The Pan-Pacific Entomologist

EDITORIAL BOARD

J. Y. Honda, Editor R. M. Bohart

Z. Nisani, Assistant to the Editor J. T. Doyen

R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.

Robert Zuparko, Treasurer Warren E. Savary

Published quarterly in January, April, July, and October with Society Proceedings

usually appearing in the following October issue. All communications regarding

non-receipt of numbers should be addressed to: Vincent F. Lee, Managing Secretary;

and financial communications should be addressed to: Robert L. Zuparko, Treasurer;

at: Pacific Coast Entomological Society, Dept. of Entomology, California Academy

of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

Application for membership in the Society and changes of address should be

addressed to: Membership 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. Jeff Honda,

Editor, Pan-Pacific Entomologist, Dept. of Biological Sciences, San Jose State

University, One Washington Square, San Jose, CA 95192-0100. See the back

cover for Information-to-Contributors, and volume 73(4): 248—255, October 1997,

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 72(4): 247 for current prices. Make checks payable to the Pacific

Coast Entomological Society.

Pacific Coast Entomological Society

OFFICERS FOR 2001

Stanley E. Vaughn, President Vincent F. Lee, Managing Secretary

Katherine N. Schick, President-elect Joshua J. Fairbanks, Recording Secretary

Robert L. Zuparko, Treasurer

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 addition-

al 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 22 March 2002

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.

The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

PAN-PACIFIC ENTOMOLOGIST

78(1): 1-6, (2002)

SURVIVAL AND DEVELOPMENT OF LACANOBIA

SUBJUNCTA (GROTE & ROBINSON) (LEPIDOPTERA:

NOCTUIDAE) LARVAE ON COMMON WEEDS AND CROP

PLANTS OF EASTERN WASHINGTON STATE

PETER J. LANDOLT

USDA-ARS, Yakima Agricultural Research Laboratory,

Wapato, WA, 98951 USA

Abstract.—Ten common weed species, four tree fruit crops and four row crops were evaluated

as hosts for larvae of Lacanobia subjuncta (Grote & Robinson), a noctuid moth pest of apple

in eastern Washington. A separate comparative evaluation was made of the suitability of five

varieties of apple as hosts for L. suwbjuncta larvae. Development was completed, from neonate

larva to adult, on nine of ten weed species and seven of eight crops tested, indicating a broad

potential host range for this insect. High rates of survival to adult, short developmental times,

and large pupal weights were noteworthy on the weeds bindweed, dandelion, and mallow, and

on potato. In the comparison of apple varieties, highest rate of survival to adult was with Red

Delicious, greatest pupal weights were with Red Delicious, Gala, and Fuji, and shortest devel-

opment times were with Gala and Golden Delicious. Strong seasonal variation (May versus July)

was indicated in the quality of apple foliage as food for L. subjuncta larvae.

Key Words:—Insecta, Lacanobia subjuncta, host plant, apple, potato.

The noctuid moth Lacanobia subjuncta (Grote & Robinson) has recently be-

come recognized as a significant pest of apple in eastern Washington and Oregon

(Landolt 1998). The moth is widely distributed in North America (McCabe 1980),

and has been present in irrigated areas of eastern Washington at least since the

1970s when it was collected in light traps in Yakima County by EF Howell (per-

sonal communication). Following an apparent increase in damage to apple attri-

buted to cutworms (Warner 1996), L. subjuncta was identified as the principal

noctuid pest on apple in eastern Washington and adjacent areas of Oregon (Lan-

dolt 1998).

Lacanobia subjuncta is bivoltine (McCabe 1980), with a flight of moths from

late May into mid June and again from late July into mid September in eastern

Washington (Landolt 1998, Hitchcox 2000). Larvae can be found on apple trees

from early June through July and again from late August until October (Hitchcox

2000). It is thought that the insect overwinters strictly as a pupa in soil. Most

larvae held in the laboratory and fed on apple foliage went through 6 instars

before pupating, although several larvae went through 7 (Hitchcox 2000).

On apple, larvae of L. subjuncta are primarily foliage feeders and occasionally

partially defoliate apple trees in commercial orchards. Damage to apple fruit oc-

curs also, with larval feeding indicated by a hollowed out scoop in the surface

of the apple that is somewhat characteristic of fruit feeding by other noctuids.

The pest status for L. subjuncta on apple is due principally to their feeding on

apple fruit and to problems in packing houses because of the presence of larvae

on fruit (Warner 1998).

The recently acquired pest status of this insect on apple in Washington and

Oregon is not understood. Hypotheses to explain this apparent change in pest

status include 1) resistance to commonly used pesticides together with escape

2. THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

from natural enemies, 2) shifts in larval host plants, 3) changes in geographic

distribution, 4) past misidentification of cutworms and fruit worms damaging ap-

ple, and 5) region-wide L. subjuncta moth population increases resulting in move-

ment of moths into apple orchards. In order to consider the latter hypothesis,

better information is needed on what plants may sustain L. subjuncta reproduction

and may then be the principal host plants contributing to regional moth popula-

tions.

Larvae of L. subjuncta have been collected on a wide variety of plants, in-

cluding trees, shrubs, and herbaceous plants (Godfrey 1972, Rings et al. 1992,

Crumb 1956, Landolt 1998, and included references), indicating a potentially high

degree of polyphagy. Recorded host plants include a number of agricultural crop

plants, such as apple, cherry, peach, blueberry fruit and foliage, cabbage, aspar-

agus, corn, and strawberry. However, the finding of larvae on a plant species is

not necessarily a good indicator of suitable host plant status. The survival and

development of larvae on plants along with the incidence of larvae on plants in

the field would be better indicators of the importance of those plants as contrib-

utors to populations of L. subjuncta. In a preliminary assessment of host suitability

of common weeds, Landolt (1998) demonstrated that L. subjuncta larvae could

complete development on the weeds Taraxacum officinale Weber (dandelion),

Sonchus oleraceus L. (annual sowthistle), Convulvulus arvensis L. (field bind-

weed), and Malva neglecta Wallr (comon mallow), but with low (20 to 28) per-

centages that survived to the adult stage.

Reported here are the results of experiments evaluating the ability of larvae of

L. subjuncta to develop on foliage of a larger number of plant species (18) that

are commonly encountered in irrigated areas of eastern Washington. This evalu-

ation included determination of larval survival to pupation and adult emergence,

larval development time, and pupal weights for L. subjuncta fed on the foliage

of each plant species or variety. The objective of the study was to determine if

these plants sustain complete development of newly hatched larvae through to

adult, and might then contribute in the field to populations of L. subjuncta. Ad-

ditionally, this study sought to identify plant species that might be further eval-

uated in the field as good hosts for L. subjuncta.

MATERIALS AND METHODS

For each plant species and apple variety evaluated, data were obtained on sur-

vival of larvae to pupation and adult emergence, on larval development time (egg

hatch to entry into soil), and on pupal weight. Plants evaluated were weed species

Sonchus asper (L.) Hill (spiny sowthistle), dandelion, common mallow, field bind-

weed, Cirsium arvense (L.) Scop. (Canada thistle), Helianthus annuus L. (sun-

flower), Chenopodium album L. (lambsquarters), Amaranthus retroflexus L. (red-

root pigweed), Cardaria draba (L.) Desv. (hoary cress), and Kochia scoparia (L.)

Schrad. (kochia), the crop plants Malus pumila Mill. (apple, var. Fuji), Pyrus

communis L. (pear, var. Bartlett), Prunus armeniaca L. (apricot), Medicago sativa

L. (alfalfa), Pisum sativum L. (dry peas, var. Columbian), Pisum sativum L. (suc-

culent peas, var. Oregon Trail), and Solanum tuberosum L. (potato, var. Russet

Burbank). In another experiment, the apple varieties Fuji, Gala, Braeburn, Red

Delicious, and Golden Delicious were evaluated as hosts for L. subjuncta larvae.

Adult L. subjuncta were obtained from a walk-in light trap at the Yakima

2002 LANDOLT: LACANOBIA SUBJUNCTA BIONOMICS 3

Agricultural Research Laboratory southeast of Yakima, Washington, in an area of

commercial irrigated apple and pear orchards. Female moths from the light trap

were held for 24 h in 50 ml plastic snap-cap vials for oviposition. Females that

did not oviposit within that time period were assumed to be unmated and were

moved to 900 ml plastic tubs with ventilated lids. These tubs contained sugar

water on cotton balls, water on cotton balls, and two males that had been collected

in the light trap. Some of these females subsequently laid fertile eggs. Newly

eclosed larvae from egg batches were used for the following experiments.

The assay unit was a 300 ml waxed paper carton with a clear plastic lid, in

which was placed plant foliage and one newly hatched larva. Cartons were held

in a room on a 16:10 light:dark photoperiod, at 25° C and 60% RH. Plant foliage

was added daily. Dried, brown, or moldy foliage was removed daily and new

cartons were provided as needed when soiled by frass. When larvae reached about

3 cm in length, 2—3 cm of damp soil was placed in the carton and the plant foliage

and larva were placed on top of this soil. Mature larvae entered the soil to pupate.

Daily records were made of larval mortality, and larval movement into soil. Four

to 6 days after larvae moved into soil, the soil was sifted to confirm the presence

of a pupa, which was then weighed and transferred to a 30 ml paper cup held

inside of the original waxed paper carton. Daily observations were made of pupae

in order to record adult emergence. Data was not obtained on pupal duration

because it was not possible to tell on what day a larva in the soil pupated. Dis-

turbing the larva at that time might interfere with successful development.

Plant foliage was obtained in the field as needed by cutting plants with scissors

and transferring foliage in 3.6 liter Ziplock® plastic freezer bags held in a cooler

until return to the laboratory. Bags of foliage were held in a refrigerator at 3° C

for up to seven days and were accessed daily to obtain foliage that was provided

to larvae. For weeds, plants were selected from areas not sprayed with insecti-

cides. For crop plants, growers were consulted for information on the timing and

application of insecticides to avoid the collection of foliage with pesticide resi-

dues.

For each plant species or apple variety evaluated, three sets of five larvae were

tested, with each set of five larvae originating from the egg batch of a different

female moth. These three sets of larvae were staggered in time (three weeks apart)

so that the individual plants evaluated were different for each set of five larvae.

RESULTS

Larvae of L. subjuncta developed to pupation and adult emergence on the

following weeds: spiny sowthistle, dandelion, bindweed, lambsquarters, mallow,

Canada thistle, hoary cress, pigweed, and kochia (Table 1). No larvae survived

to pupation on sunflower. Highest percentages of larvae surviving to pupation

were fed on field bindweed. Larvae also developed to pupation and adult emer-

gence on the following crop plants: cherry, apple, pear, alfalfa, potato, dry peas,

and succulent peas (Table 1). No larvae survived to pupation on apricot. Highest

percentages of larvae that pupated were on cherry, pear, potato and alfalfa. Larvae

developed to pupation and adult emergence on all 5 apple varieties tested, but

with significant differences in percentage of pupation among these varieties (Table

2). Significantly more larvae on Gala apple foliage died than did on Golden

Delicious foliage.

Table 1. Mean (=SE) % pupation, adult emergence, pupal weight, and larval development times

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 78(1)

for Lacanobia subjuncta on 10 weedy plant and 8 crop species. Yakima, Washington, 2000.?

Pupal weight Larval time

Plant species % Pupated % Adult (mg) (days)

Hoary Cress 66.7 + 13.3c 60.0 + 11.5 SIO aeaRe 44.3 + 1.6a

Common Mallow 80.0 + 11.5be 80.0 + 11.5 363 + 15b 26.1 + 0.7c

Spiny Sowthistle 60.0 + 20.0c 60.0 + 20.0 436 + 10a 26.9 + 1.3bc

Dandelion 86.7 + 6.7b 86.7 + 6.7 392 + 5b 25.7 HLT be

Canada Thistle 46.7 + 6.7d 46.7 + 6.7 249 + l6e 28.7 + 1.5be

Common Sunflower 00.0 + 00.0e 00.0 + 00.0 it ——

Lambsquarter 85.0 + 5.0b 73.4 + 6.7 342 + Ile 28.4 + 1.6bc

Redroot Pigweed 70.0 + 10.0d 70.0 + 10.0 280 + 10de 35.1 + 2.9ab

Field Bindweed 100 + 00.0a 100 + 00.0 374 + 8b 25.1 + 0.6c

Kochia 40.0 + 11.5d 40.0 + 11.5 270 + 12e 35.8 + 2.7ab

Apple 66.7 + 6.7c 66.7 + 6.7 389 + 7b 25.1 + 0.7c

Pear 100.0 + 00.0a 100 + 00.0 333 + 8c 26.5 + 0.7bec

Cherry 93.4 + 6.7ab 86.7 + 6.7 323 + 12cd 30.4 + 1.1b

Apricot 00.0 + 00.0e 00.0 + 00.0 —+— a

Potato 100 + 00.0a 100 + 00.0 397 = )2b 24.7 + 0.9c

Alfalfa 100 + 00.0a 100 + 00.0 297 + 8d 29.9 + 0.9b

Dry Peas 86.7 + 13.3b 60.0 + 23.0 285 + 10de 0 kao mecca El

Succulent Peas 73.4 + 17.6be 66.7 + 24.0 286 + 8d 29.8 + 0.9b

4 Means within a column followed by the same letter are not significantly different by Tukey’s test

at P > 0.05.

Mean development rates for larvae on weeds ranged from 25.1 days on bind-

weed to 44.3 days on hoary cress (Table 1). Other weeds supporting rapid de-

velopment of larvae were spiny sowthistle, dandelion, mallow, lambsquarter, and

Canadian thistle. Mean development rates for larvae on crop plants ranged from

24.7 days for potato to 31.3 days on dry peas (Table 1). Other crops supporting

rapid development were apple and pear. Among apple varieties tested, mean de-

velopment times were similar for Gala, Red Delicious, Fuji and Golden Delicious,

while development on Granny Smith foliage was significantly slower (Table 2).

