Polyergus (Formicinae) are prob-
ably really absent, because they are immediately noticed when they are present
in a biotope. Other genera which have not been found on the island include
parasitic and very rare species; it is possible that they will be discovered there in
the future.
Other Mediterranean Regions.—The myrmecofaunas of fifteen Mediterranean
regions where reliable faunistic surveys have been published in recent decades
were compared to that of Corsica (Table 2). At the genus level, these regions
have a high number of genera in common with Corsica.
At the species level, their degree of similarity with Corsica was estimated using
the Jaccard’s coefficient (JC) according to the following formula : JC = (Cab X
100)/(A + B — Cab), with Cab = number of species in common in regions A
and B; A and B = total number of species in region A and region B. By con-
1996 CASEVITZ-WEULERSSE: ANT FAUNA OF CORSICA 195
Table 2. Comparison between the myrmecofaunas of fifteen Mediterranean regions and Corsica
(8720 km’, 26 genera, 83 species).
Total no. of Total no. of
genera (no. of species (no. of
Surface genera common species common Jaccard’s
Regions (km?) with Corsica) with Corsica) coefficient
Sardinia! 24,100 24 (24) 64 (52) 54.74
Provence? 26,135 34 (25) 116 (60) 43.17
Tuscan Archipelago? 1000 22 (21) 45 (35) 37.63
Liguria‘* 5415 29 (24) 78 (42) g5229
Mt Ventoux? 2 21 (16) 63 (38) 35.19
Sicily® 25,500 29 (23) 79 (41) 33.88
North of Spain’ 252,370 34 (25) 136 (54) 32.74
Latium® 17,200 22 (19) 53 (33) 32.04
Tuscany? 29,000 26 (21) 63 (35) 31.53
All of Spain! 504,748 44 (26) 214 (63) 26.92
Yugoslavia!! 225,800 38 (25) 161 (51) 26.42
Portugal? 90,000 20 (17) 73 (30) 23.81
Baleoric Islands” 4980 19 (19) 42 (24) 23.76
Greece!* 131,944 39 (25) 222 (46) 17.76
Algerian Forests!> ? 35 (23) 114 (24) 13.87
' Baroni-Urbani 1971, Casevitz-Weulersse 1974. ? Soyer 1951, Ovazza 1954, Bernard 1968, 1983,
Gaspard 1968. ? Baroni-Urbani 1971. * Baroni-Urbani 1971. > Du Merle 1978. © Baroni-Urbani 1971.
7 Collingwood 1978, Collingwood & Yarrow 1969, Espadaler 1979, 1980, 1981, 1986a, b, Espadaler
& Collingwood 1982, Espadaler & Restrepo 1983, Espadaler & Riasol 1983, Espadaler & Roda 1984,
Espadaler et al. 1984, Martinez-Ibanez & Espadaler Gelabert 1986, Tinaut 1985, 1987a, b, 1988.
8 Baroni-Urbani 1971. ? Baroni-Urbani 1971. '° (see 7). '! Agosti & Collingwood 1987. !* Collingwood
1978. Collingwood & Yarrow 1969, Comin del Rio & De Haro Vera 1980, Comin del Rio &
Espalader Gelabert 1984. '* Agosti & Collingwood 1987. '5 Cagniant 1968, 1973.
vention, A = number of species in Corsica (83) and B = total number of species
in the country being compared (Table 2). The distribution of all the species in-
cluded in the Corsican myrmecofauna and their presence in these 15 Mediterra-
nean regions had been listed in a preceding paper (Casevitz-Weulersse 1992).
Note that Corsica (with an area of 8720 km’) has 83 species, but Sardinia (area
2.8 times larger) has only 64 species and Sicily has 79. Several large continental
regions also have a smaller number of species in relation to their area when
compared to Corsica. On contrast, Mount Ventoux, the Tuscan archipelago, and
the Balearic islands have a diversified and rather rich myrmecofauna. These ob-
servations are partly due to a lack of faunistic surveys and taxonomic studies in
many southern European countries.
In spite of missing or incomplete data, comparisons are still interesting. As
expected, considering the palaeogeographical history of the Mediterranean region,
Sardinia (JC: 54.74) has the highest biocoenotic similarity with Corsica, followed
by Provence (JC: 43.17). Similarity with western Mediterranean regions, conti-
nental or insular, is very low (all of Spain, Balearic islands, Portugal), as it is for
more distant regions (Greece and Algerian forests).
Table 3 indicates that 26 species of the Corsican myrmecofauna have a wide
geographical distribution in the Mediterranean region. Most of them also have a
large worldwide distribution. Only two species are strictly Mediterranean, the rest
belong to the European, palaearctic, or holoarctic domains.
Table 3. 26 Corsican species with a wide geographical range in 15 Mediterranean regions and their world distribution.
961
OAANINNBRWN KE
No. of
Mediterranean
regions
Species inhabited Absent from: World distribution
Ponera coarctata (Latreille) 15 european
Aphaenogaster subterranea (Latreille) 13 Tuscany, Portugal palaearctic
Messor capitatus (Latreille) 14 Greece mediterranean
Messor structor (Latreille) 14 Algerian forests european
Pheidole pallidula (Nylander) 15 palaearctic
Crematogaster scutellaris (Olivier) 15 european
Solenopsis fugax (Latreille) 12 Portugal, Balearic Islands, Algerian forests palaearctic
Myrmecina graminicola (Latreille) 15 palaearctic
Leptothorax nylanderi (Forster) 13 Balearic Islands, Algerian forests european
Le ptothorax recedens (Nylander) 12 Tuscan Archipelago, Tuscany, Portugal european
Leptothorax tuberum (Fab.) 12 Tuscany, Balearic Islands, Algerian forests european
Tetramorium caespitum (L.) 14 Algerian forests holoarctic
Tetramorium semilaeve André 12 Liguria, Mt Ventoux, Algerian forests palaearctic
Tapinoma nigerrimum (Nylander) 12 Mt Ventoux, Yugoslavia, Greece mediterranean
Plagiolepis pygmaea (Latreille) 15 Algerian forests european
Lasius alienus (Forster) 15 holoarctic
Lasius emarginatus (Olivier) 13 Balearic Islands, Algerian forests palaearctic
Lasius flavus (Fab.) 14 Balearic Islands holoarctic
Lasius niger (L.) 15 holoarctic
Camponotus aethiops (Latreille) 12 Portugal, Balearic Islands, Algerian forests palaearctic
Camponotus lateralis (Olivier) 15 palaearctic
Camponotus piceus (Leach) 14 Algerian forests european
Camponotus truncatus (Spinola) 13 Latium, Portugal palaearctic
Camponotus vagus (Scopoli) 13 Tuscany, Balearic Islands european
Formica cunicularia Latreille 13 Balearic Islands, Algerian forests european
Formica fusca (L.) 12 Tuscan Archipelago, Sicily, Balearic Islands holoarctic
LSIDO'IONOLNA OIMIOVd-NVd FHL
(D)ZL IOA
1996 CASEVITZ-WEULERSSE: ANT FAUNA OF CORSICA 197
As is the case for the Corsican populations, the myrmecofaunas of the entire
Mediterranean region are primarily European.
To sum, Corsica has a diversified myrmecofauna with a rather high number of
species, considering its area. However, this fauna is not original and it is consti-
tuted mostly of generalist, ubiquitous species with a wide distribution.
ORIGIN OF THE CORSICAN ANT POPULATIONS
Ants are social insects. Reproduction of their colonies and propagation of their
species follow particular rules. The creation of new colonies and their implanta-
tion at a given location differ depending on the species and can occur from the
aerial arrival of winged, impregnated females; along or in groups, from the move-
ment of portions of colonies including one or several egg-laying wingless females
and workers, or from the isolation of workers with larvae of future queens and
males. In some cases, groups of isolated workers without brood (eggs, larvae,
pupae) can also originate new colonies by various ways (inseminated workers or
parthenogenesis). Species can reach an island either by active (flight of winged
sexual individuals) or passive transportation. However, the queens of many ant
Species are unable to cover long distances by air, unless they are small enough
to be carried by air currents.
As with ants, an European or a palaearctic component of the fauna is often
demonstrated for other insects and invertebrates from Corsica. However, endem-
ism rates vary considerably among the groups, depending on their age and mode
of dispersal. For many groups, endemism is rather high and occurs at the species,
or more often subspecies level but rarely at the generic level. Corsican populations
of Coleoptera (Sainte-Claire Deville 1906/1914, 1926; Jeannel 1942, 1961a, b),
Plecoptera, Trichoptera, blepharocerid Diptera (Giudicelli 1975, Cianficconi &
Moretti 1990), Araneae (Canard 1989), Diplopoda (Mauries 1969) and terrestrial
Oligochaeta (Omodeo 1961, Omodeo & Rota 1987) are largely composed of
species which have evolved locally from a Tertiary fauna that found a refuge on
the island, particularly on its high mountains.
The situation is different with the ants. The last link between Corsica and the
continent seems to have occurred between 6 and 5 million years ago during the
Messinian episode, and also during the early ice ages. Corsica has been a true
island only in the last 100,000 years (Gauthier & Vigne 1987). When links existed
with the continent, at the time of the early ice ages at the beginning of the
Quaternary era, the myrmecofauna probably already included genera and species
identical to those that can be observed today (Casevitz-Weulersse 1992). The
intermittent indirect links between Corsica and continental Europe at the time of
the first ice ages allowed, on several occasions, the passage of various species
which were already modern. If the low endemism seen here is true, it would
demonstrate that there has been almost no local ant evolution.
Recent studies in Corsica and Sardinia on vertebrate, particularly mammalian
faunas, have shown that the current situation is essentially due to voluntary and
involuntary human action (Vigne & Alcover 1985). All present day taxa in Cor-
sica were absent before the Holocene (Vigne 1987, 1992) and their endemism is
low, at the level of a few subspecies. In the same manner as the mammals, it is
quite possible that ant populations were constituted in part from species introduced
198 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
by Man to this island, which was one of the first to be colonized in the Mediter-
ranean.
Finally, it should be noted that our conclusions on the Mediterranean Basin
myrmecofauna are very close to those of Blondel (1982, 1984) concerning the
biogeographical origin of avifaunas. Blondel observed the same European aspect
of the populations and low endemism in birds nesting in Corsica and in the rest
of the Mediterranean region that is also observed in ants.
The introduction of ants in Corsica probably occurred before the final separa-
tion of the island from the continent. The paleogeographical history of the island
and the paleontology of the Formicidae explain why this distribution of subfam-
ilies is similar to that of most temperate countries (Casevitz-Weulersse 1992).
However, the present composition of this fauna can also be explained by taking
into account the early arrival of Man, whose essential role in the modern com-
position of the Corsican faunas we are only now beginning to understand.
In addition to human exchanges between the island and the continent during
the post-Pliocene, several factors came into play for the creation of the myrme-
cofauna as we can observe it today. As emphasized by Bernard (1968) and La-
motte (1986), chance plays a large role in the installation of diverse species on
an island. The early comers occupy all the habitats and leave no room for others.
The lack of success in Corsica of a species as conquering as Linepithema
humile, the “‘Argentine ant’’ (Casevitz-Weulersse 1986b, 1992) and the cohabi-
tation in the same biotope of several species on the same genus, which seem to
exploit the same food sources, e.g., Messor wasmanni, M. minor, and M. capitatus
on backshores (Casevitz-Weulersse 1990c) or Lasius niger and L. emarginatus in
citrus orchards (Casevitz-Weulersse 1993) are a few of many examples that dem-
onstrate competition and association phenomena. They will have to be analyzed
in detail if we want to understand how the ant populations in Corsica were con-
stituted and how they operate.
Competition, intrinsic capacity of species to occupy new territories, high col-
onizing power are characteristics which have allowed certain species to become
established at the expense of others. The particular diversity of Corsican environ-
ments has certainly also contributed to the persistence or the implantation of
widely varied species. All of these factors have contributed in the present char-
acteristics of ant populations in Corsica.
ACKNOWLEDGMENT
I am specially grateful to Dr Judith Najt for the critical reading of the manu-
script and her helpful comments.
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Received 6 Jun 1995; Accepted 14 Feb 1996
PAN-PACIFIC ENTOMOLOGIST
72(4): 202-208, (1996)
GENETIC VARIATION IN BOMBUS APPOSITUS CRESSON
(HYMENOPTERA: APIDAE)
RICHARD W. RUST AND JENNIFER L. PORTER
Biology Department, University of Nevada, Reno, Nevada 89557
Abstract.—Genetic variation was measured in five populations of Bombus appositus Cresson.
Scoring of 29 loci in 18 enzyme systems resulted in polymorphic loci in all populations and a
low estimated mean heterozygosity of 0.028 + 0.004 (SE). Neighbor-joining analysis did not
resolve the geographic pattern of the populations.
Key Words.—Insecta, Hymenoptera, Apidae, population genetics, allozymes, Bombus appositus
Low levels of genetic variation have been reported in most Hymenoptera when
compared to other insects (Graur 1985, Crespi 1991, Yik-Yuen et al. 1991). The
exceptions are in Argidae and Tenthredinidae (Sheppard & Haydon 1986). This
low level of variation is especially true for the bees (Apoidea) (Graur 1985;
Pamilo et al. 1978, 1984; Owen 1985; Packer & Owen 1989; Scholl et al. 1990;
Owen et al. 1992; Mullen & Rust 1994). Within the bumble bees (Bombus spe-
cies) significantly low levels have been reported for 27 species (Pamilo et al.
1984, Owen et al. 1992) or about 10% of all Bombus species (Thorp et al. 1983).
Heterozygosity estimates range from zero (0.0) in several species, Bombus nev-
adensis Cresson, B. californicus EF Smith, B. rufocinctus Cresson, B. mixtus Cres-
son, B. perplexus Cresson (Owen et al. 1992), to 0.044 in Bombus balteatus
Dahlbom (Pamilo et al. 1984). Owen et al. (1992) found the mean expected
heterozygosity for 16 Bombus species from North America to be low (0.008 +
0.007 (SE)). Mullen & Rust (1994) found low (0.008 + 0.004) but similar levels
of heterozygosity in commercially reared and natural populations of B. occiden-
talis Greene.
There are several explanations for the low levels of genetic variation in Bombus
(Pamilo & Crozier 1981; Graur 1985; Owen 1985, 1988; Owen et al. 1992). The
degree of social evolution in Hymenoptera appears to be negatively correlated
with genetic variability with the eusocial species having the lowest levels (Packer
& Owen 1989).
This study examines the genetic variation within and between five populations
of the bumble bee Bombus appositus Cresson. Bombus appositus was selected
for study because it is member of the subgenus Subterraneobombus (Thorp et al.
1983). This subgenus contains only two species in North America B. appositus
and B. borealis (Kriby) and has received no genetic study. Populations were
sampled from the southern limits of the species range in the isolated mountain
ranges of Great Basin Desert and the Rocky Mountain regions of western North
America (Thorp et al. 1983).
MATERIALS AND METHODS
Female B. appositus were collected from three sites in northern Nevada and
two sites in northern Utah (Table 1). Individual workers were collected from
flowers while hiking (approximately 5 km distance) in the collection area over
two or more days.
1996 RUST & PORTER: GENETIC VARIATION IN BOMBUS APPOSITUS 203
Table 1. Bombus appositus population sample sites (State, County, mountain range, latitude and
longitude, and average elevation) and number of female individuals examined.
Population sites '
Elevation
County Location Latitude & Longitude (meters) Number
Utah
Cache Wasatch Mountains
White Pine Lake 111°40’ x 41°55’ 2500 21
Salt Lake Wasatch Mountains
Mill Creek Canyon 111°40’ x 40°41’ 2400 16
Nevada
Elko Independence Mountains
Mill Creek 116°00’ X 41°31’ 2500 28
Elko Ruby Mountains
Thomas Creek Canyon 115°25' X 40°38’ 2500 22
Elko East Humboldt Range
Angle Lake 115°05' X 41°02’ 2500 20
All field captured individuals were placed in individual plastic vials with cotton
plugs and transported to the laboratory on ice and stored at —80° C. Individuals
were prepared for electrophoresis by separating the head and thorax from the
abdomen and homogenizing each in 0.05 ml of cold extraction buffer (Tris HCl
0.05 M, pH 7.0; May 1992). After 20 min of cold incubation and low-speed
centrifugation, the supernatant was pipetted into 1.5 ml eppendorf tubes and stored
at —80° C until used for electrophoresis. Supernatant was applied to 14% hori-
zontal starch gels (50% Connaught and 50% Sigma) using filter-paper wicks
(Whatman #3). Gels ran approximately five hours. All individuals were analyzed
within two weeks of preparation. We used the methods and staining procedures
described by May (1992). Twenty-eight enzyme systems were initially surveyed
and 18 were used for population analysis.
Genotype frequencies were obtained by direct count from the phenotypes ob-
served on the gels, and electromorph (allozyme) frequencies were calculated from
genotype frequencies. All polymorphisms were photographed. The most common
electromorph at each locus was designated as “‘C’’, with relatively faster migrating
allozymes scored as “‘B”’ and relatively slower migrating allozymes scored as
me aes
Expected genotypic frequencies, expected fit to Hardy-Weinberg equilibrium,
Nei’s unbiased expected heterozygosity and unbiased genetic identity (Nei 1978),
and Wright’s F-statistics (Wright 1978) were calculated using BIOSYS-1 (Swof-
ford & Selander 1981). Neighbor-Joining unrooted tree construction (Saitou &
Nei 1987) was produced with NJTREE (Jin & Ferguson 1990).
RESULTS
Seven of the 28 enzyme systems (AAT, AC, ADH, LDH, ODH, SDH, XDH)
yielded weak or no bands on the gels (Table 2). Two (GDH and G6PDH) were
inconsistent or too smeared to score. CK was determined to yield the same band-
ing pattern as AK and was dropped from scoring. The remaining 18 enzyme
systems yielded 29 scorable loci (Table 2). Seven of the 18 enzyme systems
204 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Table 2. Enzyme systems, loci, and electrophoretic conditions used to assay Bombus appositus,
tagged enzymes (*) were used in the genetic analysis.
Enzyme (loci) Symbol E.C. Number! Buffer? _— Tissue?
Aconitase AC 4.2.1.3 TC-1 A
Alcohol dehydrogenase ADH 1.1.1.1 TC-l A
*Adenylate kinase (1) AK 2.7.4.3 TC-1 A
Aspartate aminotransferase AAT 2.6.1.1 R A
Creatine kinase CK Ded. 2 4 T
*Diaphorase (NADH) (2) DIA 1.8.1.4 4 A
*Diaphorase (NADPH) (1) DIAP 1.8.1.4 R A
*Esterase (2) EST 3.1.1.- 4 ‘tt
*Galactosaminidase (1) GAM Meat ae C A
*Glyceraldehyde-3-phosphate dehydrogenase (1) GAPDH 1.2.1.12 C TA
Glucose dehydrogenase GDH 1.1.1.118 TC-1 A
*a-Glycerophosphate (3) dehydrogenase G3P 1.1.1.8 R TA
Glucose-6-phosphate dehydrogenase G6PDH 1.1.1.49 TC-1 A
*Glucokinase (1) GK seat wl Lae) 4 a5
*General Protein (1) GP nonspecific R T
*Glucosephosphate (2) isomerase GPI 5.3.1.9 R TA
*Hydroxybuteric (1) dehydrogenase HBDH 1.1.1.30 TC-1 A
*Isocitrate dehydrogenase (2) IDH 1.1.1.42 R T
Lactate dehydrogenase LDH Lolal27 4 A
*Leucine aminopeptidase (2) LAP 3.4.11.1 R TA
*Malic enzyme (2) ME 1.1.1.40 4 T,A
*Malate dehydrogenase (1) MDH 1B) Dis les 4 TA
Octanol dehydrogenase ODH L131,73 R A
*Peptidase Leu-Ala (2) PEP-LA 3.4.11 R T,A
*Phosphoglucomutase (3) PGM 5.4.2.2 4 T,A
Succinate dehydrogenase SDH 1.3.99.1 TC-1 A
*Superoxide dismutase (1) SOD 1.15.1.1 R LA
Xanthine dehydrogenase XDH 1.1.1.204 R A
' Nomenclature Committee International Union of Biochemistry 1984.
? Gel and tray buffer systems (May 1992).
3T = head and thorax, A = abdomen.
(DIAP, GAM, GAPDH, GP, IDH, LAP, MDH, and SOD) were fixed for the same
allele in all individuals.
Table 3 contains the allele frequencies in the B. appositus populations sampled.
The two Wasatch Mountains populations had both the fewest and most polymor-
phic loci, White Pine contained two (EST-2, PGM-2) and Mill Creek contained
six (AK-1, GK-1, GPI-2, ME-1, PGM-1, PGM-2). In the Great Basin mountain
populations, Ruby contained three (GK-1, GPI-2, PEP-1), Humboldt four (DIA-
1, DIA-2, GPI-2, G3P-3), and Independence six (DIA-1, DIA-2, EST-2, GPI-2,
PGM-2, PGM-3). All of the Humboldt and Ruby loci conformed to Hardy-Wein-
berg expectations. One locus each in the Independence (GPI-2, x? = 19.06, df =
1, P = 0.000) and White Pine (EST-2, x? = 5.33, df = 1, P = 0.02) samples and
five loci in the Mill Creek samples (AK-1, x? = 27.04, df = 1, P = 0.000; GPI-
2, x? = 31.03, df = 1, P = 0.000; ME-1, x? = 31.03, df = 1, P = 0.000; PGM-
1, x? = 31.03, df = 1, P = 0.000; PGM-2, x? = 31.03, df = 1, P = 0.000) did
not conform to Hardy-Weinberg expectations. All deviations were heterozygote
deficiencies. Genetic variability was estimated within each sample with three mea-
1996 RUST & PORTER: GENETIC VARIATION IN BOMBUS APPOSITUS 205
Table 3. Allele frequencies at polymorphic loci and number of individual females analyzed in
Bombus appositus.
eres a Utah Nevada
alleles White Pine Mill Creek Humboldt Ruby Independence
n 21 16 20 22) 28
AK-1 B 0.000 0.062 0.000 0.000 0.000
C 1.000 0.938 1.000 1.000 1.000
DIA-1 B 0.000 0.000 0.000 0.000 0.036
C 1.000 1.000 0.975 1.000 0.964
D 0.000 0.000 0.025 0.000 0.000
DIA-2. B 0.000 0.000 0.125 0.000 0.035
C 1.000 1.000 0.875 1.000 0.965
EST-2. B 0.333 0.000 0.000 0.000 0.000
C 0.667 1.000 1.000 1.000 0.938
D 0.000 0.000 0.000 0.000 0.072
GK-1 B 0.000 0.031 0.000 0.000 0.000
C 1.000 0.969 1.000 0.864 1.000
D 0.000 0.000 0.000 0.136 0.000
GPI-2. B 0.000 0.062 0.050 0.000 0.000
C 1.000 0.938 0.950 0.955 0.893
D 0.000 0.000 0.000 0.045 0.107
G3P-3 B 0.000 0.000 0.025 0.000 0.000
c 1.000 1.000 0.975 1.000 1.000
ME-1 C 1.000 0.938 1.000 1.000 1.000
D 0.000 0.062 0.000 0.000 0.000
PEP-1 B 0.000 0.000 0.000 0.045 0.000
C 1.000 1.000 1.000 0.955 1.000
PGM-1 C 1.000 0.062 1.000 1.000 1.000
D 0.000 0.938 0.000 0.000 0.000
PGM-2 B 0.188 0.000 0.000 0.000 0.000
C 0.812 0.938 1.000 1.000 0.965
D 0.000 0.062 0.000 0.000 0.035
PGM-3 B 0.000 0.000 0.000 0.000 0.143
C 1.000 1.000 1.000 1.000 0.857
sures: mean expected heterozygosity, percentage of polymorphic loci, and mean
number of alleles per locus (Table 4). Mean expected heterozygosity ranged from
0.014 (Humboldt) to 0.03 (Independence). Percentage of polymorphic loci was
low and ranged from 6.9% (White Pine) to 20.7% (Mill Creek and Independence).
Mean number of alleles per locus was 1.1 or 1.2, in all populations.
A Neighbor-Joining phenogram (Saitou & Nei 1987) based on Nei’s (1978)
unbiased genetic identity (Table 5) was unresolved and did not represent the
geographical relationships of the surveyed populations of B. appositus (Fig. 1).
First, there was almost no separation of the populations (patristic values 0.0002)
and branch length were identical (0.49). Second, the two Wasatch Mountain and
three Great Basin mountain populations did not nest together; White Pine was
associated with Humboldt (cycle 1), Mill Creek with Independence (cycle 3), and
Ruby was placed between the two groups (cycle 2).
Hierarchical F-statistics for the 12 polymorphic loci indicate a high inbreeding
coefficient within the populations (F,, = 0.382) and a moderate level (Hartl &
Clark 1989) of differentiation among the populations (F,, = 0.105). The combined
206 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Table 4. Genetic variability at 29 loci in all populations of Bombus appositus. Values in parentheses
are standard errors.
Mean number of Percentage of Mean expected
Population alleles per locus polymorphic loci! heterozygosity?
Utah
White Pine 1.1 (0.0) 6.9 0.030 (0.021)
Mill Creek 1.2 (0.1) 20.7 0.024 (0.009)
Nevada
Humboldt 1.1 (0.1) 13.8 0.014 (0.008)
Ruby 1.1 (0.1) 10.3 0.015 (0.009)
Independence 1.2 (0.1) 20.7 0.025 (0.011)
' A locus is considered polymorphic if the frequency of the most common allele does not exceed
0.99.
2 Unbiased estimate (Nei 1978).
effects of nonrandom breeding and random genetic drift in the populations is also
high (F,, = 0.447).
DISCUSSION
The mean heterozygosity (H,,, 0.021 + 0.011) in B. appositus is very low in
relation to other Hymenoptera and insects (Graur 1985, Owen 1985, Crespi 1991).
However, it is relatively high with respect to other species in the genus Bombus.
Of the 27 species analyzed, only B. balteatus (0.044), B. sylvicola Kirby (0.042),
B. melanopygus Nylander (0.037), B. terrestris (L.) (0.037), and B. hypnorum (L.)
(0.025) have reported higher heterozygosity measures (Pamolo et al. 1984, Owen
et al. 1992). The observed variation did not however provide any information on
the genetic relatedness of the populations. High levels of inbreeding (F,, = 0.382)
and the combined effects of inbreeding and random genetic drift were apparent
in the populations (F;, = 0.447). These results reflect the primitive eusocial be-
havior of Bombus (Michener 1974, Brian 1983). They also suggest that the in-
dividual females sampled may have been sisters from the same colony, even
though an attempt was made to obtain a broad sample of individuals from “‘many”’
colonies.
The Neighbor-Joining tree provided no information on the origin of the Great
Basin mountain range populations with respect to the Rocky Mountains (Wasatch
Mountains). The unresolved tree results from the lack of variation in the popu-
lations sampled.
Table 5. Matrix of genetic similarity of Bombus appositus Cresson populations as measured by
Nei’s (1978) unbiased genetic identity.
Utah Nevada
Population White Pine Mill Creek Humboldt Ruby Independence
White Pine cares 0.998 0.997 0.997 0.998
Mill Creek eo 0.999 0.999 0.999
Humboldt paul: 0.999 0.999
Ruby seek ek 0.999
Independence 2k 2K 2K
1996 RUST & PORTER: GENETIC VARIATION IN BOMBUS APPOSITUS 207
0.497
0.0002 White Pine
—>
Humboldt
Ruby
Mill Creek
0.0002 -Y
Figure 1. Neighbor-Joining tree of five populations of Bombus appositus Cresson based on Nei’s
(1978) unbiased genetic identities. Numbers along branches are branch lengths (patristic values) and
numbers in parentheses are the analysis cycles. The tree is unresolved with basal branch lengths of
essentially 0.0 (0.0002).
Independence
ACKNOWLEDGMENT
We thank H. Britten, Biology, University of Nevada and A. Scholl, Zoologisch-
es Institut, Universitat Bern for reviewing the manuscript; T. Rust for figure prep-
aration; and P. Brussard and the Nevada Biodiversity Initiative for partial funding
of this research.
LITERATURE CITED
Brian, M. V. 1983. Social insects. Chapman and Hall, New York, New York.
Crespi, B. J. 1991. Heterozygosity in the haplodiploid Thysanoptera. Evolution, 45: 458-462.
Graur, D. 1985. Gene diversity in Hymenoptera. Evolution, 39: 190-199.
Hartl, D. L. & A. G. Clark. 1989. Principles of population genetics. Sinauer Associates, Sunderland,
Massachusetts.
Jin, L. & J. W. H. Ferguson. 1990. Neighbor-joining tree and UPGMA tree software. Univ. Texas
Healt. Sci. Cen., Houston, Texas.
May, B. 1992. Starch gel electrophoresis of allozymes. Jn A. R. Hoelzel (ed.). Molecular genetic
analysis of populations. Oxford University Press, New York, New York.
Michener, C. D. 1974. The social behavior of the bees. Harvard University Press, Cambridge, Mas-
sachusetts.
Mullen C. & R. W. Rust. 1994. Isozyme variability of acommercially-reared population of the bumble
bee, Bombus occidentalis Greene. BeeScience, 3: 124-127.
Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of
individuals. Genetics, 89: 583-590.
Owen, R. W. 1985. Difficulties with the interpretation of patterns of genetic variation in the eusocial
Hymenoptera. Evolution, 39: 201—205.
Owen, R. W. 1988. Selection at two sex-linked loci. Heredity, 60: 415-419.
Owen, R. W., L. J. Mydynski, L. Packer & D. B. McCorquodale. 1992. Allozyme variation in bumble
bees (Hymenoptera: Apidae). Biochem. Genet., 30: 443-453.
Packer, L. & R. E. Owen. 1989. Allozyme variation in Halictus rubicundus (Christ): A primitively
social halictine bee (Hymenoptera: Halictidae). Can. Entomol., 121: 1049-1057.
208 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Pamilo, P. & R. H. Crozier. 1982. Measuring genetic relatedness in natural population: methodology.
Theor. Pop. Biol. 21: 171-193.
Pamilo, P, S. Varvio-Aho & A. Pekkarinen. 1978. Low enzyme gene variability in Hymenoptera as
a consequence of haplodiploidy. Hereditas 88: 93-99.
Pamilo, P., S. Varvio-Aho & A. Pekkarinen. 1984. Genetic variation in bumblebees (Bombus, Psi-
thyrus) and putative sibling species of Bombus lucorum. Hereditas, 101: 245-251.
Saitou, N. & M. Nei. 1987. The Neighbor-joining method: A new method for reconstructing phylo-
genetic trees. Mol. Biol. Evol., 4: 406-425.
Scholl, A., E. Obrecht & R. E. Owen. 1990. The genetic relationship between Bombus moderatus
Cresson and the Bombus lucorum auct. species complex (Hymenoptera: Apidae). Can. J. Zoo.,
68: 2264-2268.
Sheppard, W. S. & S. L. Haydon. 1986. High levels of genetic variability in three male-haploid
species (Hymenoptera: Argidae, Tenthredinidae). Evolution, 40: 1350-1353.
Swofford, D. L. & R. B. Selander. 1981. BIOSYS-1: a FORTRAN program for the comprehensive
analysis of electrophoretic data in population genetics and evolution. J. Hered., 72: 281—283.
Thorp, R. W., D. S. Horning & L. L. Dunning. 1983. Bumble bees and cuckoo bumble bees of
California (Hymenoptera: Apidae). Bull. Calif. Insect Surv., 23: 1-79.
Wright, S. 1978. Evolution and the genetics of populations. Vol 4: Variability within and among
natural populations. Univ. Chicago Press, Chicago, Illinois.
Yik-Yuen, G., G. W. Otis, M. Mardan & S. G. Tan. 1991. Allozyme diversity in asian A pis. In Smith,
D. R. (ed.). Diversity in the genus Apis. Westview Press. Boulder, Colorado.
Received 19 Jan 1996; Accepted 4 Mar 1996.
PAN-PACIFIC ENTOMOLOGIST
72(4): 209-212, (1996)
ESTABLISHMENT OF LARINUS MINUTUS GYLLENHAL
(COLEOPTERA: CURCULIONIDAE) FOR BIOLOGICAL
CONTROL OF DIFFUSE AND SPOTTED KNAPWEED IN
THE WESTERN UNITED STATES
R. FE Lana,! J. M. Story,* AND G. L. PIPER?
‘United States Department of Agriculture, Animal and Plant Health Inspection
Service, Plant Protection and Quarantine,
Forestry Sciences Laboratory, Montana State University,
Bozeman, Montana 59717;
Western Agricultural Research Center, Montana State University,
580 Quast Lane, Corvallis, Montana 59828;
3Department of Entomology, Washington State University,
Pullman, Washington 99164
Abstract.—Larinus minutus Gyllenhal (Coleoptera: Curculionidae) is a capitulum-infesting nat-
ural enemy of diffuse and spotted knapweed (Centaurea diffusa Lamarck and C. maculosa
Lamarck) (Asteraceae). United States Department of Agriculture, Animal and Plant Health In-
spection Service, Plant Protection and Quarantine personnel and research collaborators released
13,791 adults from 1991 to 1993 for the biological control of these knapweeds. Weevil estab-
lishment was confirmed in Montana in 1992, and in Washington and Wyoming in 1993. Redis-
tribution began in 1994 from one site each in Montana, Washington, and Wyoming. As of 1994,
L. minutus had also been released in Minnesota, Nebraska, Oregon, and Utah for a total of seven
states and 24 counties.
Key Words.—Insecta, biological control, weed, Larinus, Centaurea
Diffuse and spotted knapweeds (Centaurea diffusa Lamarck and C. maculosa
Lamarck) (Asteraceae) are European plants accidentally introduced into North
America that have become serious weeds of rangelands, pastures, and waste areas.
Spotted knapweed, initially reported from Montana in 1927, now infests 1,912,181
ha in the state (Roché 1994b). In 1984, the estimated loss of forage in Montana
attributable to C. maculosa was $4.5 million (French & Lacey 1983). Washington,
in 1993, was estimated to have 851,700 ha of diffuse and spotted knapweed-
infested rangeland (Roché 1994a). Knapweeds are aggressive, invasive competi-
tors that rapidly displace desirable forage grasses and forbs, including native
plants. Diffuse knapweed is a biennial and spotted knapweed is a short-lived
perennial, and both species reproduce exclusively by seed. In order to reduce seed
production and spread, various European capitulum-feeding insects have been
introduced into North America for the classical biological control of these Cen-
taurea spp. (Piper & Rosenthal 1995, Story 1995). These insects include Metz-
neria paucipunctella Zeller (Lepidoptera: Gelechidae), Chaetorellia acrolophi
White and Marquardt, Terellia virens Loew, Urophora affinis (Frauenfeld), U.
quadrifasciata (Meigen) (Diptera: Tephritidae), Bangasternus fausti (Reitter), Lar-
inus obtusus Gyllenhal, and L. minutus Gyllenhal (Coleoptera: Curculionidae).