Also of interest, mean development time for larvae on Fuji apple foliage was

25.1 + 0.7 days when evaluated in the first study and was then 38.2 + 1.0 days

when evaluated in the second study, along with other apple varieties. The first

evaluation used apple foliage collected during May, while the second evaluation

used apple foliage collected during July.

Table 2. Mean (+ SE) % pupation, adult emergence, pupal weight, and larval development times

for Lacanobia subjuncta on foliage of 5 apple varieties. Yakima, Washington, 2000.*

Pupal weight Larval time

Apple variety % Pupated % Adult (mg) (days)

Gala 40.0 + 11.5b 33.3 + 6.7 336 + 30ab 37.4 + 6.6bc

Red Delicious 73.3 + 6.7ab 73.3 + 6.7 387 + 9a 38.0 + 1.0bc

Fuji 73.3 + 13.3ab 60.0 + 11.5 357 + 8ab 38.2 + 1.0b

Golden Delicious 80.0 + 20.0a 60.0 + 11.5 321 + 12b 35.7 + 1.0bc

Granny Smith 66.7 + 13.3ab 50.0 + 10.0 284 + 22b 47.7 + 2.1a

4Means within a column followed by the same letter are not significantly different by Tukey’s test

at P > 0.05.

2002 LANDOLT: LACANOBIA SUBJUNCTA BIONOMICS 5

Mean pupal weights for L. subjuncta reared on weeds ranged from 249 mg on

Canada thistle to 436 mg on spiny sowthistle (Table 1). Other relatively heavy

mean pupal weights were 392 mg for larvae reared on dandelion, 374 mg for

larvae reared on bindweed, and 363 mg for larvae reared on mallow. In addition

to Canada thistle, pigweed and kochia yielded low pupal weights (Table 1). Mean

pupal weights for larvae fed crop plants ranged from 285 mg for dry peas to 377

mg for potato and 389 mg for Fuji apple (Table 1). When mean pupal weights

for apple varieties were compared, they ranged from 284 mg for larvae fed Gran-

ny Smith to 387 mg for larvae fed Red Delicious apple foliage (Table 2).

DISCUSSION

These results clearly indicate a broad potential plant host range for L. subjuncta

larvae and the potential for many plants in eastern Washington to contribute to

regionally high population densities contributing to crop losses. Complete devel-

opment from egg hatch to adult emergence was documented for 9 of the 10 weeds

tested and 7 of the 8 crops tested, with sunflower and apricot not supporting larval

development to pupation in this study. The plants selected for this study and

supporting L. subjuncta development are taxonomically diverse and include the

families Compositae, Cruciferae, Chenopodiaceae, Convulvulaceae, Malvaceae,

Solanaceae, Rosaceae, and Leguminaceae. Undoubtedly, many other plants pre-

sent in the region are probably equally suitable as hosts for L. subjuncta.

Just as the collection of larvae on a plant species is not proof that the species

is a good host plant, the demonstrations of survival and development to adult on

these plant species does not demonstrate that L. subjuncta utilizes these plants.

Additional information on patterns of adult female egg laying, of larval dispersal

and movement under field conditions, and of larval numbers on these and other

plant species would be more conclusive in assessing the potential of these plants

as hosts. Such studies clearly should incorporate not only common weeds but

additional crops that have not been reported to be infested with L. subjuncta.

There were differences in the performance of larvae on Fuji apple foliage col-

lected in May versus July that indicate possible seasonal variation in the suitability

of foliage of apple as food for L. subjuncta larvae. There are two broods of L.

subjuncta in Washington, with most larvae feeding in June/July and again in

August/September (Hitchcox 2000). Larvae reared on Fuji apple foliage in May

developed more rapidly and yielded larger pupae than larvae reared on Fuji apple

in July. Despite these possible differences in apple suitability as a host for L.

subjuncta, larvae are readily found on apple in the field during both time periods

(Hitchcox 2000). Such differences in host suitability could be due to a variety of

factors, such as accumulated leaf chemical defenses, increases in average leaf age,

accumulated responses to disease and herbivore challenges, and nutritional chang-

es in leaves. There is potential for much variation in foliage quality as food for

larvae for each of the plant species tested and care must be exercised in using

the data presented here for comparative purposes among plant species.

Also of concern is the potential impact of the apparent variance of plant suit-

ability as food for L. subjuncta larvae on phenology models used in IPM programs

for apple orchards. Such models may be used to predict L. subjuncta egg hatch

for the purpose of accurately timing pesticide applications. If larval development

rates are dependent in part on seasonal parameters affecting plant physiology,

6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

these must be incorporated into insect developmental models. Also, differential

rates of development on different host plants will contribute to variance in adult

emergence, oviposition, and egg hatch when multiple plants are used as hosts.

ACKNOWLEDGMENT

Technical assistance was provided by L. Biddick, J. Brumley, J. Dedlow, and

D. Lovelace. This work was supported in part by funding from the Washington

State Tree Fruit Research Commission.

LITERATURE CITED

Crumb, S. E. 1956. The larvae of the Phalaenidae. USDA Tech. Bull. No. 1135. Washington D.C.

Godfrey, G. L. 1972. A review and reclassification of the subfamily Hadeninae (Lepidoptera: Noc-

tuidae) of America north of Mexico. USDA Tech. Bull. 1450.

Hitchcox, M. E. 2000. Seasonal phenology and monitoring of Lacanobia subjuncta (Noctuidae: Lep-

idoptera) in apple orchards of Washington State. M.Sc. Thesis, Washington State University,

Pullman, WA. 75 pp.

Landolt, P. J. 1998. Lacanobia subjuncta (Lepidoptera: Noctuidae) on tree fruits in the Pacific North-

west. Pan-Pacific Entomol., 74: 32—38.

McCabe, T. L. 1980. A reclassification of the Polia complex for North America (Lepidoptera: Noc-

tuidae). N.Y. St. Mus. Bull. No. 432, Albany.

Rings, R. W., E. H. Metzler, E J. Arnold & D. H. Harris. 1992. The owlet moths of Ohio. Order

Lepidoptera, family Noctuidae. Ohio Biol. Survey Bull. (NS), 9: 1-184.

Warner, G. 1996. Goodbye codling moth, but hello cutworms. Good Fruit Grower, 47 (April 15):

17-18.

Warner, G. 1998. New pest causes havoc in orchards and warehouses. Good Fruit Grower, 49 (May

15): 28-30.

Received 29 June 2001; Accepted 16 Sept. 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 7-16, (2002)

A NEW SPECIES OF HETEROSPILUS (HYMENOPTERA:

BRACONIDAE) ASSOCIATED WITH THE DEATHWATCH

BEETLE, HEMICOELUS GIBBICOLLIS (LECONTE)

(COLEOPTERA: ANOBIIDAE)

BRIAN J. CABRERA!, PAUL M. MARSH?, VERNARD R. LEwISs?, &

STEVEN J. SEYBOLD*

'Rort Lauderdale Research and Education Center, University of Florida,

Fort Lauderdale, Florida 33314

7P. O. Box 384, North Newton, Kansas 67117

3Department of Environmental Science, Policy and Management,

University of California, Berkeley, California, 94720

‘Departments of Entomology and Forest Resources, University of Minnesota,

St. Paul, Minnesota 55108-6125

Abstract.—Heterospilus luridostigmus Marsh, a new braconid wasp species, is described. This

wasp was found in abundance emerging from pieces of Douglas-fir, Pseudotsuga menziesii (Mir-

bel) Franco, from an outdoor wooden deck in Daly City, California, that was infested with the

deathwatch beetle, Hemicoelus gibbicollis (LeConte). Adult Heterospilus luridostigmus began

emerging in May 1999 followed by emergence of Hemicoelus gibbicollis about 4 weeks later.

Both species continued to emerge throughout the summer of 1999 and were the only species

found in the boxes. During the summer of 2000, rearing boxes containing infested wood from

the original source and from several homes in Alameda County, California yielded both Heter-

os pilus luridostigmus and Hemicoelus gibbicollis as well as Odontocolon polymor phum Cushman

(Hymenoptera: Ichneumonidae). Although we found no direct evidence of parasitism of Hemi-

coelus gibbicollis, wasps in the genera Heterospilus and Odontocolon are known to parasitize

anobiids, and Heterospilus flavicollis (Ashmead) is a parasitoid of an eastern deathwatch beetle

species, Hemicoelus carinatus (Say). This suggests that Heterospilus luridostigmus is a parasitoid

of Hemicoelus gibbicollis. The synchrony of emergence of these two species during both years

also is indicative of a host/parasitoid relationship between the two species. The discovery of a

new insect species in a heavily populated urban environment is noteworthy and serves as a vivid

reminder of the untold number of insect species that have yet to be discovered.

Key Words.—Insecta, Braconidae, Anobiidae, Ichneumonidae, deathwatch beetle, parasitoid,

emergence hole.

The genus Heterospilus Haliday is a member of the braconid subfamily Do-

ryctinae. It can be identified by using the key to New World genera of Braconidae

(Marsh 1997). In North America the genus is easily distinguished from the other

doryctine genera by the reduction of fore wing vein 2RS (Fig. 1), which is always

desclerotized and often completely absent, and the presence of a stigma in the

hind wing of the male (not pictured). The small genus Pioscelus Muesebeck and

Walkley, which also has fore wing vein 2RS absent, is separated by having no

basal tubercle on the hind coxa. Heterospilus is the most species-rich genus in

the Doryctinae, with an estimated 200 species in the Nearctic Region and 300 in

the Neotropical Region. Most of these species are undescribed and the genus is

badly in need of revision.

All Heterospilus are idiobiont ectoparasitoids (Shaw & Huddleston 1991) and

this genus also has the most diversified host range in the Doryctinae. Species of

the genus parasitize a very wide range of endophytic, mostly stem-boring, hosts

8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

Figure 1. Wing venation of female Heterospilus luridostigmus, n. sp.

(Marsh 1982, Shaw 1995) including those in the coleopteran families Anobiidae,

Bostrichidae, Bruchidae, Buprestidae, Cerambycidae, Curculionidae, Languriidae,

Mordellidae and Scolytidae, and those in the lepidopteran families Gelechiidae,

Incurvariidae, Pyralidae and Tortricidae. Several other species have been reared

from other hosts including stem-boring symphytan Hymenoptera. A few species

are known to attack pemphredonine sphecid wasps (Marsh & Melo 1999).

In North America, five species have been recorded from Anobiidae, including

the new species described in this paper whose authorship is attributed solely to

P. M. Marsh. V. R. Lewis obtained beetle-infested wood was obtained by V. R.

Lewis. Wasps and beetles were reared and collected by V. R. Lewis and B. J.

Cabrera. S. J. Seybold and V. R. Lewis are principal investigators on a deathwatch

beetle, Hemicoelus gibbicollis (LeConte), pheromone project that includes the

work presented in this paper.

HETEROSPILUS LURIDOSTIGMUS MARSH, NEW SPECIES

Types.—Holotype, female. CALIFORNIA: SAN MATEO CoO.: Daly City, 393

Oriente St., Oct. 1998, V. R. Lewis, ex Pseudotsuga menziesii (Mirbel) Franco

boards from wooden deck. Deposited in NMNH, Washington, D.C.

Paratypes.—CALIFORNIA: same data as holotype, 41 females, 7 males. De-

posited in the California Academy of Sciences, San Francisco, NMUNH, Washing-

ton, D.C. and the University of Minnesota Insect Collection, University of Min-

nesota, St. Paul.

Description:—Female. Color: head, mesosoma and metasoma dark brown; scape, pedicel and basal

flagellomeres light brown, apical flagellomeres dark brown; fore and middle legs yellow, femora

marked with raised brown spot dorsomedially, tibiae marked with brown; hind leg with coxa and

femur brown, trochanters yellow, tibia yellow marked with brown, tarsus yellow; wings slightly dusky,

veins brown, stigma and vein C+SC+R light yellow, stigma sometimes nearly white. Body size: 2.5—

2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 9

4.0 mm. Head: face smooth and with dense long gold hair, frons and vertex transversely striate (Fig.

2), occasionally weakly so and nearly smooth; temple smooth; malar space two-thirds eye height;

ocell-ocular distance about 3 times diameter of lateral ocellus; occipital carina not meeting hypostomal

carina; 19—23 antennomeres. Mesosoma: pronotum rugose, with dense gold hair along weak pronotal

groove; mesonotal lobes (Fig. 3) coriaceous, rugose along notauli, notauli scrobiculate, meeting before

scutellum in rugose area with longitudinal short carinae, dense long gold hair along notauli; scutellum

smooth; mesopleuron smooth, subalar area carinate, sternaulus deep and longitudinally striate, dense

long gold hair on subalar area and along posterior edge; propodeum (Fig. 4) rugose, basal median

areas smooth, median carina and areola distinct, dense gold hair laterally. Legs: hind coxa with small

but distinct antero-ventral basal tubercle or tooth. Wings (Fig. 1): fore wing vein r as long as or

slightly longer than 3RSa, vein 2RS indicated by weak infuscate line, vein r-m not sclerotized but

distinct, vein m-cu arising distad from vein 2RS; hind wing vein M+CU longer than vein 1M, vein

m-cu a distinct infuscated line. Metasoma (Fig. 5): first tergum longitudinally carinate, length slightly

less than apical width, median raised area distinct, defined by carinae only on basal half; second

tergum longitudinally carinate; third tergum smooth with carinate area across basal half; remainder of

tergum smooth; ovipositor about two-thirds length of metasoma.