Larinus minutus feeds only on diffuse and spotted knapweed. Overwintered
adults begin emerging in mid-May and feed upon stems, foliage, flower buds, and
flowers (Groppe 1990). Females deposit eggs among the florets of opened capit-
ula. Upon hatching, larvae feed within the capitula upon the developing seeds.
210 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Table 1. Larinus minutus releases, recoveries and establishments, 1991-1993.
Year(s) No. oO.
State County released released recovered Establishment
Montana Gallatin 1991-1993 1693 4 Yes
Lewis & Clark 1991-1992 2592 55 Yes
Park 1992 pallies 0 No
Powell 1991 50 0 No
Sweet Grass 1992-1993 1572 6 Yes
Nebraska Holt 1992 200 0 Destroyed
Oregon Deschutes 1992 200 — Yes
Hood River 1992 200 — Yes
Washington Spokane 1991-1993 1045 210 Yes
Stevens 1991-1993 406 44 Yes
Whitman 1992-1993 400 180 Yes
Wyoming Natrona 1991 175 4 Yes
Teton 1992 1500 — Yes
—, Not recorded.
One larva can destroy all the seeds in a diffuse knapweed capitulum; one or more
larvae reduce seed production by 25 to 100% in the larger capitula of spotted
knapweed. Pupation occurs within the damaged capitula (Groppe 1990). Adults
emerge during late summer and feed upon the foliage of knapweed rosettes for a
short period of time before seeking overwintering sites in the soil or debris in the
vicinity of their host plants. The weevil has one generation per year. This paper
documents the establishment and redistribution of L. minutus in the western Unit-
ed States.
MATERIALS AND METHODS
Larinus minutus was approved for release in the United States on 26 June 1991
by the United States Department of Agriculture (USDA), Animal and Plant Health
Inspection Service (APHIS), Plant Protection and Quarantine (PPQ). Adult wee-
vils were collected in Europe (Greece and Romania) by personnel from the Com-
monwealth Agricultural Bureaux International Institute of Biological Control, De-
lémont, Switzerland. The beetles were received and held in quarantine at the
USDA-APHIS-PPQ Mission Biological Control Laboratory, Mission, Texas, prior
to field release.
Uncaged releases of L. minutus adults on diffuse and spotted knapweed were
made by cooperators in Montana, Nebraska, Oregon, Washington, and Wyoming
between 1991 and 1993 (Table 1).
To document establishment, sites were sampled the year following release. As-
sessments were conducted in early May, late June, or early July, and September
through October. The undersides of rosette leaves were examined for feeding
adults in early May. The adults were found feeding on the leaves and flower buds
of bolted plants in late June and early July. If no adults were observed during the
summer months, sweep samples were taken by walking in an expanding circle
from the release point to approximately 6 m out from the center. Visual and sweep
sampling were undertaken during mid-morning (1000 h) and late afternoon (1400
h) on warm, sunny days. The knapweed stands were also checked for the occur-
1996 LANG ET AL.: LARINUS MINUTUS ESTABLISHED 211
rence of adult emergence holes in late September through October as another
method to confirm establishment. The large emergence holes, positioned in the
centers of mature capitula, can readily be seen while walking through a knapweed
infestation.
Weevil establishment was considered to be achieved when adults were recov-
ered the year after the initial release. Collection of L. minutus adults for redistri-
bution from field insectary sites occurred when it was determined that 25% of
the capitula within 30 m of the release point contained emergence holes.
RESULTS AND DISCUSSION
In 1992, L. minutus larvae or adults from the 1991 releases were recovered
from two sites in Montana: East Gate in Lewis & Clark County (24 Jul, n = 11)
and Droulliard Fishing Access in Gallatin County (12 Aug, n = 4) (Table 1). No
other recoveries were reported from the 1991 releases. By 1994, Montana had
five established L. minutus field insectaries. In 1994, the East Gate release location
provided 2685 adults which were released in Montana [Gallatin (n = 1166) and
Sweet Grass (n = 84) County], Minnesota (Becker County) (7 = 400), and Ne-
braska (Pierce County) (7 = 1035). In 1993, Washington reported recoveries of
the weevil in three of four counties and from all four counties in 1994. Washington
redistributed 4195 adults from Whitman County in 1994 to Columbia, Franklin,
Grant, Okanogan, Stevens, Walla Walla, and Whitman Counties. Wyoming re-
covered L. minutus adults in 1993, and in 1994 was able to redistribute 562 adults
from Natrona County to two other sites within the county. Oregon reported re-
covery from Deschutes and Hood River Counties in 1994. The Nebraska (Holt
County) 1992 release site was destroyed in 1994, with no weevil recovery being
recorded (Table 1). A release of 375 L. minutus from Greece was made in Utah
(Weber County) in 1994.
Larinus minutus is well-established in Oregon, Montana, Washington, and Wy-
oming (Table 1). The Montana weevil populations increased rapidly despite an
unseasonably cold summer in 1993. By summer’s end in 1994, this beetle had
been released in 24 counties in seven states, with release material originating from
both European and domestic sources. It appears that collections from most field
insectaries will be possible within three years of the initial establishment. Popu-
lations of L. minutus now established in Oregon, Montana, Washington, and Wy-
oming should continue to expand and provide additional weevils for redistribution
purposes in these and other western states with diffuse and spotted knapweed
infestations.
LITERATURE CITED
French, R. A. & J. R. Lacey. 1983. Knapweed: its cause, effect and spread in Montana. Montana
Coop. Ext. Serv. Circ., 307.
Groppe, K. 1990. Larinus minutus Gyll. (Coleoptera: Curculionidae), a suitable candidate for the
biological control of diffuse and spotted knapweed in North America. CAB International In-
stitute of Biological Control, European Station, Delémont, Switzerland. Report.
Piper, G. L. & S. S. Rosenthal. 1995. Diffuse knapweed, Centaurea diffusa Lamarck (Asteraceae),
Chapter 64. pp. 237-241. In Nechols, J. R., L. A. Andres, J. W. Beardsley, R. D. Goeden &
C. G. Jackson (eds.). Biological control in the western United States: accomplishments and
benefits of regional research project W-84, 1964-1989. Univ. Calif. Div. Agric. Nat. Res. Publ.
3361. Oakland, California.
212 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Story, J. M. 1995. Spotted knapweed, Centaurea maculosa Lamarck (Asteraceae), Chapter 70. pp.
258-263. In Nechols, J. R., L. A. Andres, J. W. Beardsley, R. D. Goeden & C. G. Jackson
(eds.). Biological control in the western United States: accomplishments and benefits of regional
research project W-84, 1964-1989. Univ. Calif. Div. Agric. Nat. Res. Publ. 3361. Oakland,
California.
Roché, B. F, Jr. 1994a. Status of knapweeds in Washington. Wash. State Univ. Coop. Ext. Serv.
Knapweed Newsletter 8(1): 2—4.
Roché, B. E, Jr. 1994b. Some thoughts about weeds. Wash. State Univ. Coop. Ext. Serv. Knapweed
Newsletter 8(3): 1—2.
Received 23 Feb 1996; Accepted 11 Apr 1996.
PAN-PACIFIC ENTOMOLOGIST
72(4): 213-219, (1996)
PTEROSTICHUS BEETLES DOMINATE THE CARABID
ASSEMBLAGE IN AN UNSPRAYED ORCHARD IN
SONOMA COUNTY, CALIFORNIA
Eric W. RIDDICK! AND NICK J. MILLS
Laboratory of Biological Control, Department of Environmental Science,
Policy and Management, University of California, Berkeley, California 94720
Abstract.—The seasonal activity of predatory carabids on the soil surface was examined in an
unsprayed apple orchard in Sonoma County. Pterostichus beetles were the most active carabids
in this orchard, comprising from 71—83% of the total number of individuals trapped during two
consecutive growing seasons. Pterostichus (Dysidius) lustrans LeConte and Pterostichus (Hy-
pherpes) spp. were most active in June 1991 and June 1992, whereas Pterostichus (Poecilus)
cursitor LeConte were most active in July and August 1991 and in July 1992.
Key Words.—Coleoptera, Carabidae, Pterostichus, seasonal activity, orchards
Adults of the carabid genus Pterostichus are important predators of insect pests
in agroecosystems (Hagley & Allen 1988, Allen & Hagley 1990, Carcamo &
Spence 1994, Clark et al. 1994, Riddick & Mills 1994, Wallin & Ekbom 1994).
In apple orchards, especially those containing semidwarf trees, Pterostichus bee-
tles help suppress codling moth (Cydia pomonella (L.)) populations by attacking
fifth-instar larvae wandering on the soil surface prior to pupation (Riddick &
Mills 1994). Determining the seasonal activity of these carabids in relation to
when C. pomonella larvae are wandering on the ground is important. If the surface
activity of Pterostichus species coincide with the time that codling moth larvae
are seeking pupation sites, these carabids can contribute significantly to the sup-
pression of this pest. First generation C. pomonella larvae leave fruit during May
or June in orchards in California (Pickel et al. 1986), and thereafter become
vulnerable to predation by Pterostichus adults on the ground.
Pesticide sprays have been shown to alter the surface activity of adult carabids
in apple orchards. For example, Harpalus pensylvanicus DeGeer were signifi-
cantly more active in plots sprayed with. granulosis virus in early July, than in
plots sprayed with Bacillus thuringiensis Berliner (Dipel) plus oil, or controls (no-
spray); Chlaenius tricolor Dejean were significantly more active in plots sprayed
with oil alone, in early June, than in plots sprayed with granulosis virus, B.
thuringiensis plus oil, or controls (Riddick & Mills 1995). In unsprayed apple
orchards, an accurate assessment of the seasonal pattern of activity for Pterosti-
chus species can be made.
MATERIALS AND METHODS
The study site was an unsprayed apple orchard located in the coastal region of
northern California (Sonoma County), near Sebastopol, a major apple-growing
district. The orchard contained trees of Golden Delicious and Rome Beauty apple
varieties. This ~2 ha orchard was bordered by a woodlot, meadow, and residence
1 Current Address: Department of Entomology, 1300 Symons Hall, University of Maryland, College
Park, Maryland 20742.
214 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
on three sides. The fourth side bordered a paved roadway that separated it from
an ~20 ha sprayed, commercial orchard. The number of trees in this orchard was
not determined, but an estimate is that it contained ~20 tree rows, each with an
average of 20 trees. The soil was disked (as a means of retaining soil moisture)
in late April to early May, and the natural grasses and herbs that were uprooted
were left on the soil surface. Pesticides were not sprayed in this orchard between
April 1991 and September 1992.
The carabid assemblage was sampled during two consecutive seasons using
pitfall traps, the standard technique for sampling carabids (Greenslade 1964, Ri-
vard 1965, Luff 1975, Morrill 1975, Adis 1979, Baars 1979a, Halsall & Wratten
1988). Trap catch estimates the activity of adult carabids on the soil surface and
whole season trapping reflects the density of carabid populations (Ericson 1979,
Hokkanen & Holopainen 1986). Traps were plastic cups (473 ml), with a 9 cm
diameter opening, sunk into the ground with the rim flush with the soil surface.
Leaf litter within 20 cm of the perimeter of each trap was removed and the soil
smoothed to facilitate the movement of carabids around the traps (Greenslade
1964, Powell et al. 1985). (However, this procedure could have introduced sam-
pling bias against species that avoid crawling onto smoothed soil.) Traps were
filled to the one-quarter mark with a solution of water and liquid detergent, so
that captured beetles sank to the bottom of the trap. A preservative was not used
because it might alter the catch and sex ratio of the trapped species (Holopainen
1992).
Six pitfall traps were positioned, one tree apart, in a single row of trees at the
center of the orchard in 1991. Traps were in place for 6-10 consecutive days
during each of four sampling periods in 1991, 4-11 June, 3-9 July, 30 July—9
August, and 4—11 September. In 1992, sampling effort was increased, such that
five traps were positioned, one tree apart, in each of three rows of trees. The three
sample rows were equidistant within the orchard; five tree rows apart. Traps were
in place for three consecutive days during each of six sampling periods, 20—23
April, 11-14 May, 5-8 June, 7-10 July, 28-31 July, and 18-21 August. Samples
were collected on the last day of each sampling period. Trapped carabids were
sorted to species or species groups in the laboratory.
Voucher specimens are deposited at the Laboratory of Biological Control, Uni-
versity of California, Berkeley, and the Department of Entomology, University of
Maryland, College Park.
RESULTS
Adults of Pterostichus species were the most active carabids in the unsprayed
orchard, representing 71—83% of the total number of individuals trapped over the
two seasons. Adult Trechus, Amara, and Anisodactylus species were less active.
In 1991, a total of 511 carabids were captured on four collection dates (Table 1).
The species that represented 3% or more of the carabids include Pterostichus
(Dysidius) lustrans LeConte, Pterostichus (Hypherpes) spp., Pterostichus (Poe-
cilus) cursitor LeConte, Trechus obtusus Erichson, Amara spp., and Anisodactylus
californicus Dejean. Pterostichus (Hypherpes) spp. represents two species, Pter-
ostichus californicus (Dejean) and Pterostichus castanipes (Ménétriés) which are
morphologically similar, and not readily distinguishable at the time that this re-
search was undertaken. Subsequent identifications of pinned specimens (collected
1996 RIDDICK & MILLS: PTEROSTICHUS IN ORCHARDS 205
Table 1. Percentage and number of carabid beetles active during two consecutive growing seasons.
1991 collection 1992 collection
Carabid species — % [No] ~ % (No}
Pterostichus lustrans LeConte 35.62 [182] 44.04 [340]
Pterostichus cursitor LeConte 13.31 [68] 26.29 [203]
Pterostichus californicus (Dejean) &
Pterostichus castanipes Ménétriés 22.11 [113] 12.30 [95]
Trechus obtusus Erichson 10.76 [55] 3.11 [24]
Amara spp. (2+ species) 8.02 [41] 5.83 [45]
Anisodactylus californicus Dejean 4.89 [25] 4.40 [34]
Anisodactylus similis LeConte 2.93 [15] 0.39 [3]
Bradycellus spp. (2 species) 0.78 [4] 1.68 [13]
Calathus ruficollis Dejean 0.39 [2] 0.39 [3]
Dicheirus piceus Menetries 0.19 [1] 0.52 [4]
Loricera foveata LeConte 0.19 [1] 0.91 [7]
Microlestes sp. 0.19 [1] — —
Agonum fossigerum Dejean 0.19 [1] — —
Tanystoma maculicolle (Dejean) 0.19 [1] — —
Scaphinotus sp. — — 0.13 [1]
during the 1991 and 1992 seasons) indicated that P. californicus comprised ~91%
and P. castanipes comprised ~9% of the P. (Hypherpes) spp. group in this or-
chard. At least two species of Amara were included in the catch; however, I was
unable to identify them to species.
In the 1992 season, 772 carabids were collected on six collection dates (Table
1). The species that represented 3% or more of the carabids include: P. lustrans,
P. cursitor, P. (Hypherpes) spp., Amara spp., A. californicus, and T. obtusus.
The seasonal activity of Pterostichus beetles was determined in both seasons.
In 1991, P. lustrans adults were the most active carabids in early June and their
activity sharply declined through July and August (Fig. 1). Pterostichus cursitor
were less active, but their activity remained constant from early June to August.
Although P. (Hypherpes) spp. were most active in early June, their activity con-
tinued into early September. In 1992, adults of P. lustrans were again the most
active carabids early in the season from late April through June. Adults of P.
cursitor appeared more active in late April and early July, and their activity was
more variable than in 1991. Adults of P. (Hypherpes) spp. were again most active
in early June. All Pterostichus beetles were active on each collection date in the
1992 season.
DISCUSSION
Pterostichus species comprised from 71-83% of the carabid assemblage in the
unsprayed orchard. In a nearby sprayed, commercial orchard (which was located
91 m from the unsprayed one), Pterostichus species represented 45% of the total
carabids trapped in the 1991 season, but only 19% of the total in 1992 (Riddick
& Mills 1996). The greater percentage of trap captures of Pterostichus species in
the unsprayed orchard may, in part, result from a lack of insecticide applications
against codling moth and the presence of suitable alternative prey for these de-
cidedly carnivorous carabids to feed on.
216 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
file P. lustrans LJ P. cursitor P. californicus «
Number
11 Jun 9 Jul 9 Aug 11 Sept
Collection Dates - 1991 Season
Mean (£SEM) Number
ye)
4
M|
QO
Vat ev,
rm LO 4
O
gy
or
23 Apr 14May. 8Jun 10 Jul 31 Jul 21 Aug
Collection Dates - 1992 Season
Figure 1. Number of Pterostichus adults per trap per day in a single tree row in the 1991 season,
and the mean (SEM) number of Pterostichus adults per tree row per trap per day in the 1992 season.