Male.—Essentially as in female; body size 1.5-3.0 mm; 17—20 antennomeres; hind wing with

elongate stigma.

Biology.—Associated with adults of Hemicoelus gibbicollis (LeConte) (Cole-

optera: Anobiidae) infesting Douglas-fir boards from a backyard deck. See details

of biology below.

Comments.—This species is distinctive by its light yellow to almost white stig-

ma in the fore wing and by vein M+CU in the hind wing being longer than vein

1M. Although this hind wing venation is not typical for the genus, in all other

characters the specimens are clearly congeneric. This species is easily distin-

guished from H. baeticatus (Provancher), H. flavicollis (Ashmead) and H. lon-

gicauda (Ashmead), which are also recorded as parasitoids of anobiids, by having

the ovipositor shorter than the metasoma. This species has a similar ovipositor

length to H. anobiidivorus Muesebeck but differs in the light stigma, completely

carinate second metasomal tergum, shorter first metasomal tergum and longer

antenna.

Etymology.—The specific name is from the Latin /uridus meaning pale yellow

in reference to the pale yellow or almost white stigma in the fore wing.

BIOLOGY AND OBSERVATIONS

Wood Collection.—Wood [Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco]

infested with the deathwatch beetle, Hemicoelus gibbicollis, was collected in Oc-

tober 1998 from the deck of a home in Daly City, San Mateo County, California.

The wood was cut into lengths of approximately 20—50 cm and stored at ambient

temperature and relative humidity in a greenhouse at the University of California,

Berkeley, in two wooden emergence boxes (122 * 122 * 122 cm) or in 58 liter

plastic storage boxes with 5-cm diameter screen-covered holes in each end for

ventilation. Wood moisture content was monitored with a moisture meter (Proti-

meter Timbermaster, Protimeter Ltd., Marlow Bucks, England) and the wood was

watered as needed to maintain approximately 14—17% moisture content (Suomi

& Akre 1992a, b). Additional pieces of infested wood (predominantly P. menzie-

sii) were collected in October, 1999 from several homes in Alameda County,

California and kept outdoors on the premises of the University of California,

Forest Products Laboratory, Richmond, California. In June 2000, this wood was

also cut and stored in plastic storage boxes as previously described.

THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

Figures 2—5. Morphological characters of Heterospilus luridostigmus, n. sp. Figure 2. Vertex and

frons. Figure 3. Mesonotum. Figure 4. Propodeum. Figure 5. Metasomal terga 1-4.

2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 11

Beetle and Wasp Emergence, 1999.—Boxes were checked occasionally for

adult Hemicoelus gibbicollis (needed for extracting sex pheromone and for be-

havioral assays) beginning in April 1999. Heterospilus luridostigmus appeared

unexpectedly in the wooden emergence boxes on 3 May and continued to emerge

through 2 Aug (Fig. 6A). The exact date of initial emergence is unknown because

the boxes were not examined on a regular basis. The first Hemicoelus gibbicollis

adults were found on 28 May with the exact date of emergence also unknown.

The last adults were collected on 16 August. This emergence was in agreement

with Suomi & Akre (1993a, b) who stated that normal emergence occurs during

June, July, and August. We collected a total of 584 beetles (336 alive, 57.5%

survival) and 179 wasps.

Beetle and Wasp Emergence, 2000.—Beetles and wasps were first collected on

15 June. The number of emerged adults of both species was considerably lower

than in 1999 (Figs. 6A and 6B), possibly because of the different collection

source, differences in the host wood, or because of exposure to sub-optimal en-

vironmental conditions prior to rearing. The wood had been kept outdoors for

seven months before it was cut and placed in the rearing boxes. A total of 218

beetles (132 alive, 60.6% survival) and 32 wasps were collected. Several adult

male and female Odontocolon polymorphum Cushman (Hymenoptera: Ichneu-

monidae) were collected in addition to Heterospilus luridostigmus.

Possible Parasitism of Hemicoelus gibbicollis.—The doryctine b raconids are

generally considered to be ectoparasitoids of wood-boring beetle larvae (Marsh

1979). However, Suomi & Akre (1992a, b, c; 1993a, b) did not mention parasit-

oids in their detailed descriptions of Hemicoelus gibbicollis biology and ecology.

Furthermore, we did not directly observe wasps emerging from any life stage of

Hemicoelus gibbicollis or from the wood and dissection of several small pieces

of wood did not yield any parasitized larvae. The need for large numbers of adult

H. gibbicollis made us reluctant to conduct a more thorough search that would

have required destruction of more beetle-infested wood. However, we believe that

Heterospilus luridostigmus is a parasitoid of Hemicoelus gibbicollis because: 1)

adults of both species appeared in our rearing boxes in relative synchrony during

both years; 2) a broad range of hole diameters was observed on the surface of

the infested wood; and 3) other species of Heterospilus are idiobiont ectoparasi-

toids of wood-destroying anobiids (e.g., Heterospilus flavicollis (Ashmead) on

Hemicoelus carinatus (Say) [Drooz 1985], the most common wood-infesting

deathwatch beetle in the northeastern United States [Simeone 1962], and Heter-

ospilus longicauda (Ashmead) on Xyletinus peltatus (Harris) [Williams et al.

1979]).

The only two live insect species to appear in 1999 in our rearing boxes were

Heterospilus luridostigmus and Hemicoelus gibbicollis. Both species were col-

lected again in 2000 from rearing boxes containing wood that was collected both

years. In 1999, adult Heterospilus luridostigmus were first observed approxi-

mately four weeks before the first emergence of Hemicoelus gibbicollis while the

following year both species were first found on the same day. Peak emergence

of both species was nearly synchronous in 1999, with the largest number of wasps

emerging approximately two weeks before the largest number of beetles emerged

(Fig. 6A). A similar trend was observed the following summer (Fig. 6B). The

observation that Heterospilus luridostigmus preceded Hemicoelus gibbicollis in

12 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

= 80

S i

- Ais

: 3

S&S 50 mi

— =

- =]

2 | 5

= 5

$ | :

s | :

2 4

> V\ =

: call i 1 Yeo

1-May 16-May 31-May 15-Jun 30-Jun 15-Jul 30-Jul 14-Aug

Date

H. gibbicollis (Total number)

(49quINU |/2}01) sdsemM

1-May 16-May 31-May 15-Jun 30-Jun 15-Jul 30-Jul 14-Aug

Date

Figure 6. Emergence of beetles and wasps. A. Hemicoeltus gibbicollis (LeConte) (bars), and Het-

erospilus luridostigmus Marsh (line), summer 1999. [] Live beetles, Hl Dead beetles, A Live wasps.

B. Hemicoelus gibbicollis (LeConte)(bars), and Heterospilus luridostigmus Marsh and Odontocolon

polymor phum Cushman (line), summer 2000. L] Live beetles, Hl Dead beetles, A Live wasps.

2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 13

emergence in 1999 but not in 2000 might be attributed to laboratory worker

inexperience, as both species are difficult to find amongst the pieces of wood in

the rearing boxes. Hemicoelus gibbicollis is especially difficult to locate because

emerged adults spend large periods of time seeking refugia in emergence holes.

We also expected a much larger emergence of Hemicoelus gibbicollis during the

summer of 2000. Dissection of several infested pieces of wood in mid-August of

that year yielded beetle larvae of mixed size and, presumably, age, indicating that

the infestations were still active. We speculate that the lower number of emerged

beetles and the shorter emergence period during the second year were partly a

result of parasitism. Williams et al. (1979) found that Heterospilus longicauda

(Ashmead) accounted for 1.3—36.6% mortality of the potential population of their

anobiid host, X. peltatus.

We frequently observed small emergence holes in the infested wood, presum-

ably made by Heterospilus luridostigmus, adjacent to much larger and more abun-

dant emergence holes, presumably made by Hemicoelus gibbicollis (Fig. 7) (Hem-

icoelus gibbicollis is considerably larger than Heterospilus luridostigmus in cross-

sectional area). Williams et al. (1979) reported that Heterospilus longicauda emer-

gence holes were about one-eighth the size of X. peltatus emergence holes, that

the wasp oviposited through the wood and onto the host larvae, and assumed

there was one parasite larva per host. However, we measured holes (n = 434)

from several pieces of wood collected in 1999 and instead of an expected bimodal

distribution we obtained an approximately normal distribution ranging from 0.46

to 2.15 mm in diameter (Fig. 8). The lack of a distinct separation among emer-

gence hole diameters prevents us from reporting species-specific emergence hole

size ranges at this time. In contrast to Williams et al. (1979), our minimum emer-

gence hole sizes all exceeded one-eighth of the maximum hole sizes. Our sample

of emergence holes probably also contained holes made by O. polymorphum.

The appearance of O. polymorphum from our rearings in 2000 reveals the

possible existence of other parasitoids of Hemicoelus gibbicollis. The wood in the

rearing boxes had been lying outdoors for seven months, thus making larvae and

pupae of H. gibbicollis within the wood readily accessible to attack by Heteros-

pilus luridostigmus and other opportunistic natural enemies. Alternatively, Het-

erospilus luridostigmus and O. polymorphum may have located and colonized

Hemicoelus gibbicollis while the wood was still in the structures from which it

was removed. Odontocolon polymorphum has been collected in Oregon, Wash-

ington, and British Columbia and has been associated with two wood-infesting

anobiid species (one unidentified, the other Ptilinus basalis LeConte) (Carlson

1979).

This paper represents the first report of O. polymorphum associated with Hem-

icoelus gibbicollis. Ovipositor length and examples of other xoridine ichneumon-

ids (Townes et al. 1960) suggest that O. polymorphum is ectoparasitic and ovi-

posits through the wood surface onto larvae and pupae of H. gibbicollis.

Whether H. gibbicollis is an obligate or facultative host for both wasp species

remains to be determined. Although there is strong evidence for a host/parasitoid

relationship between Hemicoelus gibbicollis and Heterospilus luridostigmus, di-

rect observation of wasp oviposition behavior, emergence of adult wasps from

the host or infested wood, detection of parasitized beetle larvae or larval beetle

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 78(1)

Adult emergence hole of Hemicoelus gibbicollis (LeConte) (A) and suspected emergence hole of Heterospilus luridostigmus Marsh (B). Bar = 1 mm.

Figure 7.

2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 15

100

® 60

2 40

0.46 0.62 0.77 0.92 1.08 1.23 1.38 1.54 1.69 1.85 2.00 2.15

Emergence hole diameter (mm)

Figure 8. Size distribution of emergence holes (n = 434) of Hemicoelus gibbicollis, Heterospilus

luridostigmus, and Odontocolon polymorphum measured from several pieces of infested Pseudotsuga

menziesii, collected in 1999 in Alameda Co., California.

head capsules present in wasp cocoons is needed to confirm the existence of this

proposed ecological relationship.

Finally, it is noteworthy that a new insect species has been found in an urban

environment. This discovery, in a heavily populated area and in close association

with human dwellings, is a vivid reminder of the tremendous diversity of the

Insecta and of the untold number of insect species that have yet to be discovered.

ACKNOWLEDGMENT

We thank G. Chow, L. Daniels, S. Garcia-Rubio & R. Raban for their assistance

in the collection and processing of wood and the collection of wasps and beetles.

We also thank D. Carver, Live Oak Structural Pest Control, Berkeley, California

and S. Kala, Daly City, California for providing the infested wood. Dr. E An-

drews, California Department of Food & Agriculture, Sacramento, California,

confirmed the identity of Hemicoelus gibbicollis. Dr. J. Luhman, Minnesota De-

partment of Agriculture, St. Paul, Minnesota identified Odontocolon polymorphum

and provided very useful information on the xoridine Ichneumonidae. Specimens

of O. polymorphum and H. gibbicollis from this study were deposited in the

University of Minnesota Insect Collection and specimens of H. gibbicollis were

deposited in the California Academy of Sciences. The scanning electron micro-

graphs were prepared by K. Hampton, Department of Entomology, Kansas State

University, Manhattan, Kansas. Figure 7 was taken by D. C. Blackford, University

of Minnesota, Department of Entomology. We thank R. A. Wharton and an anon-

ymous reviewer for their comments and suggestions on the manuscript. The work

16 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

described in this paper was supported by California Department of Consumer

Affairs, Structural Pest Control Board grant 849017-07 to VRL and SJS.

LITERATURE CITED

Carlson, R. W. & B. D. Bunks. 1979. Family Ichneumonidae. pp. 315-740. Jn Krombein, K. V., P. D.

Hurd Jr. & D. R. Smith (eds.). Catalog of Hymenoptera in America North of Mexico. Volume

1. Smithsonian Press, Washington D.C.

Drooz, A. 1985. Insects of eastern forests. USDA Forest Service, Misc. Pub. No. 1426.

Marsh, P. M. 1979. Family Braconidae. pp. 144-294. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith

& B. D. Burks. (eds.). Catalog of Hymenoptera in America North of Mexico. Volume I. Smith-

sonian Press, Washington, D.C.

Marsh, P. M. 1982. Two new species of Heterospilus (Hymenoptera: Braconidae) from Mexico being

introduced against the cotton boll weevil, Anthonomus grandis (Coleoptera: Curculionidae).

Proc. Ent. Soc. Wash., 84: 849-854.

Marsh, P. M. 1997. Subfamily Doryctinae. pp. 207—233. In Wharton, R. A., PB. M. Marsh & M. J.

Sharkey (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera).

Special Pub. Int. Soc. Hymenopterists No. 1.

Marsh, P M. & G. A. R. Melo. 1999. Biology and systematics of New World Heterospilus (Hyme-

noptera: Braconidae) attacking Pemphredoninae (Hymenoptera: Sphecidae). J. Hym. Res., 8:

13-22.