Three tree rows were sampled in 1992. *, represents two morphologically similar species, P. califor-
nicus (~919% of the Hypherpes group in sample) and P. castanipes (~9%).
1996 RIDDICK & MILLS: PTEROSTICHUS IN ORCHARDS 217
The most active carabids in the sprayed orchard were Amara species; they
comprised 31% of the carabids in 1991 and 63% in 1992. Amara species were
significantly less active on at least one collection date, in both seasons, in plots
managed with conventional insecticides (Riddick 1993). The Amara species be-
came dominant in a sprayed orchard despite some individuals being affected by
the insecticides.
The presence of woodlot and meadow adjacent to the unsprayed orchard may
have contributed to the greater activity of Pterostichus beetles, as well. It is un-
clear why other species, such as Amara spp. did not experience increased activity,
but Amara are infrequently captured in woodlands, possibly due to the lack of
grass seeds, a preferred food source for adults (see Niemela et al. 1992). Exchange
of carabids and spiders between cultivated fields (or orchards) and semi-natural
habitats has been documented (Altieri & Schmidt 1986, Coombes & Sotherton
1986, Duelli 1990, Dennis & Fry 1992, Kajak & Lukasiewicz 1994). Pterostichus
beetles overwintering in the adjacent woodlot or meadow, might have dispersed
into the unsprayed orchard in the spring (perhaps in early April). In contrast,
movement from the woodlot or meadow into the sprayed, commercial orchard
would have been limited, because both natural areas were located on the opposite
side of the roadway. Roadways can become a barrier to adult carabids that dis-
perse primarily by crawling (Mader 1984, Mader et al. 1990).
Adult Pterostichus species were never seen flying at the margins of either
orchard. Both P. californicus and P. castanipes are flightless. Pterostichus lus-
trans and P. cursitor are fully-winged (macropterous) and probably disperse by
flight, to some extent, from overwintering sites.
The seasonal activity of the four Pterostichus species was similar in the un-
sprayed orchard. Pterostichus lustrans adults were the most active, especially in
June or July. But the reduced activity of P. lustrans in August in both orchards
may have resulted from inadequate moisture and high temperatures at the soil
surface. The climate in the Sebastopol region is Mediterranean, characterized by
hot, dry summers and mild, wet winters (Altieri & Schmidt 1986). The lack of
ideal conditions for activity may have influenced the adults to remain beneath the
soils. surface, late in the season.
The high June activity of P. lustrans adults suggests that they have the greatest
potential for searching for and consuming fifth-instar larvae of the codling moth
on the soil surface. First generation C. pomonella larvae drop from fruit and
search for pupation sites in early June (Pickel et al. 1986). The ability of these
carabids to locate the wandering larvae may depend on the hunger level of the
individuals. Hungry carabids may forage for prey more effectively than satiated
individuals (Baars 1979b, Mols 1987, Wallin & Ekbom 1994). The seasonal ac-
tivity and feeding behavior of P. lustrans larvae is unknown, but it is conceivable
that these predatory larvae could attack and kill C. pomonella \arvae that enter
the soil to pupate.
The conservation of Pterostichus populations in apple orchards is necessary if
these natural enemies are to become components of an integrated pest manage-
ment program. This current research suggests that the conservation of the more
predatory carabids in California orchards may involve a reduction in pesticide
usage. Maintaining natural areas (meadow, woodlot or hedgerows) in the imme-
diate vicinity of orchards, or non-crop plants within orchards, might help conserve
218 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
and enhance Pterostichus species populations during the season (see Altieri &
Schmidt 1985, Altieri 1991, Thomas et al. 1991, Lys & Nentwig 1992).
ACKNOWLEDGMENT
We thank David Kavanaugh (Department of Entomology, California Academy
of Sciences, San Francisco) for his assistance with some carabid identifications,
and Sue Blodgett (former IPM Advisor, North Coast Counties (University of
California Cooperative Extension, Santa Rosa, California)) for directing us to the
unsprayed orchard.
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Received 20 Nov 1995; Accepted 16 Apr 1996.
PAN-PACIFIC ENTOMOLOGIST
- 72(4): 220-226, (1996)
ESTABLISHMENT OF THE SQUASH BUG PARASITOID,
TRICHOPODA PENNIPES FABR. (DIPTERA: TACHNIDAE),
IN NORTHERN CALIFORNIA
C. H. PIcKETT!, S. E. SCHOENIG!, AND M. P.. HOFFMANN?
1Biological Control Program California Department of Food & Agriculture
3288 Meadowview Rd., Sacramento, California 95832;
Department of Entomology, Cornell University, Ithaca, New York 14853
Abstract.—The squash bug, Anasa tristis (De Geer) (Hemiptera: Coreidae), is an important pest
of squash and pumpkin plants in California. Its pest status is partially due to a lack of specific
nymphal or adult natural enemies in California. A nymphal-adult parasitic fly, Trichopoda pen-
nipes Fabr. (Diptera: Tachnidae), commonly associated with squash bugs in eastern United States,
was imported and released beginning in 1992 at several locations in Yolo, Solano, and Sacra-
mento counties in northern California. This fly has successfully overwintered at four locations
and has been recovered at two locations three years after initial releases.
Key Words.—Insecta, Trichopoda pennipes, biological control, squash bug, Anasa tristis
The squash bug, Anasa tristis (De Geer) (Hemiptera: Coreidae), is distributed
from South America north to Canada, where it is one of the most important pests
of cucurbits (e.g., squash, melons, cucumbers) (Beard 1940, Johannsen 1957,
Nechols 1987). It is a frequent and particularly serious problem in California for
producers and home growers of organically grown squash (personal observation,
Flint 1990). Cultivars derived from Pepo maxima L. such as kabocha, red kuri,
and hubbard cannot be grown in some areas without pesticides. The squash bug
attacks all stages of the plant with the seedling stage being most susceptible to
damage (Beard 1940). It reportedly injects a toxin during feeding and may trans-
mit plant diseases (Eichmann 1945). During 1993 pesticide usage was reported
on 8300 acres of California squash and pumpkins, of which some portion was
used for control of squash bug (State of California 1993).
Trichopoda pennipes Fabr. is a nymphal-adult parasitoid that attacks squash
bug infesting cucurbits in northeastern United States. It is the most widely dis-
tributed member of Trichopoda, found throughout North America south to Ar-
gentina and from the West Indies to California (Beard 1940). In the United States
it reportedly has three geographically isolated strains (Dietrich & van den Bosch
1957). Only the strain from northcentral and northeastern United States attacks
the squash bug. In Connecticut, it has been reported parasitizing up to 84% of
overwintering adults (Beard 1940). A second strain occurs in the southern United
States where it attacks pentatomids (stink bugs), and a third exists in California
where it attacks the bordered plant bug, Euryopthalmus cinctus californicus Van
Duzee, a largid which is sometimes found in cucurbits (van Driesche 1970, Die-
trich & van den Bosch 1957). Dietrich & van den Bosch (1957). tried unsuccess-
fully to cross flies from southern California with those from Connecticut. They
also observed that flies from Connecticut did not attack the bordered plant bug.
Trichopoda pennipes collected in southern California readily attacked the bor-
dered plant bug but did not attack squash bugs. It is possible that the different
1996 PICKETT ET AL.: TRICHOPODA PENNIPES ESTABLISHED papal
strains are sibling species, morphologically identical but incapable of interbreed-
ing and biologically distinct.
The absence of squash bug natural enemies in the Pacific Northwest prompted
the state of Washington to import 7. pennipes from northeastern United States
during 1943-1944 (Clausen 1978). Observations in 1951 found ca. 50% parasit-
ism over a large area of the state in late summer and autumn. Dietrick & van den
Bosch (1957) imported T. pennipes from Connecticut in the 1950s but failed to
established permanent populations in southern California. The lack of squash
plants the following spring near their release site depleted the local squash bug
population, and thus hosts for the parasitoid (Dietrick personal communication).
The successful importation of 7. pennipes in Washington and limited efforts at
establishment during the 50s in California warranted a second effort at importing
this fly into this state. Furthermore, our surveys have recovered only small num-
bers of the egg parasitoid Ooencyrtus californicus Girault (Hymenoptera: Sce-
lionidae) in late season when densities of squash bug are at their highest levels.
Specific natural enemies of the nymph or adult squash bug stage have never been
observed in our surveys or by others (Dietrick & van den Bosch 1957). We report
on the importation, rearing, release, initial distribution, and colonization of T.
pennipes for control of squash bug.
MATERIALS AND METHODS
Field collection and shipment of parasitoids ——From June through August,
1992 and 1993, parasitized bugs were collected by one of the authors (MPH)
from plantings of cucurbits (primarily zucchini) from four organic farms in Schuy-
ler, Cayuga, and Seneca Counties of New York, one to eight collections per farm
per summer. Bugs were considered parasitized if found with one or more tachinid
egg(s) attached to their body (Fig. 1). The bugs were placed in 3.8 liter paper
cartons, ca. 40 bugs per carton, with screen tops and with fresh sections of cu-
cumbers or squash every two days. Environmental conditions were ca. 21° C
(room temperature) and natural day length. Parasitoids were sent to the California
Department of Food & Agriculture’s Biological Control Program facility in Sac-
ramento as puparia or as larvae inside of adult bugs, using overnight express mail.
Rearing and release of parasitoids—tIn 1992 and 1993, 335 and 151 adult
flies, respectively, emerged and were either directly released into squash fields or,
under laboratory conditions, used to parasitize locally collected bugs that were
subsequently released. These bugs were held inside standard sleeved cages
(DeBach 1964) in rooms set at 20 to 27° C (14L:10D) and fed zucchini squash.
No evidence of pathogens or hyperparasitoids were observed among the imported
flies and bugs. In 1992, parasitoids were released at three locations in northern
California with high squash bug populations: an organic private home garden in
Davis (Yolo Co.), and two organic farms, one near Guinda (Yolo Co.), and a
second near Winters (Solano Co.). None of these growers used insecticides. A
total of 243 adult flies and 947 parasitized adult bugs were released among these
three locations. In late summer 1993, 2440 parasitized bugs were released at the
Student Experimental Farm of the University of California (UC), Davis and an-
other 260 at an organic farm in Sacramento County.
In 1994 and 1995 an outdoor insectary or nursery site was established at the
UC Davis Student Experimental Farm to provide a large number of squash bugs
222 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Figure 1. TJ. pennipes eggs on squash bug: (A) close-up view of single egg identified by arrow
(photo by Jack Kelly Clark), and (B) cluster of eggs.
1996 PICKETT ET AL.: TRICHOPODA PENNIPES ESTABLISHED 223
Yolo Co.
LEGEND
& Released and
established
O Sacramento Co!
© Released and not yet
established
Winters
Recovery away ,
+ from release site Sacramento
MAP LOCATION
Figure 2. Release, recovery, and establishment locations for T. pennipes. Establishment implies
T. pennipes has been recovered for at least one year since its release.
as hosts for overwintering flies from the 1993 releases. Crops planted at the
Student Experimental Farm tolerate pest populations far above the economic
threshold of commercial growers. A mixture of zucchini and kabocha squash were
planted to a 0.8 ha site on three dates beginning in late spring. Twenty thousand
squash bugs were collected from surrounding farms and released to augment the
local population. Every 5th row in the field contained a strip of annual buckwheat
to provide with-in field nectar sources for T. pennipes. Honey and sugar water
have been shown to increase the longevity and fecundity of 7. pennipes in the
lab (Shahjahan 1968), and during 1994 many flies were observed feeding atop
the buckwheat flower heads. In 1994 and 1995 ca. 2100 flies were transported
from this site and released at new release sites, including a former site located in
Guinda.
Recovery.—Years following releases, adult squash bugs were collected from
squash plants located within several hundred meters of original release sites. Adult
bugs were collected two to three times at each location during summer months.
Ten to twenty adult 7. pennipes per site per summer were reared from field
collected bugs with fly eggs (Fig. 1) to confirm identity of the parasitoid.
RESULTS AND DISCUSSION
In 1993, T. pennipes was recovered at all three 1992 release sites (Fig. 2).
However, we were unable to recover parasitoids at the Guinda site in late summer.
We recovered parasitized squash bugs from the UC Davis site in 1994, 1995, and
1996, representing a fourth site in which the parasitoids successfully overwin-
224 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
tered. In 1995 we observed squash bugs with tachinid eggs in Winters (Everything
Under the Sun Farm), and the following spring we recovered the same at Terra
Firma showing that the fly population persisted at these two sites through three
winters, producing multiple generations. The same parasitoid in northeastern Unit-
ed States produces three generations during summer months while the squash bug
produces one (Beard 1940). Populations of introduced natural enemies that have
persisted for at least three generations or three years are most likely permanently
established (DeBach 1964). Trichopoda pennipes dispersed long distances from
release sites. In 1994, T. pennipes was found 8.0 km from the original release
site in the Winters area (Terra Firma Farm) and in a residential garden 2.5 km
distant from the UC Davis student farm insect nursery site.
The percentage of bugs with parasitoid eggs varied from 1.3 to 92.2 at release
sites (Table 1) and no other species of Tachinidae were reared from these indi-
viduals. These values underestimate the full potential of this parasitoid because
we were adding unparasitized bugs and/or removing parasitized ones while these
data were being recorded. Furthermore, sufficient generations have not yet passed
for this parasitoid to have become established over a large area.
Releases of T. pennipes have persisted for three years at two locations in north-
ern California and the parasitoids have dispersed up to 8.0 km from original
release sites. This is the first specific natural enemy successfully established
against the squash bug in California. Observations in New York and Washington
suggest that this parasitoid could become widely established and have a major
impact on squash bug. Although squash bug can be considered a problem to some
growers of cucurbits in New York, most of the bugs collected there for shipment
to California were parasitized. The squash bug in Washington was rated as only
a minor pest in the mid 1970s (Clausen 1978). Plantings of tachinid insectary
plants in and around fields of squash could increase the impact of flies on squash
bug (Pickett & Bugg in press). For organic farmers of specialty crops in Califor-
nia, the presence of T. pennipes may help reduce control costs and allow for the
production of highly susceptible varieties of squash that command a higher profit
margin. Similarly, home gardeners may not have to use pesticides or other control
measures for the squash bug helping to reduce widespread urban use of chemical
pesticides for this pest.
ACKNOWLEDGMENT
We thank Mark van Horn of the UC Davis Student Experimental Farm for
providing space and services for the cultivation of our T. pennipes nursery site,
and Full Belly Farm, Sky High Farm, Everything Under the Sun Farms, Marv
Kinsey, Francis and Jan Thompson, Brook and Jennifer Murphy for providing
release sites or squash bugs, and the organic farmers in New York for use of their
farms to collect squash bugs. We are grateful to anonymous reviewers for their
advice and review of this manuscript. The following individuals participated in
the rearing, establishment or data collection of this project: Giang Lam, Jorge
Lopez, Pauline Pao, Caroline Garcia, Dion Onizuka, Tunya Morizawa, Jason Pal-
omares, Debra Kress and Robin Wall. Robert L. Bugg of the UC Sustainable
Agriculture Research and Education Program brought to our attention the potential
importance of Trichopoda pennipes to organic growers in California.
Table 1. The number and percentage of adult squash bugs sampled with one or more parasitoid eggs attached to their bodies, 1993-1996.
Total number of parasitoids
Years following released (as immatures Total number of Number with Percentage of adults
release Site location (year recovered) or adults)! adult bugs examined parasitoid eggs with parasitoid eggs
One Davis—UCD Student Farm (1994) 2440 1300 205 15.8%
Davis—private homeowner (1993) 192 100 20 20.0%
Davis—Village Homes Community Garden (1994) immigrated 141 130 92.2%
Winters—Everything Under the Sun Farm (1993) 564 300 60 20.0%
Winters—Terra Firma Farm (1994) immigrated 906 350 38.6%
Guinda—Full Belly Farm (1993) 432 1350 18 1.3%
Guinda—Full Belly Farm (1996) 4434 68 14 20.5%
Two Davis—UCD Student Farm (1996) 2000 683 516 75.5%
Winters—Terra Firma Farm (1995) resident 2622 1476 56.3%
Three Winters—Everything Under the Sun (1995) resident 30 15 50.0%
Winters—Terra Firma Farm (1996) resident 65 ol 78.5%
'“Gmmigrated’’ implies that recovered parasitoids had dispersed to a site, and “‘resident’’ implies that no additional parasitoids had been released the previous
year(s).
9661
CHHSTTAV.LSH SYdINN4d VOOdOHDIEL * TW LA LLAMOId
SCC
226 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
LITERATURE CITED
Beard, R. L. 1940. The Biology of Anasa tristis DeGeer, with Particular Reference to the Tachinid
Parasite, Trichopoda pennipes Fabr. Connecticut Agricultural Experiment Station, New Haven.
Bull., 440.
Clausen, C. P. 1978. Introduced Parasites and Predators of Arthropod Pests and Weeds: a World
Review. Agriculture Handbook No. 480. United States Department of Agriculture, Agricultural
Research Service.
DeBach, P. 1964. Biological control of insect pests and weeds. Reinhold Publishing Corporation, NY.
Dietrick, E J. & R. van den Bosch. 1957. Insectary propagation of the squash bug and its parasite
Trichopoda pennipes Fabr. J. Econ. Entomol., 50: 627-629.
Eichmann, R. D. 1945. Squash bug depredations in Washington. J. Econ. Entomol., 38: 110-112.
Flint, M. L. 1990. Pests of the Garden and Small Farm: A Grower’s Guide to Using Less Pesticide.
Statewide Integrated Pest Management Project, University of California. Publication 3332.
Johannsen, C. A. 1957. History of biological control of insects in Washington. Northwest. Sci., 31:
57-92..
Nechols, J. R. 1987. Voltinism, seasonal reproduction, and diapause in the squash bug (Heteroptera:
Coreidae) in Kansas. Environ. Entomol., 16:.269—273.
Pickett, C. H. & R. L. Bugg. (in press). Enhancing Biological Control: Habitat Management to
Promote Natural Enemies of Agricultural Pests. AgAccess, Davis, California.
Shahjahan, M. 1968. Effect of diet on the longevity and fecundity of the adults of the tachinid parasite
Tricho poda pennipes pilipes. J. Econ. Entomol., 61: 1102-1103.
State of California. 1993. Pesticide Use Report, California Department of Pesticide Regulation.
van Driesche, R. 1970. Aspects of the Biology of the Common Bordered Plant Bug Largus cinctus
H.-S. (Hemiptera: Heteroptera: Largidae) and its Parasite Trichopoda pennipes (FE) (Diptera:
Tachinidae). Senior Thesis, Oregon State University.
Received 14 Feb 1996; Accepted 17 May 1996.
PAN-PACIFIC ENTOMOLOGIST
72(4): 227-234, (1996)
THERMAL-SHOCK TOLERANCE OF THREE SPECIES OF
AQUATIC INSECTS IN A NORTHERN CALIFORNIA,
GEOTHERMALLY INFLUENCED STREAM
JOHN R. Woop!, Eric P- MCELRAVY, AND VINCENT H. RESH
Division of Entomology and Parasitology, University of California,
Berkeley, California 94720
Abstract.—This study assessed the response of aquatic insects to short-term thermal shocks, such
as those encountered upon drifting into heated areas of geothermally influenced streams. The
upper incipient lethal temperature (UILT) thresholds were determined for three common species
in Big Sulphur Creek, Sonoma County, California, and the effect of acclimation temperature on
the UILT was determined for two caddisflies, both of which overwinter in the stream as larvae.
When acclimated at 28° C, the LT,, of Centroptilum convexum (Ide) ranged from 36.3° to
38.5°, that of Gumaga nigricula (McLachlan) from 37.8° to 41.0°, and that of Helicopsyche
borealis (Hagen) from 38.5° to 41.4°. When acclimated at 5.5°, the LT.) for G. nigricula ranged
from 35.7° to 38.9° and that of H. borealis from 36.6° to 41.1°. Helicopsyche borealis has the
highest UILT of any caddisfly, 39.0° for 30 min. The most important factors determining the
UILTs in these insects are acclimation temperature, shock duration, and the maximum temper-
ature of exposure.
Key Words.—Insecta, Trichoptera, Ephemeroptera, thermal-shock, geothermal, bioassay, thermal
tolerance
Aquatic insects inhabiting streams in geothermal areas are often subject to high
water temperatures. Insects that drift in these streams may encounter elevated
temperatures for brief periods, whereas those in benthic communities downstream
from thermal inputs (e.g., hot springs) generally experience longer exposures,
particularly during periods of reduced streamflow. The severity of stress on an
insect in such conditions depends upon several variables, including: 1) duration
and magnitude of temperature change; 2) the maximum temperature experienced;
3) acclimation temperature; and 4) body size, life stage, and physiological con-
dition of the organism (Hutchison 1976).
Most studies of thermal tolerance of aquatic insects have examined either short-
term (i.e., exposures of up to several days) or chronic (i.e., exposure throughout
the immature stages) effects of heated discharges. Short-term effects are typically
determined by a standard bioassay using either 24 to 96 h exposure times (e.g.,
Nebeker & Lemke 1968, Gaufin & Hern 1971, deKozlowski & Bunting 1981) or
the Critical Thermal Maximum (CTM) procedure (e.g., Moulton et al. 1993). For
CTM the animals are heated at a constant rate (usually 0.5° C per min) until
locomotory activities become disorganized (Ernst et al. 1984). Chronic effects
have been studied using a number of methods (e.g., Cairns 1976, Gartman &
Lake 1979). The effect of rapid immersion, called short-term thermal shock, has
received much less attention (Sherberger et al. 1977, Salmela & Anderson 1978).
A short-term bioassay is more appropriate than the 24 or 96 h or CIM test in
some cases, such as determining the tolerance of insects drifting into heated dis-
charges. Exposure times during entrainment in power plant cooling systems, for
' Present Address: The King’s University College, 9125 50th St., Edmonton, Alberta, Canada T6B
2H3.
228 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
example, seldom exceed 1 h (Schubel et al. 1978). These authors argue that when
exposure times are brief, rapid exposure to test temperatures is better than the
gradual increases of the critical thermal maximum (CTM) method.
Naturally heated waters present a unique opportunity for investigating the in-
fluence of thermal gradients on structural and functional aspects of benthic com-
munities. Consequently, biological communities in geothermal waters have been
well studied (e.g., Lamberti & Resh 1985, Resh & Barnby 1987 and references
therein). The potential that such waters offer for field experiments has been noted
by Brock (1970, 1975) but only a few investigators have used these habitats for
this purpose (e.g., Pritchard & Pelchat 1977, Schott & Brusven 1980, Lamberti
& Resh 1983). Although lethal temperature limits have been established for some
aquatic insects exposed to power plant discharges (e.g., Garten & Gentry 1976,
Tennessen & Miller 1983), little is known about thermal tolerance of insects that
typically occur in geothermal waters.
This study assessed the response of three stream insects to short-term thermal
shocks in the geothermal waters of Big Sulphur Creek, The Geysers, Sonoma
County, California. Specifically we ask: 1) What are the Upper Incipient Lethal
Temperature (UILT) thresholds for these insects? 2) What effect does acclimation
temperature have on the UILT? and 3) Does the magnitude of the temperature
change (AT, i.e., the difference between the acclimation temperature and the low-
est LT,, observed) during a thermal shock alter the UILT?
MATERIALS AND METHODS
Study area.—We conducted this research in Big Sulphur Creek (BSC), Sonoma
County, California, USA (38° 46’ N, 122° 45’ W, elevation 680 m), a third-order
stream that flows northwesterly through The Geysers Known Geothermal Re-
sources Area. The Geysers is the largest geothermal electric power-producing
facility in the world, but natural surface expressions are confined to hot springs
and a few steam vents. Our experiments were conducted approximately 250 m
below the confluence of BSC and its tributary Little Geysers Creek (LGC). The
lower portion of LGC is heated by a series of hot springs. The waters of LGC
cool rapidly when they mix into BSC, which is not affected by geothermal inputs
upstream of LGC. The thermal influence of LGC is not detectable after the first
200 m of mixing in BSC (Lamberti & Resh 1985). For a more detailed description
of this site including water chemistry, thermal, and geological conditions see
Lamberti & Resh (1983), McColl et al. (1978), McMillan (1985) and McElravy
et al. (1989).
Thermal-shock bioassays were conducted in BSC on 17 Jul 1982 (28° C ac-
climation temperature) and 14 Jan 1983 (5.5° C acclimation temperature). Im-
mature stages of three species of aquatic insects common in BSC, the caddisflies
Helicopsyche borealis (Hagen) and Gumaga nigricula (McLachlan) and the may-
fly Centroptilum convexum (Ide) were tested in July; C. convexum, however, was
not present in sufficient numbers for winter testing.
Design of heat shock experiment.—All tests were conducted at streamside,
eliminating the need for the 1-3 week pre-test holding period usually required in
laboratory-based bioassays, and reducing many non-thermal stresses (e.g., han-
dling, transportation). Additionally, this procedure allowed the test-cell environ-
1996 WOOD ET AL.: THERMAL SHOCK TOLERANCE 229
Se Ses
. Ss aoe .
eB ay.
Figure 1. Floating racks used to hold test cells and monitor effects of thermal-shock on stream
macroinvertebrates in Big Sulphur Creek, The Geysers, Sonoma Co., California.
ment to fluctuate with ambient conditions during holding periods and provided a
more realistic bioassay.
Test insects were collected at the study site and placed in groups of ten into
each of 42 PVC test cells (5 cm diameter with screened bottoms) supported in
the stream by a floating platform. Test cells were arranged in a 6 X 7 time-
temperature matrix described below. Three replicates of cells were prepared and
supported in BSC by a floating platform (Fig. 1) for <12 h prior to testing. For
each species a total of 1260 insects were tested in the three replicate experiments.
Seven controlled-temperature water baths in portable ice chests (20 * 50 X 20
cm) were set up next to the stream. Submersible aquarium heaters were used in
each bath to maintain constant temperature, and an air pump provided aeration
and mixing. Water temperatures were monitored manually and fluctuations in the
baths were kept to <0.5° C by hand additions of warm water when necessary.
Results of preliminary tests were used to establish a site-specific time-temperature
matrix that would bracket from 0 to 100% mortality. In setting time levels for
this matrix, we considered the minimum possible drift times (determined with a
dye tracer) through thermal portions of LGC to the confluence with the cooler
waters of BSC. The six test-temperature baths were 33°, 36°, 39°, 41°, 43° and
45° C; the control bath was held at ambient stream temperature. Six exposure
times were used (5, 10, 15, 20, 30, and 60 min) at each temperature.
Experimental procedure.—The thermal shocks were administered by quickly
transferring the test cells from the platform in BSC to the pre-heated water baths
on shore, and immediately immersing them. After the appropriate time interval
230 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
the cells were removed from the bath, returned to the platform, and percent mor-
tality determined after 24 h: Individuals were considered dead when they failed
to respond to repeated probing. The median resistance temperature (LT.,) for each
exposure time was determined graphically following the method of Litchfield &
Wilcoxon (1949). The upper incipient lethal temperature (UILT) was computed
for each time-temperature combination from the replicate LT;, values (Hutchison
1976, Schubel et al. 1978). Exposure times (min) were log-transformed [Log,,
(X; + 1)] to correct for non-linearity (Zar 1974). Simple linear regression of
median LT’, temperature against exposure time was used to determine the UILT
equations. Two-way ANOVA (Zar 1974) was used to compare the median LT..’s
among species and acclimation temperatures.
RESULTS AND DISCUSSION
Helico psyche borealis.—When acclimated at 28° C, this caddisfly has the high-
est 1 h resistance to thermal shock among the three species tested (LT;), = 38.5°
C). It endured a 1 h exposure at 36° C in both summer and winter with no
mortality. The UILT equations of H. borealis for July and January were signifi-
cantly different (ANOVA, P < 0.0005). The 5 and 60 min LT,,. temperatures
estimated by the UILT equations ranged from 41.4 to 38.5° C in summer and 41.1
to 36.6° C in winter (Fig. 2). This species’ resistance to thermal shock is slightly
lower in winter (1—2° C) than in summer, as indicated by the greater negative
slope of the winter UILT equation, especially at the longer durations tested. Pre-
vious reports have set the maximum thermal tolerance for H. borealis at 34° C,
but these were based on field observations (Wiggins 1977), not bioassays. If the
exposure time is limited to less than 30 min, H. borealis can survive temperatures
as high as 39° C. This is a higher temperature tolerance than that reported for
any other caddisfly (Moulton et al. 1993). It is not surprising then that H. borealis
is widespread across North America (Resh et al. 1984) and that it tolerates rela-
tively low oxygen levels (Williams et al. 1983).
The AT experienced by H. borealis during these tests was 9.7° C in summer
and more than three times as large (31.4° C) in winter. Thus, the total temperature
change experienced by this species appears to have little effect on its survival
threshold.
Gumaga nigricula.—When acclimated at 28° C, this caddisfly had a 1 h LT,
of 37.8° C. It was able to survive. a temperature of 36° C for 1 h during the
summer with no mortality, but could not do so in winter. Gumaga nigricula has
a lower UILT during winter than in summer (ANOVA, P < 0.0005). The 5 and
60 min LT, temperatures from the UILT equations ranged from 41 to 37.8° C in
summer and 38.9 to 35.7° C in winter (Fig. 3). The AT in summer was 8.9° C
and it was 29.3° C in winter.
Centroptilum convexum.—The 1h LT, for C. convexum was 36.3° C. However,
this mayfly had some mortality at all test conditions, compared with no mortality
at 36° C for the two caddisflies when acclimated to 28° C. The summer UILT for
this species was significantly different (ANOVA, P < 0.01) from either caddisfly,
indicating that this mayfly was less tolerant of high temperatures for all durations
tested. The LT, temperature for 5 and 60 min ranged from 38.5 to 36.3° C in
summer. Mayflies have consistently been shown to be less tolerant of temperature
1996 WOOD ET AL.: THERMAL SHOCK TOLERANCE 231
HELICOPSYCHE BOREALIS
é
fs
2
|
5 10 15-20 30 60
EXPOSURE TIME (MIN)
GUMAGA NIGRICULA
a
a
|
5 10 15 20 30 60
EXPOSURE TIME (MIN)
CENTROPTILUM CONVEXUM
2
|
5 °10 15 20 30 60
EXPOSURE TIME (MIN)
Figure 2-4. LT, thermal-shock values and the associated UILT regression lines at two acclimation
temperatures. Designations, summer (28° C) —-——@- ——@- — -; winter (5.5° C) { }}—_[_ }—. Fig-
ure 2. Helicopsyche borealis. Summer UILT equation is Y = —2.87x + 43.65; r? = 0.84, P < 0.0001.
Winter UILT equation is Y = —4.47x + 44.59; r* = 0.88, P < 0.0001. Figure 3. Gumaga nigricula.
Summer UILT equation is Y = —3.24x + 43.53; 7° = 0.76, P < 0.0001. Winter UILT equation is Y
= —3.2x + 41.36; r? = 0.62, P = 0.0001. Figure 4: Centroptilum convexum. Summer UILT equation
is Y = —2.11x + 40.1; 7? = 0.61, P = 0.0001.
232 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
increases than caddisflies (e.g., Gaufin & Hern 1971, deKozlowski & Bunting
1981, Tennessen & Miller 1983).
The UILT for aquatic insects can be expected to vary with respect to accli-
mation temperature in three ways (Precht et al. 1973): 1) UILT may be unaffected
by acclimation temperature and show no variation; 2) UILT may be positively
related to acclimation temperature, thus increasing and decreasing with seasonal
temperature fluctuations (this is the most commonly observed relationship in
aquatic organisms); or 3) UILT may be negatively related to the acclimation
temperature. We found that the UILTs of H. borealis and G. nigricula are lower
in the winter than in the summer and that this seasonal affect is greater for H.
borealis. This diminished thermal tolerance may be due to the physiological state
of this caddisfly. It overwinters in a quiescent state, attached at the base of boul-
ders (Williams et al. 1983, Resh et al. 1984b).
Moulton et al. (1993) found a similar positive correlation with acclimation
temperature in Hydropsyche simulans Ross, Ceratopsyche morosa (Hagen), Chi-
marra obscura (Walker), and Chimarra aterrima Hagen. However, their results
and ours contrast with those for two other caddisflies, Hydropsyche sp. (Sherber-
ger et al. 1977) and Brachycentrus americanus (Banks) (Salmela & Anderson
1978), both of which were unaffected by acclimation temperature. The differences
may be due, in part, to variations in the experimental protocols. Salmela & An-
derson (1978) used fewer time-temperature combinations than in our matrix. Sher-
berger et al. (1977) lowered the post-shock temperature slowly, whereas we used
an instantaneous decrease. We held the insects in the field for a period of only
24 h for post-shock evaluation.
During this study the maximum temperature at the study site in BSC was 30.5°
C. In summer, the aquatic insects in this portion of the stream are living near
their UILT and a relatively small increase in stream temperature (i.e., a AT of 6—
9° C) would exceed the UILT determined for all three test species. This was
illustrated in a related study of the thermally influenced portion of LGC upstream
from our study site. Resh et al. (1984a) found that very large numbers of ma-
croinvertebrates (primarily Chironomidae and Oligochaeta) occurred in the after-
noon drift. These animals were probably induced to drift catastrophically as the
substrate temperature exceeded their UILTs. In winter, there is a small reduction
in the UILT for the two caddisflies. However, since ambient water temperatures
are low, and both insects can withstand a AT > 30° C, these species would thus
be less susceptible to the effect of thermal discharges at this time of year.
An insect drifting into a thermal hot spring discharge would not be killed by
rapid heating, provided the temperature did not exceed its UILT. We conclude, as
did Sherberger et al. (1977), that the magnitude of the thermal shock is not con-
sequential in inducing mortality unless it approaches the UILT. The most impor-
tant factors determining the UILTs in H. borealis, G. nigricula, and C. convexum
appear to be acclimation temperature, shock duration, and the maximum temper-
ature of exposure.
ACKNOWLEDGMENT
We thank R. W. Flowers, Florida Agricultural and Mechanical University, for
confirming the identification of Centroptilum convexum (Ide). The research lead-
ing to this report was supported by the Office of Water Research and Technology,
1996 WOOD ET AL.: THERMAL SHOCK TOLERANCE 233
USDI, under the Annual Cooperative Program of Public Law 95-467, and by the
University of California Water Resources Center, as part of Office of Water Re-
search and Technology Project A-084-CAL and Water Resources Center Project
UCAL-WRC-W-612.