Shaw, S. R. 1995. Braconidae. pp. 431—463. In Hanson, P. & I. Gauld (eds.). The Hymenoptera of

Costa Rica. Oxford University Press.

Shaw, M. R. & T. Huddleston. 1991. Classification and biology of braconid wasps (Hymenoptera:

Braconidae). Handbooks for the Identification of British Insects 7: 1-126.

Simeone, J. B. 1962. Survey of wood-feeding Anobiidae in northeastern United States, including a

study of temperature and humidity effects on egg development of Hadrobregmus carinatus

(Say). pp. 326-335. 11th Int. Kongr. Entomol., Wien. Aug. 17—25 1960. Verh. Band II.

Suomi, D. A. & R. A. Akre. 1992a. Characteristics of structures attacked by the wood-infesting beetle,

Hemicoelus gibbicollis (Coleoptera: Anobiidae). J. Entomol. Soc. Brit. Columbia, 89: 63-70.

Suomi, D. A. & R. A. Akre. 1992b. Control of the structure-infesting beetle Hemicoelus gibbicollis

(Coleoptera: Anobiidae) with borates. J. Econ. Entomol., 85: 1188-1193.

Suomi, D. A. & R. A. Akre. 1992c. Distribution of economically important, wood-infesting anobiid

beetles in the Pacific Northwest. J. Entomol. Soc. Brit., Columbia, 89: 57-62.

Suomi, D. A. & R. A. Akre. 1993a. Biological studies of Hemicoelus gibbicollis (LeConte)(Coleoptera:

Anobiidae), a serious structural pest along the Pacific coast: adult and egg stages. Pan-Pacific

Entomol., 69: 155-170.

Suomi, D. A. & R. A. Akre. 1993b. Biological studies of Hemicoelus gibbicollis (LeConte)(Coleoptera:

Anobiidae), a serious structural pest along the Pacific coast: larval and pupal stages. Pan-Pacific

Entomol., 69: 221-235.

Townes, H. M., S. Walley, L. Walkley, D. Habeck & G. Townes. 1960. Ichneumon flies of America

north of Mexico. Vol. 2. Ephaltinae, Xoridinae, and Acaenitinae. USNM Bulletin No. 216, Part

I, Washington, D.C.

Williams, L. H., H. M. Barnes & H. O. Yates, III. 1979. Beetle (Xyletinus peltatus) and parasite exit

hole densities and beetle larval populations in southern pine floor joists. Environ. Entomol., 8:

300-303.

Received 4 April 2001; Accepted 16 August 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 17-22, (2002)

A NEW SPECIES OF YELICONES CAMERON

(HYMENOPTERA: BRACONIDAE) FROM THAILAND

BUNTIKA AREEKUL!” & DONALD L. J. QUICKE!

'Department of Biological Sciences, Imperial College at Silwood Park, Ascot,

Berkshire, SL5 7PY, UK

*Department of Entomology, The Natural History Museum, South Kensington,

London, SW7 5BD, UK

Abstract.—Yelicones siamensis Areekul & Quicke, NEW SPECIES is described and illustrated

based on two adult females collected at light in Thailand. This wasp is the ninth species of

Yelicones described from the East Palaearctic and Oriental regions. A modification to the key

of Quicke et al. (1997: J. Nat. Hist. 31: 779-797) is included to differentiate Y. siamensis from

similar species.

Key Words.—Insecta, Hymenoptera, Braconidae, Yelicones, Thailand.

Wasps of the genus Yelicones Cameron are solitary endoparasitoids of lepi-

dopteran larvae, whose remains they mummify before pupating within the host

(Quicke & Chishti 1997). For many years after its original description (Cameron

1887) the genus was known only from a handful of specimens from the New

World (Shenefelt 1975, Quicke & Kruft 1995). However, over the last 20 years

a number of new species have been described, extending the known range of

Yelicones into the Indo-Australian, Afrotropical and Palaearctic regions. The ge-

nus is now known to be widely distributed throughout the Old and New Worlds

(Fischer 1961, 1962 [as Pectenopius Fischer]; Togashi 1980; Papp 1985, 1989,

1991, 1992; Belokobylskij 1993a, b; Quicke & Kruft 1995; Quicke et al. 1996,

1997, 1998; Quicke & Chishti 1997; Shaw 1998).

In this paper a new species of Yelicones is described based on two female

specimens from Thailand, the ninth for the East Palaearctic and Oriental regions

(Quicke et al. 1997). It is being described because it has been used to generate

DNA sequence data which will be published elsewhere as part of another study.

Male morphology and biology are unknown. The genus Yelicones can be recog-

nized using the keys of van Achterberg (1995), Chen and He (1997) or Shaw

(1997). A brief diagnosis is provided below.

MATERIALS AND METHODS

Two specimens were collected by light trapping in Chon Buri, Thailand and

preserved in absolute alcohol. Three legs on one side of the body were taken

from the paratype specimen for DNA sequencing and both specimens were then

mounted for description and photography. Measurements were made with an eye-

piece micrometer graticule. Terminology follows van Achterberg (1979, 1988)

and Quicke et al. (1997).

Genus Yelicones Cameron, 1887

Yelicones Cameron 1887: 387; van Achterberg, 1995: 147 (literature). Type spe-

cies, Yelicones violaceipennis Cameron, designated by Viereck (1914).

* Author for correspondence

18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

Figure 1. Yelicones siamensis NEW SPECIES, female holotype.

Rhopalotoma Cameron 1911: 318. Type species, Rhopalotoma crassitarsis Cam-

eron, monotypic.

Pectenopius Fischer 1961: 156. Type species, Pectenopius paradoxus Fischer,

original designation.

YELICONES SIAMENSIS AREEKUL & QUICKE, NEW SPECIES

Types.—Holotype, female (Fig. 1); data: THAILAND. CHON BURI: Khao

Kheow, 20-31 March 2001, D. L. J. Quicke and N. Laurenne, light trap; depos-

ited: British Museum (Natural History). Paratype: same data as holotype, 1 fe-

male; deposited Insect Collections of Chulalongkorn University, Bangkok, Thai-

land.

Description —Female (holotype) Length. Body 4.5-5.0 mm, and fore wing 3.5 mm. (Fig. 1). Color.

Yellow, antennae yellow basally, gradually brown on distal 0.4; wing veins dark brown, pterostigma

basal 0.4 ivory, distal 0.6 dark brown (Fig. 1). Head. Antennae with 26 flagellomeres, terminal fla-

gellomere pointed, approximately 2.3 longer than wide; first flagellomere 1.1 X and 1.4X longer than

the second and third respectively; first flagellomere 1.4 longer than wide; third flagellomere as long

as wide; malar space unsculptured, length of malar space 0.04 height of eye; height of clypeus:

inter-tentorial distance: tentorio-ocular distance = 1.0: 3.4: 0.8; clypeus slightly punctate, with long,

dense setae; face with subtransverse carinae below the antennal sockets, punctate ventrally (Fig. 2),

densely covered with long setae, with weak but distinct mid-longitudinal ridge (Fig. 3); height of eye:

width of face: width of head = 1.0: 0.9: 1.7; length of face = 0.5X width of face; eyes glabrous;

frons with sparse, long setae, impressed behind the antennal socket, mid-longitudinal ridge strongly

developed, posteriorly with two curved transverse carinae; occiput and temples densely punctate;

horizontal length of eye: horizontal length of head behind eye = 1.6: 1.0; post-ocellar length: trans-

2002 AREEKUL & QUICKE: A NEW YELICONES FROM THAILAND 19

Figures 2-6. Yelicones siamensis NEW SPECIES. Figure 2, front view of head. Figure 3, front

and lateral aspect of head. Figure 4, mesoscutum. Figure 5, lateral view of prothorax. Figure 6,

propodeum.

20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

= ‘

a ss

’ ae ee

Py..- ‘oe be

- Umer sre

‘ we se we Hog

Figures 7-8. Yelicones siamensis NEW SPECIES. Figure 7, fore wing. Figure 8, dorsal view of

metasomal tergites 1-3.

verse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.0: 2.0:

2.7; occipital carina nearly complete, absent for a small distance medially. Mesosoma. Shiny, densely

punctured and setose, 1.8 longer than high; mesoscutum postero-medially with longitudinal groove-

like impressions (Fig. 4); notauli weakly impressed throughout length of mesoscutum; scutellar sulcus

with 7 carinae between the two outer ones; scutellum shiny and sparsely setose with small punctures;

median area of metanotum medially without pit (Fig. 6); mesopleuron densely setose, with transverse

carinae anteriorly, densely punctate posteriorly; precoxal suture weakly impressed, crenulate, upcurved

posteriorly, impressed 0.8 length of mesopleuron (Fig. 5); propodeum strongly aerolate-rugose, antero-

medially without a prominent U- or V-shaped carina (Fig. 6). Wings. Fore wing—length of veins SR1:

3SR: r = 3.3: 0.4: 1.0; vein 1-SR+M more or less straight; vein r arising 0.5 distance from base of

pterostigma; length of veins 2-SR: 3-SR: r-m = 1.0: 1.0: 1.1; length of veins 2-SR+M: 2-M: m-cu =

1.0: 0.7: 0.7; length of veins 2-CU1: 3-CU1 = 2.4: 1.0; veins C+SC+R and 1-SR forming an angle

of 60° (Fig. 7). Hind wing—length of veins lr-m: SC+R1 = 1.0: 1.8; vein 2-SC+R interstitial; vein

SR posteriorly weak, more or less straight at apex; vein 2m-cu strongly postfurcal, length of vein 1M

4.4X vein 2M, vein 2m-cu more or less straight; marginal cell, basal cell and base of wing densely

setose. Legs. Length of fore femur: tibia: tarsus = 1.0: 1.3: 1.0; fore femur 2.0X longer than maximum

depth; fore tibia without mid-longitudinal ridge; hind femur 2.7X longer than maximum depth; length

of hind femur: tibia: basitarsus = 1.8: 2.5: 1.0; hind basitarsus 3.4 longer than maximally depth.

2002 AREEKUL & QUICKE: A NEW YELICONES FROM THAILAND 21

Metasoma. Metasomal tergites shiny, first and second tergite with sparse setae, 3rd—8th tergites mod-

erately setose; first and second tergite with punctate-rugulose sculpture; first metasomal tergite 1.3 X

wider than medially long, dorsal carina weakly impressed, uniting before the level of spiracles; second

metasomal tergite 2.4 wider than medially long, without smooth triangular area anteriorly and with-

out mid-longitudinal carina; second suture narrow, smooth; third metasomal tergite 2.4 wider than

medially long; third tergite anteriorly finely punctate, posteriorly smooth (Fig. 8); 4th—6th metasomal

tergites smooth. Tip of ovipositor pale.

Diagnosis.—Yelicones siamensis Areekul & Quicke keys out to couplet 8 using

the key to East Palaearctic and Oriental species of Yelicones (Quicke et al. 1997).

It can be distinguished from Y. flavus Chen and Quicke by the following char-

acters: face not transversely imbricate; mesoscutum postero-medially rugose; fore

wing vein 1-SR+M more or less straight not sinuous; hind wing vein 2-SC+xR

interstitial not transverse; dorsal carinae of first metasomal tergite uniting anteri-

orly not near mid-length, without median carina; second metasomal tergite ante-

rior-medially without smooth triangular area and mid-longitudinal carina; color

yellow, wings without distinct pattern of brownish blotches, pterostigma bicolored

not unicolorous.

Modification to the key to the species of Yelicones of the East Palaearctic and

Oriental region (Quicke et al. 1997) to accommodate the new species.

8(7). Mesoscutum with line of notauli piceous on anterior two thirds and

usually with a dark line connecting them in front of the posterior

margin; propleuron and fore coxae whitish ......... Y. koreanus Papp

- Mesoscutum uniformly yellow-brown; propleuron and fore coxae

brownish yellow

8a(8). Forewing with distinct pattern of brownish blotches; pterostigma uni-

formly, pale brown; antennae with 33 flagellomeres; second meta-

somal tergite antero-medially with smooth triangular area ..........

ees, cee Nea Teiticie gat Avot dB MN SORES dtath git cdc Y. flavus Chen and Quicke

- Forewing without distinct pattern of brownish blotches; pterostigma

bicolored, whitish basally, brown distally; antennae with fewer than

33 flagellomeres; second metasomal tergite antero-medially without

smooth triangular area ............... Y. stamensis NEW SPECIES

Variation in Females.—Body length 4.5—6.0 mm; antennae with 26—28 flagel-

lomeres; mesoscutum postero-medially with punctate-rugose sculpture or longi-

tudinal groove-like impressions; scutellar sulcus with 6—7 carinae between the

outer ones.

Male.—Unknown.

Distribution.—Thailand.

Etymology.—The name is derived from the old name for Thailand.

Material Examined.—See Types.

ACKNOWLEDGMENT

We thank Angoon Lewvanich and Sura Pimpasalee for help in collecting the

specimens, and Apidet Singhaseni for permitting collecting at Khao Kheow Zoo.

Robert Belshaw, Gavin Broad and David Orme reviewed the manuscript and

provided helpful comments. Andrew Polaszek helped with Automontage imaging

facilities.

22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

LITERATURE CITED

Achterberg, C. van. 1979. A revision of the subfamily Zelinae auct. (Hymenoptera, Braconidae).

Tijdschr. Ent., 122: 241-479.

Achterberg, C. van. 1988. Revision of the subfamily Blacinae Foerster (Hymenoptera, Braconidae).

Zool. Verh., Leiden, 249: 1-324.

Achterberg, C. van. 1995. Generic revision of the subfamily Betylobraconinae (Hymenoptera: Bra-

conidae) and other groups with modified fore tarsus. Zool. Verh., Leiden, 298: 1-242.