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Ernst, M. R., T. L. Beitnger & K. W. Stewart. 1984. Critical thermal maxima of nymphs of three
Plecoptera species from an Ozark foothill stream. Freshwat. Invertebr. Biol., 3: 80-85.
Garten, C. T, Jr. & J. B. Gentry. 1976. Thermal tolerance of dragonfly nymphs. II. Comparison of
nymphs from control and thermally altered environments. Physiol. Zool., 49: 206-213.
Gartman, D. K. & R. W. Lake. 1979. The effect of a thermal discharge on the benthos of a Virginia
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Gaufin, A. R. & S. Hern. 1971. Laboratory studies on tolerance of aquatic insects to heated waters.
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Hutchison, V. H. 1976. Factors influencing thermal tolerances of individual organisms. pp. 10—26. In
G. W. Esch & R. W. McFarlane (eds.). Thermal Ecology II. ERDA Symposium series, CONE
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Lamberti, G. A. & V. H. Resh. 1983. Geothermal effects on stream benthos: separate influences of
thermal and chemical components on periphyton and macroinvertebrates. Can. J. Fish. Aquat.
Sci., 40: 1995-2009.
Lamberti, G. A. & V. H. Resh. 1985. Distribution of benthic algae and macroinvertebrates along a
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Litchfield, J. T. Jr. & FE Wilcoxon. 1949. A simplified method of evaluating dose-effect experiments.
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McMillan, L. E. (ed.). 1985. Geysers-Calistoga KGRA-ARM program 1982-1983 annual report. 2
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Received 24 Jun 1994; Accepted 23 Apr 1996.
PAN-PACIFIC ENTOMOLOGIST
72(4): 235-237, (1996)
Scientific Note
EFFECTS OF COPULATION FREQUENCY ON
EGG-LAYING AND EGG HATCH IN THE
WALNUT HUSK FLY, RHAGOLETIS COMPLETA
CRESSON
The mating system of Rhagoletis completa Cresson (WHF) has been charac-
terized as a resource-based polygamous system (Opp, S. B. et al. 1996. In: Steck,
G. J. and B. A. McPheron (eds.), Fruit Fly Pests: A World Assessment of Their
Biology and Management. St. Lucie Press, Fla.), in which males actively defend
walnuts to gain exclusive mating opportunities with females seeking oviposition
sites. Post-copulatory mate guarding and multiple matings by males and females
have been observed in field studies (Opp, et al. 1996) but nothing is known about
the effects, negative or positive, of multiple mating behavior on female fertility
and fecundity. This study determined the effects of copulation frequency (single
vs. multiple with each of two mates) on WHE egg-laying propensity and egg
hatch in the laboratory.
WHE adults developed from larvae that were collected from infested walnuts
from Ardenwood Historic Farm (East Bay Regional Parks, Newark, Calif.). The
larvae pupariated, overwintered, and adults eclosed in the lab. Flies were sepa-
rated by sex at eclosion to insure virginity and were held at least two weeks under
laboratory temperatures of 24 + 2° C to insure that both sexes had reached re-
productive maturity (Boyce, A. M. 1934. Hilgardia, 8: 363-579) prior to use in
mating experiments. Adult flies were fed a mixture of sugar and hydrolyzed yeast,
and given water.
Matings were conducted in the lab by placing virgin males and females in a
25 cm? Plexiglas and screen communal cage which was observed continuously
during the mating trial for any copulating pairs. Copulating pairs were carefully
removed in copula from the communal cage using a small, clear plastic cup and
then were held separately in 470 ml plastic cages where they were supplied with
food and water. All initial copulations were timed and only pairs which remained
in copula for at least 5 min were used in the trials (average copulation duration
= 9 min in this study and previous field studies (Opp, et al. 1996). During a
particular mating session, we separated the total number of copulating pairs equal-
ly into two mating treatments (single copulation vs. multiple copulations with
each of two mates). The single copulation treatment consisted of a male and
female pair allowed to copulate once, and, after the pair disengaged, the male
was removed from the cage and the female was allowed to oviposit for a period
of one week. After one week, a mated female was placed in another container
with five virgin males and continually observed for any new matings. The second
male was also removed after the pair naturally disengaged, and the female was
again allowed to oviposit freely for several weeks. Thus, the single copulation
treatment consisted of female flies which had copulated once with each of two
males. In the multiple copulation treatment, the male was not removed after the
first copulation but rather was transferred along with the female to a cage where
236 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
MM Laia Eggs
Produced Larvae
% Mated Females
|
Single Multiple
Mating Treatment
Figure 1. Effect of numbers of matings on egg-laying and larval hatch in the walnut husk fly. (N
= number of females in each mating treatment.)
they remained together for 48 h with food and water and an oviposition substrate.
The pair was allowed to copulate at will, however the exact duration and fre-
quency of subsequent copulations were not recorded. Most pairs were observed
to mate frequently although pairs were not observed continuously beyond the first
copulation. After 48 h, the male was removed and the female was allowed to
continue ovipositing for the remainder of the week. The following week she was
placed in the company of five new virgin males and observed for any matings.
A resulting pair was again removed and transferred to a separate cage and allowed
to copulate at will for 48 h. The second male was also removed after 48 h, and
the female was allowed to oviposit freely for several weeks. Thus, the multiple
copulation treatment consisted of females allowed to copulate many times with
each of two males.
Females in both treatments received artificial substrates which mimicked the
green walnut husks and allowed easy penetration for oviposition (Telang, A. 1995.
M.S. Thesis). The artificial walnuts were constructed of 1.8% agar in distilled
water colored with green food dye which was poured and allowed to set in plastic
snap-together Easter egg molds before being covered with a thin layer of Paraf-
ilm®. Fly eggs were collected intact from the agar walnuts and placed on black
moistened construction paper stacked on top of layers of moistened filter paper
and kept in petri dishes in the dark. Petri dishes were checked daily and egg hatch
recorded. Mean numbers of eggs laid per female and mean numbers of hatching
larvae per female were compared between treatments with Mann-Whitney U-tests
(Sokal & Rohlf. 1981. Biometry. Freeman and Co.). Propensities of females of
each mating treatment to lay eggs and to produce fertile eggs were compared
with G-tests of independence (Sokal & Rohlf. 1981).
Singly-copulated females were both less likely to lay eggs and less likely to
produce larvae than multiply-copulated females (Fig. 1) though these differences
were not statistically significant (G = 0.36, df = 1, P > 0.05). Among females
1996 SCIENTIFIC NOTE 237
laying eggs, multiply-copulated females laid more eggs (i.e., showed greater fe-
cundity) than singly-copulated females (multiply-copulated = 49.1 eggs/female;
singly-copulated = 17.2 eggs/female) even though these differences were not
Statistically significant (W = 126.5, P = 0.38). Multiply-copulated females also
demonstrated greater fertility than singly-copulated females (multiply-copulated
= 37.2 hatching eggs/female; singly-copulated = 5.0 hatching eggs/female)
though, again, no statistically significant difference was detected (W = 5.0, P =
0.33).
We concluded that females allowed multiple copulations had higher levels of
egg-laying and egg hatch than females only allowed a single copulation with each
mate. The lack of statistically significant differences likely resulted from low
sample sizes and high variability but does not detract from the biological signif-
icance of these findings. Repeated copulations may grant females a nutritional,
stimulatory or sperm replenishment benefit as reported for a congener, R. pomo-
nella (Walsh) (Opp & Prokopy. 1986. Ann. Entomol. Soc. Amer., 79: 705-710).
Our study, however, was not designed to test any of the above mentioned hypo-
thetical benefits of multiple copulations in the WHF but to determine the optimal
mating conditions for collection of WHE offspring for laboratory paternity anal-
ysis. Based on our study, we recommend that WHF females be allowed to mate
multiply (i.e., at least twice) with each mate to increase laboratory offspring pro-
duction. These results correlate well with field studies which have reported that
female WHF on average mate approximately twice a day, often with the same
male, and may mate as many as nine times a day with one or more males (Opp
et al. 1996).
Acknowled gment.—We thank the East Bay Regional Parks for permission to
collect and study walnut husk flies in Ardenwood Historic Farm. This paper is in
partial fulfillment of the M.S. Degree of A. Telang and was funded by U.S.D.A.
grant #9204321 to S. B. Opp.
Aparna Telang,! Sha S. Hammond, Susan B. Opp, Department of Biological
Sciences, California State University, Hayward, California 94542. |Current mail-
ing address: University of Arizona, IDP in Insect Science, 410 Forbes Bldg.,
Tucson, Arizona 85721.
Received I Sep 1995; Accepted 14 Feb 1996
PAN-PACIFIC ENTOMOLOGIST
72(4): 238-242, (1996)
Proceedings of the Pacific Coast Entomological Society, 1995
FIVE HUNDRED FIFTEENTH MEETING
The 515th meeting of the Pacific Coast Entomological Society was held on 20 January 1995 at 8:00
PM in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco with President Curtis Y. Takahashi Presiding.
The minutes of the 9 December 1994 meeting were read and accepted. Ms. Cheryl Barr of the
University of California was proposed and accepted as a new regular member.
President Curtis Y. Takahashi announced that the California Department of Food and Agriculture
would be conducting the State Entomologist examination, and also announced an opening for an
Agricultural Biologist position in Contra Costa County as well as several seasonal employment op-
portunities with the CDFA.
Ms. Cheryl Barr presented a unique exhibition of silphid beetles.
The featured speaker, Dr. Rollin Coville presented a slide lecture entitled ‘Photographing Insects
and Spiders in Costa Rica’’. Dr. Coville discussed some of the techniques he utilized in photographing
various arthropods, as well as sharing an entertaining account of his experiences in Costa Rica. The
meeting was adjourned at 9:43 PM, and was followed by a social hour in the Department of Ento-
mology conference room.—Stan Vaughn, Recording Secretary.
The following 45 persons were present. (32 members): P. H. Arnaud Jr., C. B. Barr, L. G. Besark,
J. E. Brandriff, T. S. Briggs, H. K. Court, R. E. Coville, L. W. Currie, D. K. Dabney, J. G. Edwards,
B. K. Eya, S. V. Fend, C. E. Griswold, J. E. Hafernik Jr.. W. Hammersky, R. L. Langston, V. F Lee,
T. Meikle, L. A. Norton, N. D. Penny, A. E. Rackett, K. Reynolds, R. G. Robertson, W. E. Savary,
J. S. Schweikert, W. D. Shepard, EF A. H. Sperling, R. E. Stecker, C. Y. Takahashi, D. Ubick, S. E.
Vaughn, and R. R. White; (13 guests) M. M. Amaud, D. Bromberger, J. E. Court, P. Coville, J. P.
Dorard, K. M. Hallit, S. Meehl, J. Pretare, W. E. Rauscher, S. Renkes, J. Robertson, N. E. Robinson,
and P. Stecker.
FIVE HUNDRED SIXTEENTH MEETING
The 516th meeting of the Pacific Coast Entomological Society was held on 17 February 1995 at
8:00 PM in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco with President Curtis Y. Takahashi presiding.
The minutes of the 20 January 1995 meeting were read, amended and adopted. Dr. Norman Penny
announced that the Academy has CAS drawers available for sale and also mentioned that he will be
offering a course in curation on 12 April 1995, to be held at the Academy. Drs. J. Gordon Edwards
and Ron Stecker of San Jose State University announced the donation of unit trays to be given away
to interested members and guests of the Society.
Leslie Saul of the San Francisco Insect Zoo announced the need for volunteers at the San Francisco
County Butterfly Count before 4 July 1995, and reported on the Butterfly Houses in Denver, Colorado;
Salt Lake City, Utah; and Moody Gardens in Galveston and Houston, Texas. Dr. Jerry Powell of the
University of California announced the retirement celebration honoring Dr. John Chemsak.
The featured speaker, Dr. Norman Penny of the California Academy of Sciences, presented a slide
lecture entitled ‘“Specimen-Level Databases’’. Dr. Penny discussed how the management of specimen-
level databases can increase information availability and accessibility to entomologists and people
beyond the systematic community. Dr. Penny also shared many insights into the future potential of
database information integration utilizing existing software such as mapping, delta keys, data matrix,
and image fields. The meeting adjourned at 9:08 PM and was followed by a social hour in the
Department of Entomology conference room.—Stan Vaughn, Recording Secretary.
The following 37 persons were present: (29 members) C. M. Brandau, T. S. Briggs, M. S. Caterino,
H. K. Court, P. G. da Silva, W. A. Doolin, J. G. Edwards, C. E. Griswold, A. Hom, M. A. Isaak, R.
L. Langston, V. FE Lee, T. C. Meikle, L. A. Norton, A. R. Olsen, J. EF Parinas, A. M. L. Penny, N. D.
Penny, J. A. Powell, K. Reynolds, K. J. Ribardo, L. S. Saul, W. E. Savary, H. I. Scudder, FE A. H.
Sperling, R. E. Stecker, C. Y. Takahashi, S. E. Vaughn, and S. C. Williams; (8 guests) T. Aweeka, K.
Chester, J. E. Court, H. Millspaugh, L. Randal, W. E. Rauscher, R. Takumi, and T. Sidebottom.
1996 PROCEEDINGS 239
FIVE HUNDRED SEVENTEENTH MEETING
The 517th meeting of the Pacific Coast Entomological Society was held on 17 March 1995 at 8:00 PM
in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco
with President Curtis Y. Takahashi presiding. Due to President Takahashi’s tardiness, the meeting was
called to order by the Recording Secretary, Mr. Stanley E. Vaughn, under the insistence of Dr. Norman
Penny and Mr Vincent E Lee.
The minutes of the 17 February 1995 meeting were read and accepted. Proposed and accepted as
new student members were Mrs. Laura Irons, Ms. Melanie Preston, and Mr. Mathew Gigliotti, all
from San Jose State University.
Mr. Vincent E Lee, of the California Academy of Sciences, announced a symposium on the Bio-
diversity of the Sierra-Nevada to be held on the 29th and 30th of April 1995.
The featured speaker, Mr. Al Hom, presented a graphic slide presentation entitled ‘Forensic Ento-
mology: An Enchanted Evening with Maggots and Dead Meat’’. Mr. Hom discussed the roles of
forensic entomologists in the investigations of human deaths and how faunal succession in insects can
be used in determining the times of death. The meeting was adjourned at 10:18 PM and was followed
by a social hour in the Department of Entomology conference room.—Stan Vaughn, Recording Sec-
retary.
The following 35 persons were present: (24 members) J. E. Brandriff, T. S. Briggs, J. G. Edwards,
C. D. Franklin II, J. Garcia, M. J. Gigliotti, W. Hamersky, A. Hom, L. A. Irons, M. A. Isaak, S. E.
Jungers, R. L. Langston, V. KF Lee, B. R. Manchester, L. A. Norton, N. D. Penny, M. A. Preston, A.
E. Rackett, K. Reynolds, L. S. Saul, H. I. Scudder, C. Y. Takahashi, D. Ubick, and S. E. Vaughn; (11
guests) R. Bander, G. Gee, S. Gee, D. H. Ludwig, M. Mann, W. E. Rauscher, J. Scheiman, T. H.
Sidebottom, G. Spaulding, E. Yorkey, and 1 illegible signature.
FIVE HUNDRED EIGHTEENTH MEETING
The 518th meeting of the Pacific Coast Entomological Society was held on 21 April 1995 at 8:00
PM in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco with President Curtis Y. Takahashi presiding.
The minutes of the 17 March 1995 meeting were read and accepted. Proposed and accepted as new
regular members were Ms. Dana H. Ludwig and Mr. Thomas H. Sidebottom, both of the Food and
Drug Administration. Mr. Bill Hamersky, of CSU Hayward, announced that on the 30th of April 1995
CSUH would be hosting a Science Festival. Mrs. Laura Irons, of San Jose State University announced
that the annual SJSU Entomology Club overnighter to Arroyo Seco will be conducted on the 6th and
7th of May 1995, and Ms. Jeanette McNicol also from San Jose State University reported that SJSU
Ent Club T-shirts would be available for purchase after the meeting.
The featured speaker, Mr Cliff Ramos of the California Department of Food and Agriculture, pre-
sented a slide lecture entitled ““The Africanized Bee: General Biology and California Update’’. Mr.
Ramos discussed the general biology of the AHB and detailed the identification methods; range,
distribution, and progression models; and described strategies for those who are attacked and at risk.
The meeting adjourned at 9:01 and was followed by a social hour in the Department of Entomology
conference room.—Stan Vaughn, Recording Secretary.
The following 30 persons were present: (24 members) P. H. Arnaud Jr., T. S. Briggs, M. R. Carbiener,
H. K. Court, L. W. Currie Jr, D. K. Dabney, W. Hamersky, A. Hom, L. A. Irons, B. Keh, R. L.
Langston, V. F Lee, J. J. McNicol. L. A. Norton, N. D. Penny, M. A. Preston, K. A. Reynolds, W. E.
Savary, J. Schweikert, H. I. Scudder, R. E. Stecker, C. Y. Takahashi, D. Ubick, and S. E. Vaughn; (6
guests) T. Aweeka, J. E. Court, J. Myatt, R. Myatt, C. Ramos, and W. E. Rauscher.
FIVE HUNDRED NINETEENTH MEETING
The 519th meeting of the Pacific Coast Entomological Society was held 15 September 1995 at 8:00 PM
in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco
with Present Curtis Y. Takahashi presiding. The minutes of the 21 April 1995 meeting were not read
and approved, due to the.absence of the Recording Secretary.