Belokobylskij, S.A. 1993a. New taxonomic data on the braconid fauna (Hymenoptera, Braconidae) of

Vietnam. Russian Entomol. J., 2: 37—67.

Belokobylskij, S.A. 1993b. Contribution to the taxonomy of Braconidae (Hymenoptera) of the Russian

Far East. Russian Entomol. J., 2: 87-103.

Cameron, P. 1887. Family Braconidae, Biologia Centrali-Americana. Hymenoptera, 1: 312-419.

Chen, X. & J. He. 1997. Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China.

Zool. Verh., Leiden, 308: 1-187.

Fischer, M. 1961. Zwei neue Opiinen Gattungen (Hym., Braconidae). AnnIn naturh. Mus. Wien, 64:

154-158.

Fischer, M. 1962. Die Opiinae des Museo Civico di Storia Naturale in Genua (Hymenoptera, Bracon-

idae). Annali. Mus. Civ. Stor. Nat. Giacomo Doria, 73: 71-97.

Papp, J. 1985. Braconidae (Hymenoptera) from Korea, 7. Acta Zool. Hung., 31: 341-365.

Papp, J. 1989. A contribution to the braconid fauna of Israel. Israel J. Ent., 22: 45-59.

Papp, J. 1991. New braconid wasps (Hymenoptera: Braconidae) in the Hungarian Natural History

Museum, 2. Annls hist.-nat. Mus. Natn. hung., 83: 145-167.

Papp, J. 1992. New braconid wasps (Hymenoptera: Braconidae) in the Hungarian Natural History

Museum, 3. Annls hist.-nat. Mus. Natn. hung., 84: 129-160.

Quicke, D. L. J. & R. A. Kruft. 1995. Species of Yelicones (Hymenoptera: Braconidae: Rogadinae)

in North America with descriptions of two new species. Ann. Entomol. Soc. Am., 88: 129-—

138.

Quicke, D. L. J.. M. J. K. Chishti, & H. H. Basibuyuk. 1996. A revision of the Yelicones species

(Hymenoptera: Braconidae: Rogadinae) from Central America, with descriptions of sixteen new

species. Zool. Meded. Leiden, 70: 17-61.

Quicke, D. L. J. & M. J. K. Chishti. 1997. A revision of the Yelicones species (Hymenoptera: Bra-

conidae: Rogadinae) from Africa and the Arabian Peninsula, with descriptions of four new

species. African Entomol., 5: 77-91.

Quicke, D.L.J.,M.J.K.Chishti, X. Chen, & R.A. Kruft. 1997. Revision of Yelicones (Hymenoptera:

Braconidae: Rogadinae) from the East Palaearctic and Oriental regions with description of four

new species. J. Nat. Hist., 31: 779-797.

Quicke, D. L. J., A. D. Austin, & M. J. K. Chishti. 1998. Revision of Yelicones (Hymenoptera:

Braconidae: Rogadinae) from the Australasian region. Invert. Taxon., 12: 897-928.

Shaw, M. R. 1998. The surprising discovery of the genus Yelicones Cameron (Hymenoptera: Bracon-

idae) in Western Europe. Br. J. Ent. Nat. Hist., 11: 15-16.

Shaw, S. R. 1997. Subfamily Rogadinae s.s. pp. 403—412. In Wharton, R. A., Marsh, P. M. & Sharkey,

M. J. (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera). Special

publication of the International Society of Hymenopterists, Number 1, Washington, D.C.

Shenefelt, R. D. 1975. Braconidae 8, Exothecinae and Rogadinae. Part 12. pp. 1115-1262. In J. van

der Vecht & R. D. Shenefelt (eds.). Hymenoptera Catalogus, Junk, The Hague.

Togashi, I. 1980. Discovery of the genus Yelicones Cameron (Hymenoptera, Braconidae) from Japan.

Kontyu, 48: 571-520.

Received I] July 2001; Accepted 23 November 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 23-33, (2002)

THE SPIDER FAUNA ASSOCIATED WITH LITTER

UNDER WOODRAT MIDDENS IN SOUTHERN

CALIFORNIA (ARACHNIDA: ARANEAE)

RICHARD S. VETTER & THOMAS R. PRENTICE

Department of Entomology, University of California,

Riverside, California 92521

Abstract.—Litter from under wood rat (Neotoma sp.) middens from southern California, was

sampled, primarily between 1977-1985, in search of latridiid beetles. As an ancillary by-product,

spiders were collected, separated and labeled with collection data but most have remained un-

determined until now. In this paper, we determine the spider species associated with the rat

middens, predominantly from five southern Californian counties, ranging in elevation from 200

to 1900 m. This study yielded 316 specimens representing 42 species, 34 genera and 20 families

of spiders. The family Linyphiidae was represented here with the greatest number of genera,

species and specimens. The most frequently collected species, Tapinocyba dietrichi Crosby &

Bishop (Linyphiidae), contributed 37.3% of the specimen total. Spirembolus erratus Millidge

(Linyphiidae) and Zanomys californica (Banks) (Amaurobiidae) were the second and third most

predominant species, contributing 16.5% and 5.1%, respectively, of the specimen total. Many of

the specimens identified in this study are members of minute species (1-3 mm), which can easily

be overlooked unless samples are carefully scrutinized. Some of the more interesting and un-

common species include Trogloneta paradoxum Gertsch (Mysmenidae), Gertschanapis shantzi

(Gertsch) (Anapidae), several Zanomys spp., and members from the families Hahniidae, Capon-

iidae and Oonopidae. Specimens collected here will contribute to the description of three new

species (2 amaurobiids, 1 linyphiid) and the male of a second linyphiid species.

Key Words—Arachnida, Neotoma, rat middens, species list, Araneae, spiders.

Woodrats of the genus Neotoma create large middens composed of vegetation

and other materials culled from the surrounding environment (English 1923). Mid-

dens can reach massive sizes, primarily through occupancy by successive gener-

ations with each occupant adding material. In so doing, the rats create a micro-

habitat that often differs from the immediate surrounding area in various biotic

and abiotic aspects (Vestal 1938, Linsdale & Tevis 1951). Dimensions of 301

middens in northern California averaged 118 cm in height and 152 cm in basal

diameter; the average volume of 572 middens was 0.713 m? (Vestal 1938). Active

rat middens are strewn with fresh food cuttings and copious fecal pellets (Linsdale

& Tevis 1951). Considering their structural features, middens provide ideal har-

borage for a plethora of animals including spiders (Linsdale & Tevis 1951). Fossil

midden remains have offered a wealth of data on the historical plant and arthropod

components of the regions of occupancy (Hall et al. 1989, Elias et al. 1992, Clark

& Sankey 1999) although, in these studies, arachnids were represented solely by

the hard-bodied ticks, scorpions and pseudoscorpions.

From 1977 to 1985, Ken Cooper of the University of California-Riverside

(UCR) harvested arthropod inhabitants from the litter under Neotoma nests in

search of beetles of the family Latridiidae. This litter was collected primarily from

five southern Californian counties in a variety of habitats ranging from the Los

Angeles-San Diego basins (ca. 300 m elevation) to the surrounding inland and

coastal mountains (up to 1900 m). An ancillary by-product of the collections

resulted in an accumulation of spiders which, until now, have languished in the

24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

UCR Entomology museum, mostly as undetermined specimens. We have identi-

fied these spiders, the majority of which were less than 3 mm in body length,

and present our findings here.

Minute spiders pose several problems that impede identification. Size alone

creates difficulties in manipulation of the specimens as well as in distinguishing

the pertinent diagnostic characters. Most of the minute species determined in this

study are members of the Linyphiidae which contains a cumbersome number of

genera and species that are often very difficult to identify. Relatively few generic

keys and current taxonomic revisions are available for this family; those that do

exist often include only regional fauna. One of the most current and definitive

identification guides for North American spiders (Roth 1993) presents keys to

genera for every family except the Linyphiidae which are simply listed alpha-

betically by genus for both the linyphiine (45 genera) and erigonine (117 genera)

spiders. What we hope to accomplish here is to assist future investigators of

Neotoma biology, add to our knowledge of Californian linyphiid distribution, and

encourage eager arachnologists to pursue taxonomic studies on the little-known

genera.

MATERIALS AND METHODS

All information regarding the history of the spiders from the Neotoma middens

was derived from personal communications with the collector (K. Cooper), the

processor (KF Andrews) of the midden material, and the former curator (S. From-

mer) of the Entomology museum collection (all former associates of UCR). Ap-

proximately 100 Neotoma (mostly N. fuscipes with afew N. lepida) middens were

sampled. The litter under each nest was shoveled into canvas bags, brought back

to the laboratory and processed in approximately 12 gallon loads, for up to 10

days, in Berlese funnels until no additional arthropods were collected. Although

most of the spiders were part of the undetermined UCR Entomology Research

Museum spider collection, it was not known how many spiders were actually

identified and incorporated as determined species. Consequently, we searched the

entire general spider collection (> 1700 vials) excepting the orb-weaver families

Araneidae, Tetragnathidae, Uloboridae, the adults of which we felt would not

likely be using the litter under rat middens as refugia. As a result of our search,

we believe that this study represents an accurate inventory of the material recov-

ered by the researchers.

Spiders were present in samples from at least 50 Neotoma middens collected

by Cooper in southern California. Also included are data from one midden from

San Bernardino County, California collected by Andrews, Hardy & Ejichlin in

1985. Elevations not present on the collection labels were approximated to the

nearest 25 meters with topographic maps or through recent communications with

K. Cooper and should be accurate to within 50 meters. Spiders are listed in Table

1 in decreasing frequency by family to show the importance of their ecological

association with Neotoma nests and then alphabetically by genus within each

family; families with equal numbers of specimens are grouped alphabetically.

All specimens have been incorporated into the reference collection of the UCR

Entomology Research Museum.

Table 1. List of the spiders identified from Neotoma rat nests.

Elevation

CcOOT

Family/Genus Species Male Fem Imm County Date (meters) Locale

Linyphiidae

Ceratino ps inflatus 2 2 Riverside 14-Jan-83 1800 0.8 km E of James Reserve, San Jacinto Mts

Ceratinopsis interventa 1 Kern 23-Sep-81 800 Red Rock Cyn campsite

“Ceratinopsis”’ palomara 1 Riverside 30-Apr-81 750 Bautista Cyn, 27.4 km SE of Valle Vista

palomara 2 Riverside 29-Apr-82 300 Clinton Keith Rd, W Murrieta near entrance to cyn

palomara 1 5 Riverside 14-Jan-83 1825 0.8 km E of James Reserve, San Jacinto Mts

Linyphantes aeronautica 1 Riverside 27-Dec-79 400 Whitewater Cyn

aeronautica l Riverside 25-Mar-82 500 Hemet, Motte Reserve

laguna 1 San Diego 13-Mar-81 1075 San Felipe

microps 1 Riverside 25-Mar-82 500 Hemet, Motte Reserve

Spirembolus demonologicus Za San Diego 13-Mar-81 1075 San Felipe

erratus 1 Riverside 13-Jan-77 1100 Bautista Cyn, 27.4 km SE of Valle Vista

erratus 1 Riverside 13-Feb-77 600 Harford Preserve, Gavilan Hills

erratus 9 Riverside 20-Mar-77 600 Gavilan Hills

erratus 10 14 1 Riverside 27-Dec-79 400 Whitewater Cyn

erratus 1 Riverside 18-Nov-83 200 Vail Lake

erratus 1 1 Riverside 18-Nov-83 200 Vail Lake

erratus 1 Riverside 18-Nov-83 200 Vail Lake “nest 2”

erratus 1 9 San Bernardino 26-Feb-77 j25 E of Mentone

erratus 1 San Bernardino 13-May-82 700 Baldy Mesa, Phelan & Transmission Line Rd

erratus 1 Santa Barbara 11-Aug-83 1075 Figueroa Mt Rd

hibernus 1 Riverside 24-Aug-83 950 El Carrizo Oaks

hibernus 1 San Diego 13-Mar-81 1075 San Felipe

redondo 1 San Bernardino 2-May-68 325 1.6 km E of Summit on Rt 138

Tapinocyba dietrichi 8 7 Los Angeles 11-Nov-80 1000 0.8 km N of Jct. Vasquez & Bouquet Cyns

dietrichi 6 6 Los Angeles 14-Aug-81 1000 Bouquet Cyn nr. SE arm Bouquet Reservoir

dietrichi 1 Los Angeles 20-Apr-82 1000 Bouquet Cyn Rd at E. end Bouquet Cyn Reservoir

dietrichi 6 Los Angeles 20-Apr-82 1000 3.2 km E toward Palmdale on Bouquet Cyn Rd

dietrichi 1 Los Angeles 20-Apr-82 1000 Bouquet Cyn nr. entrance Vasquez Rd

dietrichi 2. 24 Riverside 14-Jan-83 1825 James Reserve Lk Fulmor

dietrichi 1 1 Riverside 4-Jun-83 750 7 km SE of Sage on R3

dietrichi 1 Riverside 24-Aug-83 950 E] Carrizo Oaks

SNHCdIN LVaY GOOM JO VNOVA YACIdS ‘HOLLNAYd 3 WALLAA

Table 1. Continued.