Proposed and accepted as new regular members were Dr. Janine Casevitz-Weulersse of the Museum
National d’Histoire Naturelle in Paris, France; Dr. David R. Horton of the USDA-ARS Yakima Ag-
ricultural Research Laboratory; Mr. D. Christopher Rogers of Sacramento; Dr. Darrel W. Ross of
240 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
Oregon State University; Dr. Charles G. Summers of the University of California; and Mr. John
Urzykowski-Orzel of Salinas. Proposed and accepted as new regular family members were Dr. David
R. Parks and Ms. Margaret Sharp of San Francisco. Ms. Denise A. Piechnik of San Francisco State
University was proposed and accepted as a new student member. Dr. Ron Stecker introduced Mr. Bob
Clement as a guest, as did Dr. Felix Sperling who introduced Mr. Anthony Cognato and Mr. John
Ascher. Dr. Charles Griswold presented a note on onlyatholipid spiders and African/Madagascan bio-
geography and Dr. Edward Smith gave a detailed note on Salix galls.
The featured speaker Dr. Steve Heydon, of the University of California at Davis, presented a slide
lecture entitled ‘““Desparately Seeking Salix’’. Dr. Heydon described, in detail, the interactions between
Hexomyza salicis and its parasite, Sphegegaster. H. salicis is widely distributed but rarely found in
collections. Dr. Heydon also noted that an undescribed eurytomid has been discovered that also par-
asitizes Hexomyza. The meeting was adjourned at 9:20 PM and was followed by a social hour in the
Department of Entomology conference room.—Stan Vaughn, Recording Secretary.
The following 45 persons were present: (29 members) R. L. Aalbu, P H. Amaud Jr., L. G. Besark,
H. K. Court, B. Deutsch, W. A. Doolin, J. G. Edwards, J. Garcia, C. E. Griswold, W. Hamersky, A.
Hom, V. F Lee, J. J. McNicol, T. Meikle, L. A. Norton, D. A. Piechnik, J. M. Ribardo, K. J. Ribardo,
R. G. Robertson, K. N. Schick, J. Schweikert, H. I. Scudder, E. L. Esith, EF A. H. Sperling, R. E.
Stecker, C. Y. Takahashi, D. Ubick, J. Urzykowski-Ozel, and R. L. Zuparko; (16 guests) M. M. Amaud,
J. Ascher, T. Aweeka, R. Clement, A. I. Cognato, J. E. Court, Mrs. A. Doolin, S. L. Heydon, E.
Lacabanne, D. Maffei, W. Maffei, J. Myatt, R. Myatt, N. Pickard, J. Robertson, and G. Spaulding.
FIVE HUNDRED TWENTIETH MEETING
The 520th meeting of the Pacific Coast Entomological Society was held on 20 October 1995 at 8:00
PM in the Goethe Room of the California Academy of Sciences, Golden Gate Park, San Francisco
with President Curtis Y. Takahashi presiding. The minutes of the 15 September 1995 meeting were
read and accepted.
Proposed and accepted as new regular members were Dr. Kee-Jeong Ahn of the University of
Kansas, Dr. Mohammed T. Aliniazee of Oregon State University, Dr. Jeng-di Lee of the National
Museum of Natural Science in Taiwan, and Dr. Richard S. Vetter of the University of California at
Riverside. Proposed and elected as new student members were Mr. Anthony Cognato of UC Berkeley,
and Ms. Elizabeth Lacabanne of San Francisco State University.
Mr. Michael Carbiener of San Jose State University announced that the SJSU Entomology Club
Christmas party will be held on the 15th of December 1995 at the home of Dr. J. Gordon Edwards
in San Jose. Mr. Vincent F Lee of the California Academy of Sciences reported the status of the
ongoing publication of the Pan-Pacific Entomologist. Dr. Robert Dowell of the California Department
of Food and Agriculture has taken over as the new editor. Mr. Lee also announced the arrival of a
new exhibit, a flea circus, at the Exploratorium called ‘‘About the Size of It”. Ms. Jean M. Ribardo
exhibited several scarab beetle larvae that were unearthed from the front flower beds at Sandoz
Corporation. Dr. Ron Stecker of San Jose State University exhibited some Cuban Laural thrips that
were collected from the Federal Building in San Jose, and Mr. Warren Savary of the Food and Drug
Administration presented a note on a beetle parasite of Shitake mushrooms.
The featured speaker, Dr. Simon Pollard of the University of Canterbury, presented an engaging
slide lecture entitled ‘Spiders: Lifestyles of the Small and Hairy’’. Dr. Pollard discussed the techniques
and mechanisms of feeding in crab spiders, demonstrating how the spider’s prey acts as an extension
of the spider’s digestive system. The prey is not only a source of food, but aids in the mechanics of
feeding. Truly outstanding photography! The meeting adjourned at 9:23 PM and was followed by a
social hour in the Department of Entomology conference room.—Stan Vaughn, Recording Secretary.
The following 57 persons were present: (38 members) P. H. Amaud Jr., C. B. Barr, L. G. Bezark,
J. E. Brandriff, T. S. Briggs, J. S. Chinn, H. K. Court, P. R. Craig, B. Deutsch, N. E. Gershenz, C. E.
Griswold, W. Hamersky, A. Hom, L. A. Irons, E. Lacabanne, R. L. Langston, V. E Lee, T. Meikle,
M. H. Niehoff, A. R. Olsen, N. D. Penny, W. J. Pulawski, K. A. Reynolds, J. M. Ribardo, K. J.
Ribardo, R. G. Robertson, E. S. Ross, L. S. Saul, W. E. Savary, K. N. Schick, H. I. Scudder, T. H.
Sidebottom, R. E. Stecker, C. Y. Takahashi, D. Ubick, S. E. Vaughn, S. P. Welles, and R. L. Zuparko;
(19 guests) M. M. Arnaud, T: Aweeka, R. Bandar, D. Banks, M. R. Carbiener, J. E. Court, F Crosby,
D. D. Giuliani, S. Haugues, M. Imberski, K. Kyle, J. Myatt, R. Myatt, W. A. Rauscher, J. Schick, S.
Schopen, P. Stecker, M. Stekanski, and C. Zuparko.
1996 PROCEEDINGS 241
FIVE HUNDRED TWENTY-FIRST MEETING
The 521st meeting of the Pacific Coast Entomological Society was held on 17 November 1995 at
8:00 PM in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San
Francisco with President Curtis Y. Takahashi presiding. The minutes of the 20 October 1995 meeting
were read and accepted.
Prior to the meeting, a reception was held in the Goethe Room honoring Dr. Paul H. Arnaud Jr. Dr.
Arnaud served as Curator of Diptera at the Academy for 37 years before retiring.
Mr. Warren Savary of the FDA announced the death of Mont Cazier.
Proposed and accepted as new regular members were Dr. Byron A. Alexander of the University of
Kansas, and Dr. Serguei V. Triapitsyn of the University of California at Riverside. One student member
was accepted, Mr. John A. Rumph of Washington State University.
The featured speaker, Mr. Warren Savary of the Food and Drug Administration, presented a com-
puter-projected lecture entitled ““There’s More on the Web than Spiders: A World Overview of World
Wide Web Resources for Entomology”’. Mr. Savary introduced techniques for accessing entomological
information on the Internet, as well as detailing strategies for information retrieval. The meeting
adjourned at 9:20 and was followed by a social hour in the Department of Entomology conference
room.—Stan Vaughn, Recording Secretary.
The following persons were present: (29 members) P. H. Arnaud Jr., C. B. Barr, L. G. Besark, T:
S. Briggs, J. S. Chin, H. K. Court, D. K. Dabney, W. A. Doolin, J. G. Edwards, E. M. Fisher, N. E.
Gershenz, W. Hammersky, A. Hom, L. A. Irons, R. L. Langston, V. FE Lee, J. J. McNicol., T. Meikle,
W. J. Pulawski, K. A. Reynolds, J. M. Ribardo, K. J. Ribardo, L. S. Saul, W. E. Savary, J. Schweikert,
H. I. Scudder, R. E. Stecker, C. Y. Takahashi, and S. E. Vaughn; (6 guests) M. M. Arnaud, J. E. Court,
E. Dunbar, W. A. Rauscher, N. M. Schiff, and S. Udayagini.
FIVE HUNDRED TWENTY-SECOND MEETING
The 522nd meeting was held on 8 December 1995 at 8:00 PM in Room 135 in the Carl D. Duncan
Hall of Science at San Jose State University, San Jose with President Curtis Y. Takahashi presiding.
The minutes of the 17 November 1995 meeting were read and accepted. There was one candidate that
was proposed and accepted as a new regular member, Dr. Miguel A. Moron from the Instituto de
Ecologia in Veracruz, Mexico.
Membership Chair William Hamersky reported that the total membership for 1995 fell by 21 from
1994, for a total of 385. Ms. Katherine Reynolds, on behalf of the Nominating Committee announced
the nominations of officers for 1996; President: Dr. Wojciech J. Pulawski, President-Elect: Mr. Warren
Savary, Treasurer: Ms. Julieta Parinas, Managing Secretary: Mr. Vincent FE Lee, and Recording Sec-
retary: Mr. Stanley E. Vaughn. All nominees accepted and were voted in by the membership.
Dr. Norman Penny of the California Academy of Sciences announced that the Audit Committee
has met, and that details would be forthcoming. Mr. Vincent E Lee announced that the Society ex-
changed the Pan-Pacific Entomologist with 52 organizations and received 71 serials in 1995. Mr. Lee
also reported the following new PCES exchanges in 1995: The Boletin del Museo de Entomologia de
la Universidad del Valle; Cali, Colombia; European Journal of Entomology (continuation of Acta
Entomologica Bohemoslovaca and Casopis) Czech Academy of Sciences, Ceske Budejovice, Czech
Republic; Folia Heyrovskyana, Prague, Czech Republic; Japanese Journal of Systematic Entomology
(continuation of Transactions of the Shikoku Entomological Society) Matsuyama, Japan; and Mittei-
lungen des Thuringer Entomologenverbandes e. V., Kranichfeld, Germany.
Dr. Penny announced the Academy has unit trays available. Mr. Warren Savary exhibited a number
of arthropods that were seized by the FDA. These arthropods were imported to provide questionable
health and recreational benefits common in some cultures.
The featured speaker, Mr. Curtis Y. Takahashi presented a slide lecture entitled “‘A Brief Introduction
to Cultural Entomology’’. Mr. Takahashi illustrated the unique roles that insects have and continue to
play in all societies. From ancient cultures to present times, arthropods have been influential in dis-
ciplines as diverse as religion to entertainment. The meeting was adjourned at 9:10 and was followed
by a social hour in the San Jose State Entomology Department Laboratories.—Stan Vaughn, Recording
Secretary.
The following 28 persons were present: (22 members) R. M. Brown, M. R. Carbiener, T: D. Cuneo,
242 THE PAN-PACIFIC ENTOMOLOGIST Vol. 72(4)
J. G. Edwards, C. D. Franklin I, W. Hammersky, L. A. Irons, M. A. Isaak, V. K Lee, J. J. McNicol,
L. A. Norton, N. D. Penny, M. A. Preston, A. E. Rackett, K. A. Reynolds, J. M. Ribardo, W. E.
Savary, N. M. Schiff, R. E. Stecker, C. Y. Takahashi, and S. E. Vaughn; (6 guests) N. S. Darby, M.
Imberski, D. Irons, J. Myatt, R. Myatt, and E. Takahashi.
PAN-PACIFIC ENTOMOLOGIST
72(4): 243-244, (1996)
PACIFIC COAST ENTOMOLOGICAL SOCIETY
NOTES TO THE FINANCIAL STATEMENTS
YEAR ENDED SEPTEMBER 30, 1995
SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES
Accounting Method
Income and expenses are recorded by using the cash basis of accounting.
Note from the Treasurer
The Pan-Pacific Entomologist, the journal of the Pacific Coast Entomological
Society, is published quarterly. However, due to editorial delays, the issues are
often not published and charged to the Society on schedule. This explains the
abnormal fluctuation in publishing costs.
Capital Expenditures
Annual capital expenditures of $5,000 or less are charged to expense.
Marketable Securities
American Telephone & Telegraph Co., Pacific Telesis Group and Air Touch Com-
munications common stocks are carried at market value. Increases and decreases
in value are reflected in income.
Income Tax
The Society is exempt from Federal Income and California franchise tax.
As Chairman of the Accounting and Tax Committee, and in accordance with the
Society’s bylaws, I have reviewed the financial records of the Society but have
not made an audit of them.
During the course of this review, nothing was noted which indicated any material
inaccuracy in the financial statements.
H. Vannoy Davis
Chairman of the Accounting and Tax Committee
THE PAN-PACIFIC ENTOMOLOGIST
PACIFIC COAST ENTOMOLOGICAL SOCIETY
STATEMENT OF INCOME, EXPENDITURES AND
CHANGES IN FUND BALANCES
YEARS ENDED SEPTEMBER 30, 1995 AND 1994
pele
Income
PES sand SUBSE TIONS a Fe fccs-n 0 steecate Fecasin ne adh orsde acaaiacn veal Genes Keduiae nn els acdeoiacessio ay adhe $ 17,066
Reprints: and aiscellaneous 2:24.05 -c205. 525-2 eet ee Sten ieee ede e ne 10,421
| CU e) oc) Be ee A Orn SUR Ainge TABATA icp MOAT TRA A SA ORL ABA A Ridin MURATA miner lee TEA A 3,730
Dividends ase Ries Vee yee net edie Meee eres a ey Te kee ge re ge eas te Oe Ba 681
Increase (Decrease) in value of capital stock:
American Telephone & Telegraph Company ...................22222e eee 940
Pacific Telesis Group and Air Touch Communications (see note) ..... $28
Fotalu Ime oie oes 097 oi ees ae ee a Ses» BSCR Howe Bo ee, Oe $ 33,366
Expenditures
Publication costs—Pan-Pacific Entomologist ..................eeceeeeeee eee $ 25,590
REprinE COSISH occu pn ptou eget sour aate hopes dee cette tty ate wees on ie 2,614
Postage, newsletter and miscellaneous expenses ...............--00eeeeeees 1,274
Gta Wxqeim Gites. p82 koe ance ate oe Pe ecistte, e Me ae lee dacs Ee sede ale Ak BS $ 29,478
Increasesin-ftundabalances 58.03. ett en eee BY ee” SM oe $ 3,888
Fund balances October 1, 1995 and 1994.0... 00 eee ee 129,370
Fund balances September 30, 1995 and 1994 ..........................0085 $133,258
STATEMENT OF ASSETS
AS OF
SEPTEMBER 30, 1995 AND 1994
1995
Cash in bank
COMMEFCAl ACCOUNT. es) esses ls thee Date eee aon Nepeele ot loee ait Meda ds $ 17,914
Special Funds:
General Fund—Wells Fargo Bank .................00eceeeeeeee cece eeeeeees 4,942
C. P. Alexander Fund—Capital Preservation Fund ...................... 54,711
Fall Memoir Fund—Wells Fargo Bank ..................02e eee eee eee e eee 34,228
Total cash in bank and special funds ........................ 00 ccc eee ee eee $111,795
Capital Stock (at market value)
American Telephone & Telegraph Co., 80 shs. ...............2. ee eee cece eee 5,260
Pacitic: Telesis, Group, 264 SHS. e, ots le ove! acg shed loge & ayase late wlopels onplace abe aleebalegticg 8,118
Air Touch Communications, 264 Shs. ............ ccc cece cece cence ceceeeueuees 8,085
21,463
Total Assets
$133,258
See accompanying notes to the financial statements
Vol. 72(4)
1994
$ 17,504
11,010
2,604
681
(370)
1,716
$ 33,145
$ 19,448
2,138
1,353
$22,939.
$ 10,206
119,164
$129,370
1994
$ 15,386
4,832
52,015
37,142
$109,375
4,320
8,118
7,557
19,995
$129,370
PAN-PACIFIC ENTOMOLOGIST
72(4): 245, (1996)
1995 SPONSORING MEMBERS OF THE
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PAN-PACIFIC ENTOMOLOGIST
72(4): 246, (1996)
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Volume 72
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PAN-PACIFIC ENTOMOLOGIST
72(4): 247, (1996)
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PAN-PACIFIC ENTOMOLOGIST
72(4): 248-249, (1996)
The Pan-Pacific Entomologist
Contents for Volume 72
Announcement—Publications of the Pacific
Coast Entomological Society ... 180, 247
BERNHARDT, P. & K. WALKER—Observations on
the foraging preferences of Leioproctus
(Filiglossa) Rayment (Hymenoptera: Col-
letidae) in eastern Australia 130
Brown, J. W.—New generic assignment, new
synonymy, and new distribution records for
the neotropical genus Icteralaria Razowski
(Lepidoptera: Tortricidae) ......... 162
BRUNNER, J. E—Discovery of Colpoclypeus
florus (Walker) (Hymenoptera: Eulophidae)
in apple orchards of Washington ...... 5
CAMBRA, R. A. & D. QUINTERO A.—The Mexican
and Central American species of Lopho-
stigma Mickel, including a new species, new
distribution records, and taxonomic notes for
the genus (Hymenoptera: Mutillidae) ... 92
CASEVITZ-WEULERSSE, J.—Biogeographic aspects
of the ant fauna of Corsica (Hymenoptera:
Formicidae) 193
COHER, E. I.—Cave-associated tropical American
Neoditomyia (Diptera: Mycetophilidae) 152
DOWELL, R. V—Emmel, T. C., M. C. Minno &
B. A. Drummond. 1992. Florissant
Butterflies: A Guide to the Fossil and
Present-day Species of Central Colorado.