Family/Genus

Amaurobiidae

Amaurobius

Zanomys

Species

dietrichi

sp. #1

latescens

californica

ochra

ultima

sp. #1

Male

Nee

—

BRAYNrF WORN ADA W

NWN RRR

County

San Bernardino

San Diego

Santa Barbara

Riverside

San Bernardino

Los Angeles

Riverside

San Bernardino

San Diego

San Bernardino

Santa Barbara

Riverside

Date

2-May-68

3-Jun-81

1-May-85

25-Jul-79

13-Mar-81

11-Aug-83

17-Jun-77

9-Dec-81

21-May-82

26-Feb-77

11-Nov-80

13-Jan-77

14-Jan-83

18-Nov-83

2-May-68

3-Jun-81

1-May-85

14-Apr-85

25-Jul-79

25-Apr-80

15-May-78

13-May-82

11-Aug-83

9-Dec-8 1

Elevation

(meters)

325

975

1100

1250

1075

725

1075

1300

1325

975

525

1000

1100

1825

200

Wa)

975

1100

1850

1250

1075

1900

700

1075

950

Locale

1.6 km E of Summit on Rt 138

18.6 km from Hwy 15 on Rt 138 nr Little Horse-

thief Ranch

Summit Vly under cottonwood

William Heise Co. Park, Pine Hills S of Julian

San Felipe

San Felipe @ S2

Figueroa Mt Rd, 1.6 km W Rngr Sta

Covington Flat

Joshua Tree Natl Pk, Hidden Valley

6.4 km N of Yucca Valley

E of Mentone

0.8 km N of Jct. Vasquez & Bouquet Cyns

Bautista Cyn, 27.4 km SE of Valle Vista

James Reserve Lk Fulmor

0.8 km E of James Reserve, San Jacinto Mts

Vail Lake “‘nest 2”

1.6 km E of Summit on Rt 138

18.6 km from Hwy 15 on Rt 138 nr Little Horse-

thief Ranch

Summit Valley

Wrightwood

William Heise Co. Park, Pine Hills S. Julian

San Felipe

Burns Cyn above Rimrock

Baldy Mesa, Phelan & Transmission Line Rd

Figueroa Mt Rd

Joshua Tree Natl Pk

9¢

LSIDO'IOWNOLNA DIFIOVd-NVd HHL

(L)8L TOA

Table 1. Continued.

Family/Genus

Genus #1

Anapidae

Gertschana pis

Oonopidae

Oonops

Orchestina

Scaphiella

Dictynidae

Dictyna

Tivyna

Yorima

Scytodidae

Scytodes

Species

shantzi

sp. #1

moaba

hespera

cholla

moaba

angelica

imm

undescr. sp.

Male

NO a

County

Riverside

Los Angeles

Riverside

San Bernardino

Riverside

Los Angeles

Riverside

Date

7-Dec-77

17-Nov-80

14-Aug-81

5-Nov-80

13-Feb-77

25-Dec-80

29-Nov-81

9-Dec-81

18-Nov-83

25-Dec-80

9-Dec-8 1

27-Dec-79

25-Dec-80

5-Nov-80

25-May-82

26-Feb-77

29-Apr-82

17-Nov-80

13-Nov-79

5-Nov-80

Elevation

(meters)

unknown

1000

400

600

325

450

950

200

325

950

400

325

400

550

525

329

1000

400

Locale

Boyd Desert Center, Coyote Creek

Bouquet Reservoir

Bouquet Cyn nr. SE arm Bouquet Reservoir

Whitewater Cyn

Gavilan Hills

Box Springs Mts nr UCR

3.2 km S of Valle Vista on Rt 74, under live oak

Joshua Tree Natl Pk

Vail Lake

Box Springs Mts nr UCR

Joshua Tree Nat] Pk

Whitewater Cyn

Box Springs Mts nr UCR

Whitewater Cyn

Twentynine Palms

E of Mentone

Clinton Keith Rd, W of Murrieta near entrance

to cyn

Bouquet Reservoir

Whitewater Cyn

SNHddIN LVa GOOM AO VNNVA YACIdS ‘AOLLNAYd FY MALLAA Z00Z

Table 1. Continued.

Elevation

Family/Genus Species Male Fem Imm County Date (meters) Locale

Caponiidae

Orthono ps imm 1 Imperial 25-Mar-80 100 Indian wash, 30 km SE of Glamis

imm 3 Los Angeles 17-Nov-80 1000 Bouquet Reservoir

imm 1 Riverside 25-Mar-82 500 Hemet, Motte Reserve

zebra | Riverside 13-Feb-77 600 Gavilan Hills

Corinnidae

Trachelas pacificus 2 Riverside 18-Nov-83 200 Vail Lake

pacificus 1 San Bernardino 28-Sep-83 425 Afton Cyn

pacificus 1 Santa Barbara 11-Aug-83 1075 Figueroa Mt Rd, 1.6 km W of Rngr Sta

Gnaphosidae

Her pyllus propinquus 1 Riverside 17-Feb-78 1100 Bautista Cyn

propinquus 1 San Diego 13-Mar-81 725 San Felipe @ S2

Micaria imm 1 Riverside 25-Dec-80 325 Box Springs Mts nr UCR

pasadena 1 San Bernardino 1-May-85 1100 Summit Vly under cottonwood

Hahniidae

Hahnia sanjuanensis 2 Riverside 24-Aug-83 950 E] Carrizo Oaks

sanjuanensis 1 1 San Diego 25-Apr-80 1075 San Felipe

Liocranidae

Phrurotim pus mateonus 1 1 Riverside 18-Nov-83 200 Vail Lake

Scotinella kastoni 1 San Diego 25-Apr-80 1075 San Felipe

Cybaeidae

Cybaeota nana 1 Riverside 15-Nov-81 1825 James Reserve Lk Fulmor

nana 1 Riverside 14-Jan-83 1800 0.8 km E of James Reserve, San Jacinto Mts

Mysmenidae

Trogloneta paradoxum ie Los Angeles 14-Aug-81 1000 Bouquet Cyn nr. SE arm Bouquet Reservoir

LSIDO'TOWNOLNY OIIOVd-NVd HHL

(L)8L ‘TOA

Table 1. Continued.

Elevation

Family/Genus Species County Date (meters) Locale

Plectreuridae

Plectreurys deserta Riverside 9-Dec-81 950 Joshua Tree Natl Pk

imm Riverside 22-May-80 275 Corn Springs, SW Desert Ctr

Salticidae

Neon pixil Riverside 20-Mar-77 600 Gavilan Hills

Sitticus dorsatus Riverside 25-Dec-80 325 Box Springs Mts nr UCR

Theridiidae

Euryo pis spinigera Riverside 9-Dec-8 1 1350 Joshua Tree Natl Pk, Jumbo Rocks Cmpgrd

spinigera Riverside 9-Dec-81 950 Joshua Tree Natl Pk

Cyrtaucheniidae

Aptostichus imm Riverside 10-Jan-81 325 Riverside

Filistatidae

Kukulcania utahana Riverside 10-Jan-81 325 Riverside

Lycosidae

Pardosa california San Bernardino 5-Jun-79 950 4.8 km S of Hesperia

Pholcidae

Psilochorus acanthus Los Angeles 14-Aug-81 1000 Bouquet Cyn nr. SE arm Bouquet Reservoir

TOTALS

SNHddIW LVa GOOM AO VNNVSA YACIdS ‘AOLLNAYd F MALLAA ZOOT

30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

RESULTS AND DISCUSSION

A total of 316 spiders representing 20 families, 34 genera and 42 species were

identified in this study of Neotoma midden litter (Table 1). The linyphiid family

provided the greatest number of genera (6), species (12) and overall specimens

(204), garnering 65.2% of the specimen total. Tapinocyba dietrichi Crosby &

Bishop (Linyphiidae) was the most frequently collected species, contributing 118

specimens (37.3% of the specimen total). 7. dietrichi was found in midden debris

in five California counties (Los Angeles, Riverside, San Bernardino, San Diego,

Santa Barbara), ranging in approximate elevation from 325 to 1825 m but pre-

dominantly found at higher elevations. From the 14 middens in which it was

found, the average number of 7. dietrichi per midden was 8.4, with a range of

1—26 specimens. The other Tapinocyba species is believed to be undescribed. The

linyphiid genus Spirembolus contributed four species and the second most com-

monly collected species, S. erratus Millidge (16.5% of specimen total). The amau-

robiid genus, Zanomys, also contributed four species and the third most prevalent

species, Z. californica (Banks) (5.1% of specimen total) with one new species to

be described as a result of this study.

Although we expected that the linyphiids would comprise the bulk of midden

spider inhabitants, we were somewhat surprised to find Tapinocyba dietrichi in

southern California, let alone its overwhelming contribution to the study’s spec-

imen total. To our knowledge, the only other California specimens of this species

were taken in Alpine and Alameda Counties in central California (Crosby &

Bishop 1933, Boe, unpublished data). This species was not discovered during

faunal studies of coastal sage scrub in either San Diego or Riverside Counties

(Prentice et al. 1998, 2001) nor has it been found in montane oak (Quercus spp.)

leaf duff in southern California (Vetter, unpublished data).

Spirembolus erratus was previously known only from samples collected in

sycamore litter (Millidge 1980), grass litter in coastal sage scrub and in oak leaf

litter (Riverside County) (Prentice & Vetter, unpublished data). In this study, S.

erratus was collected most often at elevations of 200 to 700 m with two specimens

taken near 1100 m. Millidge (1980) states that virtually nothing is known of the

natural history of Spirembolus species in general, including our other listed Spir-

embolus species, S. redondo (Chamberlin & Ivie), S. hibernus Millidge, and S.

demonologicus (Crosby).

Zanomys californica has previously been collected from Neotoma middens by

J. Linsdale at the Hastings Reserve in central California (specimens at California

Academy of Sciences, examined). In our study, Z. californica was more prevalent

at high elevations (predominantly 1000—1850 m). Leech (1972) records both Z.

californica and Z. ochra Leech (holotype) from dry leaf duff, the latter also taken

from a rat midden in Juab County, Utah. Z. californica is a common inhabitant

of montane oak leaf duff (moist or dry) in southern California (Vetter, unpublished

data).

In addition, the concomitant collection of several females of “‘Ceratinopsis”’

palomara Chamberlin along with the male (which is undescribed) in rat nest litter

in the San Jacinto Mountains in corroboration with similar recent contempora-

neous collections of both sexes in oak leaf duff in the same mountain range (Vetter

2002. VETTER & PRENTICE: SPIDER FAUNA OF WOOD RAT MIDDENS 31

& Prentice, unpublished data) lead to the conclusion that this species does not

belong in the genus.

Members of many of the spider families identified from our southern Califor-

nian pack rat midden study are commonly found within (and possibly restricted

to) the leaf litter strata. In several ecosystems, especially those within the desert,

Neotoma middens represent a drastic vegetative change from the immediate sur-

roundings and may indeed be “‘oases”’ of increased survival potential. Occupied

middens contain fresh, nocturnally-harvested vegetation and copious amounts of

fecal pellets (Vestal 1938, Linsdale & Tevis 1951), both of which may attract and

support potential spider prey in the middens. Active and abandoned middens alike

provide refugia (for the local inhabitants) that are structurally more stable and

offer protection from environmental extremes than many of the other niches with-

in the surrounding environment. Vorhies & Taylor (1945) showed that over a

year’s time, temperatures inside an Arizona midden were consistently 11 to 17° C

lower than soil surface temperatures.

Platnick (1995) states that rat middens may provide a “‘vestige refuge’’ for the

araneophagous Orthonops spp. due to habitat destruction in various regions of

southern California. However, in an undisturbed site in the Colorado Desert south

of Joshua Tree National Park (400 m elevation), the caponiids Orthonops iceno-

glei Platnick and Tarsonops sp. were frequently collected from pitfall traps but

not from the remains of a Neotoma midden at the site (Vetter, unpublished data).

It may be that, in undisturbed areas, caponiids are not restricted to rat middens

nor to leaf duff.

Several of the species listed here were previously taken from Neotoma middens

during various studies. Gertsch (1960) discovered both Gertschanapis shantzi

(Gertsch) and Trogloneta paradoxum Gertsch while Platnick & Forster (1990)

recorded only G. shantzi during their examination of Linsdale’s wood rat material

from the Hastings Reserve. Ryckman & Lee (1956) sampled Neotoma middens

for reduviid bugs (Triatoma spp.), primarily in Riverside and San Bernardino

Counties and recorded the arachnids found therein. However, for the spiders, they

failed to provide frequency data which would have allowed percent species com-

position comparisons with our study. Species listed in both their study and ours

include the following: Zanomys californica, Trachelas pacificus Chamberlin &

Ivie, Dictyna cholla Gertsch & Davis, Herpyllus propinquus (Keyserling), Liny-

phantes laguna Chamberlin & Ivie, Pardosa californica Keyserling, and Orches-

tina moaba Chamberlin & Ivie. Assuming that their determinations were based

on mature specimens, the average size of the species sampled was larger than

that of the species that we examined which is probably due to sampling differ-

ences of whole nests by Ryckman and Lee in comparison to litter under the nests

by Cooper.

Vestal (1938) observed general aspects of Neotoma and their nests in Berkeley,

California and although most of his article focused on Neotoma behavior, he did

mention some arthropod associates. The only arachnids listed are mites (Histios-

toma sp.) and a pseudoscorpion (Apocheiridium fumeroides). Walters and Roth

(1950) sampled 30 nests in Oregon, recording a myriad of arthropod inhabitants,

but list only two genera and four families of spiders, Orchestina (Oonopidae),

Calymmaria (Hahniidae), Linyphiidae, and Lycosidae. Linsdale and Tevis (1951)

allocated 80 pages of their comprehensive book to animal associates of Neotoma

32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

nests at the Hastings Reserve in central California. However, they merely state

that “‘spiders occur everywhere within the house except in the used nest’

(“‘house”’ meaning the structure and “‘used’’ meaning the occupied portion of the

structure). Vorhies & Taylor (1940) investigated middens of N. albigula albigula

in Arizona and found that, within 100 nests, opilionids were the most common

arachnid encountered (74% of the nests), followed by miscellaneous spiders

(46%), black widow spiders (12%), scorpions (6%) and one tarantula (1%).

In our work here, we have documented the partial diversity of the Araneae in

Neotoma middens. Members of many of the families that were sampled here are

infrequently collected, possibly because of their rarity, secretive habits, and/or

very small size. Three of the species on our list are believed to be previously

undescribed taxa, one belonging to an undescribed genus in the family Amauro-

biidae (D. Ubick, personal communication). To our knowledge, two of the listed

linyphiid species, Ceratinops inflatus (Emerton) and Ceratinopsis interventa

Chamberlin, constitute new records for the state of California. Collecting material

from underneath wood rat middens should provide a rewarding experience for the

interested arachnologist by producing spiders that are rarely seen or that possess

bizarre somatic features (such as Trogloneta and Gertschanapis) and, thus, would

be a fruitful challenge.

ACKNOWLEDGMENT

We thank K. Cooper for the tremendous effort spent in collecting the Neotoma

nests, and EK Andrews and S. Frommer for providing information regarding the

history of the spiders used in this collection. This project was funded in part by

Humbug Mountain Engineering Services P-62 (RSV).

LITERATURE CITED

Clark, W. H. & J. T. Sankey. 1999. Late Holocene Sonoran desert arthropod remains from a packrat

midden, Catavifia, Baja California Norté, Mexico. Pan-Pac. Ent., 75: 183-199.

Crosby, C. R. & S. C. Bishop. 1933. American spiders: erigoneae, males with cephalic pits. Ann.

Entomol. Soc. Am., 26: 105-182.

Elias, S. A., J. I. Mead & L. D. Agenbroad. 1992. Late Quaternary arthropods from the Colorado

plateau, Arizona and Utah. Great Basin Natur., 52: 59-67.

English, P. EK 1923. The dusky-footed wood rat (Neotoma fuscipes). J. Mammal., 4: 1-9.

Gertsch, W. J. 1960. Descriptions of American spiders of the family Symphytognathidae. Amer. Mus.

Novit., 1981: 1-40.

Hall, W. E., C. A. Olson & T. R. Van Devender. 1989. Late Quaternary and Modern arthropods from

the Ajo Mountains of Southwestern Arizona. Pan-Pac. Entomol., 65: 322-347.

Leech, R. 1972. A revision of the nearctic Amaurobiidae (Arachnida: Araneida). Mem. Entomol. Soc.

Can., 84: 1-182.

Linsdale, J. M. & L. P. Tevis, Jr. 1951. The dusky-footed wood rat. Univ. Calif. Press, Berkeley,

California, 664 pp.

Millidge, A. EK 1980. The erigonine spiders of North America. Part 2. The genus Spirembolus Cham-

berlin (Araneae: Linyphiidae). J. Arachnol., 8: 109-158.

Platnick, N. I. 1995. A revision of the spider genus Orthonops (Araneae, Caponiidae). Amer. Mus.

Novit., 3150: 1-18.

Platnick, N. I. & R. R. Forster. 1990. On the spider family Anapidae (Araneae, Araneoidea) in the

United States. J. New York Entomol. Soc., 98: 108-112.

Prentice, T. R., J. C. Burger, W. R. Icenogle & R. A. Redak. 1998. Spiders from Diegan coastal sage

scrub (Arachnida: Araneae). Pan-Pac. Entomol., 74: 181-202.

Prentice, T. R., J. C. Burger, W. R. Icenogle & R. A. Redak. 2001. Spiders from Riversidian coastal

2002 VETTER & PRENTICE: SPIDER FAUNA OF WOOD RAT MIDDENS 33

sage scrub with comparisons to Diegan scrub fauna (Arachnida: Araneae). Pan-Pac. Entomol.,

77: 90-122.

Roth, V. D. 1993. The Spider Genera of North America (3rd ed.). Amer. Arachnological Society,

Gainesville, Florida, 203 pp.

Ryckman, R. E. & R. D. Lee. 1956. Spiders and phalangids associated with mammals (Citellus and

Neotoma) in southwestern United States and northern Mexico. Ann. Entomol. Soc. Amer., 49:

406-409.

Vestal, E. H. 1938. Biotic relations of the wood rat (Neotoma fuscipes) in the Berkeley Hills. J.

Mammal., 19: 1—36.

Vorhies, C. T. & W. P. Taylor. 1940. Life history and ecology of the white-throated wood rat, Neotoma

albigula albigula Hartley, in relation to grazing in Arizona. Univ. Arizona Agric. Exp. Sta.

Tech. Bull., 86: 455-529.

Vorhies, C. T. & W. P. Taylor. 1945. Water requirements of desert animals in the Southwest. Univ.

Arizona Agric. Exp. Sta. Tech. Bull., 107: 487-525.

Walters, R. D & V. D. Roth. 1950. Faunal nest study of the woodrat, Neotoma fuscipes monochroura

Rhoads. J. Mammal., 31: 290-292.

Received 30 June 2000; Accepted 4 September 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 34-42, (2002)

REDESCRIPTION OF GONATOCERUS ATRICLAVUS

GIRAULT (HYMENOPTERA: MYMARIDAE), WITH

NOTES ON OTHER EGG PARASITOIDS OF

SHARPSHOOTERS (HOMOPTERA: CICADELLIDAE:

PROCONIIND IN NORTHEASTERN MEXICO

SERGUEI V. TRIAPITSYN!, LARRY G. BEZARK’, & DAVID J. W. MORGAN?

‘Department of Entomology, University of California,

Riverside, California 92521

*Integrated Pest Control Branch, California Department of Food and

Agriculture, Sacramento, California 95814

Abstract—The mymarid wasp Gonatocerus atriclavus Girault, NEW STATUS, described orig-

inally from Trinidad as a variety of G. triguttatus Girault, is redescribed and illustrated based

on specimens reared from an egg mass of the sharpshooter Oncometopia clarior (Walker), col-

lected in Ciudad Victoria, Tamaulipas, Mexico. This is the first known host record for this

parasitoid species. Proconiine leafhopper host associations are also reported for three other eco-

nomically important Gonatocerus species from the ater species group in North America: G.

ashmeadi Girault, G. morrilli (Howard), and G. triguttatus Girault.

Key Words.—Insecta, Cicadellidae, Proconiini, Mymaridae, Gonatocerus, egg parasitoid, Mex-

ico.

The glassy-winged sharpshooter, Homalodisca coagulata (Say), is native to the

southeastern United States and northeastern Mexico. This species recently invaded

California and has become established in the southern part of the State. In 2000,

several sharpshooter populations were detected north of Kern County. In order to

control populations of this xylem-feeding species, an integrated program employ-

ing several control options is needed. One cornerstone of an integrated pest man-

agement program will be classical biological control.

Egg parasitoids of H. coagulata were discovered through survey activities con-

ducted in California and the southeastern United States in 1996 and 1997 (Triap-

itsyn et al. 1998). Levels of parasitism from one dominant species, Gonatocerus

ashmeadi Girault (Hymenoptera: Mymaridae), have been high in southern Cali-

fornia, but only on the summer and fall generations of the host. The level of

parasitism by other local parasitoids, such as Gonatocerus morrilli (Howard) and

Ufens spp. (Hymenoptera: Trichogrammatidae), has been generally very low. It

is clear that parasitoids attacking the spring generation of H. coagulata need to

be imported, screened through quarantine, reared, registered and released through

the appropriate permit process, and monitored, in order to impact this pest. Several

field expeditions to locate natural enemies of H. coagulata and its close relatives

were undertaken, including two trips to the Mexican states of Nuevo Leon and

Tamaulipas in early March and April 2000. Preliminary results of these activities

were reported by Morgan et al. (2000); this paper provides taxonomic notes on

the parasitoid species discovered through our survey and also indicates their host

associations, most of which are new records.

Members of the tribe Proconiini (Homoptera: Cicadellidae: Cicadellinae), to

which Homalodisca Stal belongs, often are referred to as sharpshooters. Distri-

2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION 35

Figure 1. Gonatocerus atriclavus Girault. Antenna, female.

bution of the 55 genera and numerous species that comprise this tribe is restricted

to the Western Hemisphere (Young 1968); only a few of those occur in the United

States, most of them are Neotropical.

In our survey, we searched for egg parasitoids of the species in two sharp-

shooter genera, Homalodisca and Oncometopia Stal, based on data obtained dur-

ing the previous year (Triapitsyn & Phillips 2000). All collections in Mexico were

made under a permit issued to our collaborator, E. Ruiz Cancino, Universidad

Auténoma de Tamaulipas at Ciudad Victoria. We searched for parasitized sharp-

shooter egg masses mainly in parks, citrus orchards, and irrigated private gardens.

Adult sharpshooters were collected directly in 70% ethanol for further identifi-

cation and association with the egg masses on their host plants. All parasitized

egg masses were sent to the University of California, Riverside (hereafter UCR),

quarantine facility under a USDA permit, where emerged parasitoids were

screened, identified, and propagated (Morgan et al. 2000).

Investigative responsibilities were divided between authors so that S.V.T. iden-

tified mymarid egg parasitoids and worked on taxonomic aspects of this study as

well as on conclusive remarks in the “‘Discussion’’, L.G.B. coordinated foreign

exploration efforts, L.G.B. and S.V.T. collected parasitized egg masses of Pro-

coniini in Mexico, and D.J.W.M. processed and reared the material in quarantine.

Terminology for morphological features used in the description is that of Huber

(1988); we use the abbreviation F1, F2, etc., to represent the first, second, etc.

funicular segments of the females, and the first, second, etc. flagellar segments of

the males. Measurements are given in micrometers (wm) as length or, if necessary,

as length/width. Abbreviations for depositories of specimens are as follows:

CSCA, California State Collection of Arthropods, Sacramento; UATM, Univer-

sidad Aut6dnoma de Tamaulipas, Ciudad Victoria, Mexico; UCRC, Entomology

Research Museum, University of California, Riverside; USNM, National Museum

of Natural History, Washington, D.C.

GONATOCERUS ATRICLAVUS_GIRAULT, 1917, NEW STATUS

(Figs. 1-4)

Gonatocerus triguttatus atriclavus Girault, 1917: 19 (as a new variety).

Gonatocerus triguttatus atriclavus Girault; Huber, 1988: 57.

Type Locality.—Mitan, Trinidad.

Types.—Lectotype 2 on point [USNM], here designated in order to maintain

stability of usage of the name, labeled: 1. ‘““Reared from egg-mass of leafhopper’’;

Vol. 781)

THE PAN-PACIFIC ENTOMOLOGIST

= =

Seo ~ “>?

=> = > ~

SS Raters

——~ SS N

—s Tey ae ae

>=S= Esse RS

—- SOR ~~. oe ay. hae.

ace) .™ Sn SS eS aN

— >= > eta RO Si. eg

= SNS tet ca

wa] SY SY 7A SS

a 28. FS i Core Dan EI? ROE Ae Pity, as

—= ~ Sah re ine PR +> =

= ere OS Say > ie Ee sce ~ he OS

— =~ ee ie ae. oy os St

== Sg Sey SY ete See ae SS! GS —~

= = ~

—s Sgeen ts Peete Ripe ae Beatie = VS SWAY

Se Fe ye oe gk eoeetea ai oN —

Og aes ble CS = — Le

Bae IS Sere SS a Se

ea A Ti Rear ne, DM a > Td ae ree

eee. ee ee oe shots

_ wm —~ = ~~ — =

eee eo ee ie er a ts aes > => ago OTS ee eae

Se eS eT RS LS ES =, Sea et

Sat ae — NS etre et “eee a as te ae

eee eS OSES ee ee

Seton Te geet | tea] ~sNe et wee

mh SS peta er eae a te es ee, Be Ll Leora ~

See ee, et ie RS ee | Bey

ou, =e, a a

_-_—_~ en ©, , ~~ ma

i 7 =

esi UE Soe a SS ee ae czy ee

(

Figure 2. Gonatocerus atriclavus Girault. Forewing, female.

2. ‘““Mitan BWI Jan. 1915”; 3. ““E W. Urich Collector’; 4. “53-41”; 5. ““Gon-

atocerus triguttatus atriclavus Gir 3 @ types.’’; 6. “Type No. 20098 U.S.N.M.’’.

Girault (1917) described this species as a variety of Gonatocerus triguttatus Gi-

rault without designating a holotype, from several “‘types’’, Catalogue No. 20098.

The type series now consists of the lectotype 2 mentioned above and 1 2 and 1

Gonatocerus atriclavus Girault. Dorsellum and propodeum, male.

Figure 3.

2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION Bh

Figure 4. Gonatocerus atriclavus Girault. Basal segments of antenna, male.

3 paralectotypes on points, here designated, labeled same as the lectotype except

label 1 is lacking and label 6 with “‘Paratype’’ instead of ““Type’”? [USNM].

Other Material Examined—MEXICO. TAMAULIPAS, Ciudad Victoria, Tamatan, ex. parasitized

egg mass of Oncometopia clarior (Walker) in hibiscus leaf collected 7 Mar 2000, L. G. Bezark and

S. V. Triapitsyn; 2 22 and 1 ¢ parasitoids emerged at UCR quarantine 20 Mar 2000 [UCRC]. 3 22

and 1 6 labeled: 1. ““MEXICO, interc.[epted] S. Antonio, Tex. VI-10-1959 Johnston’’; 2. “‘59-21950

3325 on palm leaves”’; 3. “‘“Gonatocerus triguttatus atriclavus Girault det. J. T. Huber 1984’? [USNM].

Original Description—The original description is inadequate as it is limited to a single sentence:

“Similar to typical form but the antenna concolorous except the club” (Girault 1917). Huber (1988)

was first to notice that G. atriclavus “‘is probably a good species, not a subspecies of triguttatus’’.