Stanford University Press. 118pp. + I-IX
FE A Oe ae Ra en aT! 173
Fox, C. W., A. D. HARBIN & T. A. MoussEAU—
Suitability of a non-host palo verde for
development of Stator limbatus (Horn)
(Coleoptera: Bruchidae) larvae ...... 31
GILL, R. J., see SORENSEN, J. T. ........ 160
GRAVES, P. L., see HALBERT, S.E. ....... 13
GREVE, J. H., see MCFADZEN, M.E....... 41
GRISWOLD, T. L.—A new Microbembex endemic
to the Algodones Dunes, California
(Hymenoptera: Sphecidae) 142
GULMAHAMAD, H.—Gnathamitermes perplexus
(Banks) (Isoptera: Termitidae): A nuisance
structural termite pest in southern California
PR are ee fh ne ee 9 Be 37
Haas, J.. D. W. Ross, R. R. Mason, T. R.
TORGERSEN & A. R. MOLDENKE—
Geographic variation in aboreal spider
(Araneae) communities on Douglas-fir in
Westen OreoOnh s.. oleh 4ue ie ee: 17
HALBERT, S. E., J. B. JOHNSON, P. L. GRAVEs, P.
M. MaArsH & D. NELSON—Aphidius
uzbekistanicus (Hymenoptera: Aphididae)
established in Idaho.............. 13
HAMMOND, S. S., see TELANG, A. ....... 235
Harsin, A. D., see Fox, C. W. ......... 31
HEADRICKS, D. H., see TRIAPITSYN, S. V... 168
HOFFMAN, M. P, see PICKETT, C. H...... 220
JOHNSON, J. B., see HALBERT, S. E. ...... 13
Lanc, R. F, J. M. Story & G. L. PIPER—
Establishment of Larinus minutus Gyllenhal
(Coleoptera: Curculionidae) for biological
control of diffuse and spotted knapweed in
the western United States......... 209
LARSEN, E.—The distribution of aquatic
Heteroptera (Notonectidae) in Hawaii 39
LIANG, A.-P—Taxonomic changes in Chinese
Lophopidae with a check list of Chinese
species (Homoptera: Fulgoroidae) ... 145
Lyon, R. J.—New cynipid wasps from the
southwestern United States (Hymenoptera:
Gymipidaes N07 ee vg a eee 181
Markow, T. A. see SNOOK,R.R. ...... 121
Marsh, P. M., see HALBERT, S.E. ....... 13
MARTINEZ, M. J.—The first North American
record for the ant Pheidole fervenas Fr.
Smith (Hymenoptera: Formicidae) 171
Mason, R.R., see HALAJ, J... 2.2.2.2... Wa
Mayer, D. F—Effects of sex ratio and female
density on progeny survival of the alfalfa
leafcutter bee (Hymenoptera: Megachilidae)
Fit Ns eee ie Oe an 138
MCcELRAVY, E. P, see Woop, J.R....... 227
McFADZEN, M. E., M. S. VEKSaAy, T. Y.
MoRISHITA & J. H. GRAvES—Northern range
extension for Haematosiphon inodorus
(Dugés) (Hemiptera: Cimicidae) ..... 41
MLLLs, N. J., see RippIcK, E.W. ....... 213
MOLDENKE, A.R., see HALAJ, J. ........ 17
MorRISHITA, T. Y., see MCFADZEN, M. E. ... 41
MOuSSEAU, T. A., see Fox, C. W......... 31
NasT, J. & W. M. WEINER—Geographical
distribution of Brachystomellinae (Collem-
bola: Neanuridae)
NELSON, D., see HALBERT, S.E.......... 13
Opp, S. B., see TELANG, A. ...........
PAcIFIC COAST ENTOMOLOGICAL SOCIETY—
Financial statement for 1991, 1992 .. 174
PAcIFIC COAST ENTOMOLOGICAL SOCIETY—
Financial statement for 1992, 1993 .. 176
PACIFIC COAST ENTOMOLOGICAL SOCIETY—
Financial statement for 1993, 1994 .. 178
1996
PAcIFIC COAST ENTOMOLOGICAL SOCIETY—
Financial statement for 1994, 1995 .. 243
PaciFIC COAST ENTOMOLOGICAL SOCIETY—
Financial statement for 1990, 1991 ... 55
PACIFIC COAST ENTOMOLOGICAL SOCIETY—
Proceedings for 1990 ............. 43
PacIFIC COAST ENTOMOLOGICAL SOCIETY—
Proceedings for 1991 ............. 47
PaAcIFIC COAST ENTOMOLOGICAL SOCIETY—
Proceedings for 1992 ............ 104
PACIFIC COAST ENTOMOLOGICAL SOcIETY—
Proceedings for 1993 ............ 109
PacIFIC COAST ENTOMOLOGICAL SOCIETY—
Proceedings for 1994 ............ 114
PACIFIC COAST ENTOMOLOGICAL SOCIETY—
Proceedings for 1995 ............ 238
PACIFIC COAST ENTOMOLOGICAL SOcCIETY—
Sponsoring members 1990-1994 .... 57
PaciFIC Coast ENTOMOLOGICAL SOcIETY—
Sponsoring members 1995 ........ 245
PICKETT, C. H., S. E. ScHOENIG & M. P.
HOFFMANN—Establishment of the squash
bug parasitoid, Trichopoda pennipes Fabr.
(Diptera: Tachinidae), in northern California
CN eee te ee ty 6 Oa Rt Mere SU 220
Piper, G. L., see LANG, R. E.......... 209
QUINTERO-A., D., see CAMBRA, R. A. ..... 92
REsH, V. H., see Woop, J. R. 2o7
Rippick, E. W. & N. J. MiILLs—Prterostichus
beetles dominate the carabid assemblage in
an unsprayed orchard in Sonoma County,
California 213
RIppIcK, E. W. & N. J. MILLsS—A comparison of
the seasonal activity of Pterostichus beetles
(Coleoptera; Carabidae) in a commercial
apple orchard in Sonoma County, California
Rocers, D. C.—Eubranchipus bundyi Forbes
(Anostraca: Crustacea), a new record from
Wali PONia. 28 = Lie eg ge 89
Ross, D. W.—Phenology of pandora moth
(Lepidoptera: Saturniidae) adult emergence
and egg eclosion in central Oregon .... 1
Ross, D. W., see HALAJ, J. ........00.0. 17
Rust, R. W. & J. L. PORTER—Genetic variation
in Bombus appositus Cresson (Hymenop-
tera: Apidae) 202
CONTENTS FOR VOLUME 72
249
SCHOENIG, S. E., see PICKETT, C. H. ..... 220
SIKES, D. S.—The natural history of Nicrophorus
nigrita, a western neartic species (Cole-
optera: Silphidae)
SNOOK, R. R. & T. A. MARKow—Possible role of
nonfertilizing sperm as a nutrient source for
female Drosophila pseudoobscura Frolova
(Diptera: Drosophilidae) ......... 121
SORENSEN, J. T. & R. J. GLL—A range extension
of Homalodisca coagulata (Say) (Hem-
iptera: Clypeorrhyncha: Cicadellidae) to
southern California 160
SORENSEN, J. T—Recent California records for
the sawfly Xiphydria mellipes Harris
(Hymenoptera: Xiphydriidae) 102
Story, J. M., see LANG, R. FE 209
TELANG, A., S. S. HAMMOND & S. B. Opp—
Effects of copulation frequency on egg-
laying and egg hatch in the walnut husk fly,
Rhagoletis completa Cresson ...... 235
The Pan-Pacific Entomologist—Index for
VOMMGe 2 p48. eee ok ee ge 250
The Pan-Pacific Entomologist—Reviewers for
AV CUTIE 7D Stee st ee i eee 246
The Pan-Pacific Entomologist—Reviewers for
Volumes 70 and 71
The Pan-Pacific Entomologist—Table of
Contents for Volume 72.......... 248
TORGERSEN, T. R., see HALAJ, J. ......... 17
TRIAPITSYN, S. V. & D. H. HEADRICKS—
Description of the male of Ceranisus
americensis (Girault) (Hymenoptera: Eu-
lophidae) 168
VEKSAY, M. S., see MCFADZEN, M.E. .... 41
WALKER, G. P-—Leaf age preference for
Oviposition by the monophagous whitefly,
Aleurotithius timberlakei (Homoptera:
Aleyrodidae).................. 164
WALKER, K., see BERNHARDT, P......... 130
WEINER, W., M. see NAJT, J... 2... we. 61
Woop, J. R., E. PR McELRAvy & V. H. REsH—
Thermal-shock tolerance of three species of
aquatic insects in a northern California
geothermally influenced stream DY:
ZUPARKO, R. L.—Hymenoptera reared from
Plagiotrochus suberi (Hymenoptera: Cyni-
pidae) galls in California .......... 27
PAN-PACIFIC ENTOMOLOGIST
72(4): 250-251, (1996)
The Pan-Pacific Entomologist
Index to Volume 72
(title and key words)
aboreal spiders 17
adult pandora moth emergence 1
Aleurotithius timberlakei 164
Aleyrodidae 164
alfalfa leafcutter bee 138
alfalfa leafcutter bee progeny survival 138
alfalfa seed 138
Algodones Dunes 142
allozymes 202
Anasa tristis 220
Andricus flocculentus NEW SPECIES 181
Anisops kuroiwae 39
Anostraca 89
ant fauna 193
Antron daileyi NEW SPECIES 181
Antron franklinensis NEW SPECIES 181
Antron madera NEW SPECIES 181
Aphidiidae 13
Aphidius uzbekistanicus 13
Apidae 202
apple orchard 5
aquatic Heteroptera 39
Arachnida 17
Aranae 17
Asiraca choui NEW COMBINATION 145
Australian Colletidae 130
biogeography 61, 193
biological control 13, 209, 220
Bombus appositus 202
Brachystomellinae 61
Bruchidae 31
Buenoa pallipes 39
burying beetle 70
California records for Xiphydria mellipes 102
California, Gnathamitermes perplexus in 37
California, Plagiotrochus suberi in 27
California, Trichopoda established in 220
carabid seasonal activity 82, 213
Carabidae 82, 213
carabids in apple orchard 213
carrion 70
caves 152
Centaurea 209
Ceranisus americensis 168
Cercidium 31
Ceropteres snellingi NEW SPECIES 181
check list of Chinese Lophopidae 145
Cicadellidae 160
Cimicidae 41
Coleoptera 31, 70, 82, 209, 213
Collembola 61
Collembola distribution 61
Colletidae 130
Coloradia pandora |
Colpoclypeus florus 5
copulation frequency 235
cork oak galls 27
Corsica 193
Crustacea 89
Curculionidae 209
Cynipidae 27, 181
degree-days 1
diffuse knapweed 209
Diptera 121, 152, 220, 235
Diuraphis noxia 13
Drosophila pseudoobscura 121
Drosophilidae 121
egg eclosion 1
egg hatch 235
egg-laying 235
ejaculate 121
endemic sphecid in California 142
Ephemeroptera 227
Eubranchipus bundyi 89
Euderus crawfordii 27
Eulophidae 5, 168
Fabaceae 31
fairy shrimp 89
Falco mexicanus 41
female density 138
foraging preferences 130
Formicidae 171, 193
Fulgoroidea 145
genetic variation 202
geographic variation 17
geothermal 227
Gnathamitermes perplexus 37
Haematosiphon indorus 41
Hawaii 39
Hemiptera 41, 160
Heteroptera 39
Histiostomidae 70
Homalodisca coagulata 160
1996 INDEX FOR VOLUME 71
Homoptera 145
host range 31
Hymenoptera 5, 13, 27, 92, 102, 130, 138, 142,
168, 171, 181, 193, 202
Icteralaria 162
Icteralaria incusa NEW COMBINATION 162
Idaho, Aphidius uzbekistanicus in 13
Idaho, Haematosiphon inodorus in 41
inquiline 181
Larinus minutus 209
leaf age preference 164
leafroller parasite 5
Leioproctus 130
Lepidoptera 1, 162
Lophomutilla 92
Lophopidae of China 145
Lophostigma 92
Lophostigma distribution records 92
Lophostigma grisselli NEW SPECIES 92
male of Ceranisus americensis 168
male-derived nutrient donation 121
Megachile rotundata 138
Megachilidae 138
Microbembex elegans NEW SPECIES 142
monothalamous gall 181
Mutillidae 92
Mycetophilidae 152
Neanuridae 61
Neoditomyia 152
Neoditomyia farri NEW SPECIES 152
Neoditomyia para NEW SPECIES 152
new California ant record 171
new California fairy shrimp 89
new host record 5
new host-parasitoid associations 27
new North American ant record 171
Nicrophorinae 70
Nicrophorus nigrita 70
nonfertilizing sperm 121
Notonecta indica 39
Notonectidae 39
Oregon, moth phenology in 1
palo verde 31
Parkinsonia 31
Pelzneria 70
Persoonia 130
Pheidole fervens 171
phenology 1
Plagiotrochus suberi 27
251
Poecilochirus carabi 70
Poecilochirus subterraneus 70
population genetics 202
predators 82
Pseudotsuga menziesii 17
Pterostrichus 82
Pterostrichus cursitor 213
Pterostrichus lustrans 213
Pterostrichus species 213
Quercus arizonica 181
Quercus pungens 181
Quercus suber 27
Quercus toumeyi 181
range extension of Haematosiphon inodorus 41
range extension of Homalodisca coagulata 160
raptor nests 41
Rhagoletis completa 235
Saturnidae 1
Schizaphis graminum 13
sex ratio 138
Silphid natural history 70
Silphidae 70
Sitobion avenae 13
Sonoma County, California 82
sperm polymorphism 121
Sphecidae 142
spider community structure 17
spotted knapweed 209
squash bug parasitoid 220
Stator limbatus 31
structural pest 37
Tachinidae 220
Tephritidae 235
termite 37
Termitidae 37
thermal tolerance 227
thermal-shock 227
Tortricidae 162
Trichopoda pennipes 220
Trichoptera 227
tropical America 152
walnut husk fly 235
Washington, new Eulophidae in 5
whitefly oviposition preferences 164
Xanthoteras pungens NEW SPECIES 181
Xanthoteras tuckeri NEW SPECIES 181
Xiphydria mellipes 102
Xiphydriidae 102
zoographic regions 61
PAN-PACIFIC ENTOMOLOGIST
Information for Contributors
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Literature Cited. — Format examples are:
Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.
Blackman, R. L.. P. A. Brown & V. FE Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide
some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. F Dixon & L. Weismann (eds.). Population structure, genetics and
taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB
Academic Publishing, The Hague, The Netherlands.
Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.
Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.
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THE PAN-PACIFIC ENTOMOLOGIST
Volume 72 October 1996 Number 4
Contents
LYON, R. J.—New cynipid wasps from the southwestern United States (Hymenoptera: Cynip-
TAG) 2 aed ert 8g, Sige WW at Beet Ra Be) si ee RD he RO ls Pee MoM Ose 181
CASEVITZ-WEULERSSE, J.—Biogeographical aspects of the ant fauna of Corsica (Hyme-
NO PREK AL OO RUPECTER AC) ea wale orcad Rr Pes AC UO See ee ME gs We ere ae Be 193
RUST, R. W. & J. L. PORTER—Genetic variation in Bombus appositus Cresson (Hymenoptera:
PTC Cy Teed lc eee Pier ee rn A ee tae ere emer ower ge To 202
LANG, R. F, J. M. STORY & G. L. PIPER—Establishment of Larinus minutus Gyllenhal
(Coleoptera: Curculionidae) for biological control of diffuse and spotted knapweed in
the “Western: UNaGeGlsS tates ptewern bern. Sele DAN A 8 Siesta te eB eas WI Veen ea 209
RIDDICK, E. W. & N. J. MILLS—Prerostichus beetles dominate the carabid assemblage in an
unsprayed orchard in Sonoma County, California — ZS
PICKETT, C. H., S. E. SCHOENIG & M. P- HOFFMANN—Establishment of the squash bug
parasitoid, Trichopoda pennipes Fabr. (Diptera: Tachnidae), in northern California _____. 220
WOOD, J. R., E. PR. MCELRAVY & V. H. RESH—Thermal-shock tolerance of three species
of aquatic insects in a northern California, geothermally influenced stream __----- 227
SCIENTIFIC NOTES
TELANG, A., S. S;. HAMMOND, & S. B. OPP—Effects of copulation frequency on egg-laying
and egg hatch in the walnut husk fly, Rhagoletis completa Cresson ___-_--------------------- 235
Pacific Coast Entomological Society, Proceedings for 1995 2a 238
Pacific Coast Entomological Society, financial statements for 1994, 1995 __ 243
Pacific Coast Entomological Society, Sponsoring Members 1995 ______ 245
PAN-PACIFIC ENTOMOLOGIST REVIEWERS, Volume 72. ___--_-_--------------------------- nanan 246
Announcement: publications of the Pacific Coast Entomological Society —___- 247
The Pan-Pacific Entomologist: Table of Contents for Volume 72. __-_-----_----------------n--n-n-nnean 248
ithe Pan-Pacific Entomologist: Indexdior Vole 72 2.1.22 2 ee 250