Redescri ption.—F emale.—Coloration: Head pale except upper face and ocellar triangle brown,

trabeculae and occiput dark brown, eyes and ocelli dusky. Antenna with scape yellow to light brown;

pedicel, F1—-F3, F4 (basally) and F7 brown; F4 (distally), F5 and F6 light brown; F8 dark brown; club

black. Neck pale, pronotum pale brown with darker spots; mesoscutum orange-brown anteriorly and

yellowish posteriorly, parapsidal furrows black; anterior scutellum light brown to brown; axilla brown

with darker spot at middle; posterior scutellum yellowish-orange-brown; dorsellum, propodeum, pro-,

meso-, and metapleura brown; lateral panels of metanotum and propodeal carinae dark brown. Legs

yellowish brown except all tarsi, middle and hind tibiae brown. Wings hyaline; venation brown to

dark brown. Petiole dark brown; gaster pale to light yellow with dark brown bands on terga; ovipositor

plates brown.

Head: Width 432. Antenna (Fig. 1) with radicle 0.2X as long as scape, scape very long and wide

for the species group, 3.0X as long as wide, very finely longitudinally striate (inner side) or almost

smooth (outer side), with several rows of strong setae; pedicel short, smooth; Fl short and without

sensilla; F2-F5 subequal in length but each slightly wider than preceding article; F6 subquadrate,

shorter than F5, F7 slightly wider than long, F8 much wider than long; F2 with 1 or 2 and F3—-F8

each with 2 longitudinal sensilla; all funicle segments densely setose; club long (length/width 2.5:1),

slightly wider than scape, and about as long as combined length of F1—F4, with 8 longitudinal sensilla,

its ventral surface covered with numerous minute, short setae and placoid sensilla, its dorsal surface

densely covered with longer setae.

Mesosoma: Pronotum divided medially, each lobe with 1 dorsal and 1 lateral strong seta. Mesoscu-

tum with a pair of strong adnotaular setae. Dorsellum rhomboidal (as in Fig. 3). Propodeal spiracle

kidney-shaped; lateral carinae well-developed, submedial carinae almost parallel except anteriorly;

propodeum (as in Fig. 3) otherwise smooth. Legs: foretibia with 6—7 conical sensilla. Forewing (Fig.

2) 3.55X as long as wide; marginal cilia very short, longest fringe seta about % maximum wing width.

Forewing blade bare immediately distal to submarginal vein, with 9 microtrichia behind marginal and

stigmal veins, cubital row of setae complete; remainder of blade densely setose. Submarginal vein

length 263, with 2 hypochaetae, marginal vein length 216, with 7 microchaetae between proximal and

distal macrochaetae, stigmal vein length 58. Hind wing blade bare except for complete rows of mi-

crotrichia along margins and several scattered discal setae at apex.

Metasoma: Petiole about as wide as long. Ovipositor about % length of gaster, barely exserted

beyond its apex. Outer plates of ovipositor each with 1 basal and 1 apical seta.

Measurements (n = 1).—Body: head: 67; mesosoma: 648; petiole: 72; gaster: 783; ovipositor: 603.

Antenna: radicle: 66; scape: 329; pedicel: 88; Fl: 47; F2: 80; F3: 84; F4: 80; F5: 80; F6: 65; F7: 51;

38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

F8: 43; club: 303. Forewing: 1503/423; longest fringe cilia: 88. Hind wing: 1107/69; venation: 513;

longest fringe cilia: 106. Legs (given as coxa, femur, tibia, tarsus): fore: 198, 306, 351, 369; middle:

126, 270, 450, 364; hind: 201, 405, 531, 441.

Male.—Coloration: Antenna brown to dark brown except base of scape yellow; face brown; ocellar

triangle and occiput dark brown, remainder of vertex and gena light brown; eyes and ocelli pinkish

brown. Neck light brown; pronotum, mesoscutum (except light brown edges of lateral lobes), axilla,

anterior and posterior scutellum, dorsellum, and propodeum shining brown; mesosomal pleura light

brown. All legs light brown except middle tibia and middle and hind tarsi slightly darker, hind tibia

brown. Petiole brown, gaster light brown with several dark cross-bands on terga. Otherwise similar

to female except for sexually dimorphic characters, as follows: antenna with scape and radicle fused

(Fig. 4); basal flagellomeres relatively wide, all flagellomeres with numerous longitudinal sensilla.

Measurements (n = 1).—Total body length (dry specimen): 1321; Antenna: scape + radicle: 153;

pedicel: 54; Fl: 160; F2: 182; F3: 171; F4: 186; F5: 193; F6: 186; F7: 187; F8: 172; F9: 176; F10:

173; Fll: 233. Forewing: 1884/532. Genitalia: 303.

Diagnosis.—This species is easily distinguished from all other described Ne-

arctic species of the ater species group (ater subgroup) of Gonatocerus that have

the forewing with the cubital row of microtrichia complete, extending to base of

marginal vein (Huber 1988) (1.e., G. ashmeadi Girault, G. fasciatus Girault, and

G. triguttatus Girault) by the transverse F8 in the female antenna. If using the

key by Huber (1988), G. atriclavus would be separated from G. fasciatus by

having the forewing hyaline, without fascia, thus keying to couplet 5 together

with G. ashmeadi and G. triguttatus. Besides the shape of F8 mentioned above,

G. atriclavus can also be easily separated from these two species by the dilated

scape, F5 and F6 lighter colored than other funicle segments, and a very long,

black club of the female antenna. In both G. ashmeadi and G. triguttatus, the

funicle of the females is uniformly dark brown to black.

Distribution.—Mexico and Trinidad. Although we collected this species in

Ciudad Victoria, Tamaulipas, which is near the border between the Nearctic and

Neotropical regions, it appears to be a mainly Neotropical species.

Hosts.—The type series was reared from a “‘leaf-hopper egg mass”’ in Trinidad

(Girault 1917). Our three specimens from Tamaulipas were reared from a sharp-

shooter egg mass on a hibiscus plant which was almost certainly laid by Oncom-

etopia clarior (Walker).

Upon emergence in quarantine, all the wasps were fed with honey, and the

single male was given time to mate with the females. The females were then

exposed to freshly laid egg masses of H. coagulata in citrus leaves several times;

despite the fact that they attempted to parasitize those eggs, we failed to obtain

any progeny. Eventually, wasps were killed in 70% ethyl alcohol to serve as

voucher specimens.

NOTES ON OTHER EGG PARASITOIDS OF PROCONIINE LEAFHOPPERS IN

NORTHEASTERN MEXICO

Gonatocerus ashmeadi Girault. This common North American species was

redescribed and illustrated in detail by Huber (1988), who also indicated its known

hosts: Cuerna costalis (FE), Homalodisca coagulata, H. lacerta (Fowler), and On-

cometopia orbona (F). Oncometopia clarior is added here to that list.

Individuals from northeastern Mexico appeared identical to specimens collected

in southern California, except for two rearings of darker-colored individuals from

Tamaulipas that were not propagated. We attribute such differences in body col-

oration to intraspecific variability, which is possibly host-induced.

2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION 39

Under quarantine laboratory conditions (22—25° C, 40-50% RH), the Ciudad

Victoria population of G. ashmeadi successfully parasitized H. coagulata eggs

laid in a variety of plants including chrysanthemum (Dendranthema sp.), crape

myrtle (Lagerstroemia indica L.), grape (Vitis vinifera L.), sweet orange (Citrus

sinensis (L.) Osbeck), toyon (Heteromeles arbutifolia (Aiton) M. Romer), and

Verbascum sp. Eggs laid by H. lacerta in chrysanthemum, crape myrtle, and sweet

orange were also accepted for parasitism. Viable offspring emerged from all ma-

terial tested. Further studies were then conducted on the biology of G. ashmeadi

as well as of the two other Gonatocerus species discussed below; the results will

be reported elsewhere.

Material Examined.—Gonatocerus ashmeadi: MEXICO: NUEVO LEON, Monterrey, UANL cam-

pus, ex. parasitized host egg masses coll. 6 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 18 2°, 7

3d em. 21 Mar 2000 at UCR quarantine (ex. Oncometopia clarior (Walker) on leaves of Fraxinus

sp.). TAMAULIPAS: Ciudad Victoria, 14 Feb 2000, S. N. Myartseva, 16 22, 2 d¢ (ex. sharpshooter

eggs on citrus leaf); Ciudad Victoria, Tamatan, ex. parasitized host egg masses coll. 7 Mar 2000, L.

G. Bezark and S. V. Triapitsyn, 19 22, 7 36 parasitoids em. in UCR quarantine 14-23 Mar 2000

(ex. Oncometopia clarior (Walker) on hibiscus leaves); Llera de Canales, ex. parasitized sharpshooter

egg mass (on hibiscus leaf) coll. 8 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 1 2, 1 3 parasitoids

em. in UCR quarantine 23 Mar 2000; Municipio Hidalgo, nr. Ejido Benito Juarez, garden in Hotel

Hacienda Santa Engracia, ex. parasitized sharpshooter egg mass (on orange leaf) coll. 10 Apr 2000,

L. G. Bezark and S. V. Triapitsyn, 7 92, 3 dd em. 14 Apr 2000 at UCR quarantine; 5 km N of

Valle Hermoso, ex. parasitized host egg masses coll. 13 Apr 2000, L. G. Bezark, 10 22,11 dd em.

20 Apr 2000 at UCR quarantine (ex. Homalodisca coagulata (Say) on hibiscus leaves) [UATM,

UCRC].

Oncometopia clarior. MEXICO. NUEVO LEON: 6 km SE of Allende, Sanatorio Naturista de

Canoas, 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn (on citrus); Monterrey, UANL campus, 6

Mar 2000, L. G. Bezark and S. V. Triapitsyn (on crape myrtle). TAMAULIPAS, Ciudad Victoria,

Tamatan, 7 Mar 2000, L. G. Bezark and S. V. Triapitsyn (on hibiscus) [CSCA].

Gonatocerus morrilli (Howard). This species is common in southern U.S.A.

and Mexico according to Huber (1988). The only previously known host of G.

morrilli, H. coagulata, was first indicated by Turner & Pollard (1959) and later

by Triapitsyn et al. (1998). We observed a female of this species ovipositing in

a sharpshooter egg mass, probably that of O. sp. nr. nigricans (Walker), on a

hibiscus plant in the garden of Hotel Rancho Mariposa, near Santander Jiménez

in Tamaulipas. Another leafhopper host of G. morrilli that we discovered in Ta-

maulipas is O. clarior.

A colony of G. morrilli, individuals of which were morphologically similar to

those collected in southern California, was established at UCR quarantine using

wasps originating from Ciudad Victoria, Tamaulipas. They were successfully

reared on both H. coagulata and H. lacerta eggs laid in host plants as described

previously for G. ashmeadi.

Material Examined —MEXICO. TAMAULIPAS: Ciudad Victoria, Tamatan, ex. parasitized host

egg mass coll. 7 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 9 22, 1 ¢ parasitoids em. in UCR

quarantine 20—21 Mar 2000 (ex. Oncometopia clarior (Walker) on hibiscus leaf); Llera de Canales,

ex. parasitized sharpshooter egg masses (on citrus and hibiscus leaves) coll. 8 Mar 2000, L. G. Bezark

and S. V. Triapitsyn, 14 2 2, 2 d6 parasitoids em. in UCR quarantine 14—23 Mar 2000; same location

and collectors, ex. parasitized sharpshooter egg mass (on orange leaf) coll. 11 Apr 2000, 2 22,1 46

parasitoids em. in UCR quarantine 17 Apr 2000; nr. Santander Jiménez, Hotel Rancho Mariposa, ex.

parasitized sharpshooter egg masses (on hibiscus leaves) coll. 9 Mar 2000, L. G. Bezark and S. V.

Triapitsyn, 7 22,3 d¢ parasitoids em. in UCR quarantine 15—20 Mar 2000; same location, 22 Apr

40 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

1999, S. Triapitsyn, 2 2 2 (on hibiscus, by sweeping); 5 km N of Valle Hermoso, 10 Mar 2000, S.

Triapitsyn, 1 d (on hibiscus, by sweeping) [UATM, UCRC].

Gonatocerus triguttatus Girault. This species was redescribed and illustrated

by Huber (1988) based on the type series from Trinidad and additional specimens

from Texas. Adult specimens of G. triguttatus were reared in northern Tamaulipas

during April 1999 from the egg masses of its only known host at that time, H.

coagulata, laid in citrus and peach leaves (Triapitsyn & Phillips 2000). In early

spring of 2000, we dissected dead specimens of G. triguttatus from an egg mass

of a sharpshooter on a citrus (orange) leaf in G6mez Farias, whichis in the tropical

part of Tamaulipas. These specimens apparently could not emerge from the egg

mass and had died not long before we found them. From the same orange tree,

we collected an adult male leafhopper, later identified by Raymond J. Gill as

Paraulacizes thunbergi (Stal); that species thus can be considered as a probable

host for G. triguttatus. Paraulacizes thunbergi, previously known from southern

Mexico (Young 1968), is a new addition to the list of proconiine leafhoppers in

Tamaulipas that feed and oviposit on citrus plants (Coronado-Blanco et al. 2000).

Additional adult specimens of P. thunbergi were collected in the Nearctic part of

Mexico in Nuevo Le6n and Tamaulipas. Other apparent hosts of G. triguttatus

are O. clarior and O. sp. (see “‘Material examined” below).

A colony of G. triguttatus was established at UCR quarantine (Morgan et al.

2000) from adults that emerged from the egg masses of H. coagulata in hibiscus

near Valle Hermoso, Tamaulipas. This colony was being reared on H. coagulata

eggs laid in chrysanthemum and orange leaves. All plants tested with G. ashmeadi

were also accepted by G. triguttatus. When G. triguttatus females were offered

eggs of the smoke-tree sharpshooter, H. lacerta, which is native to California,

they readily parasitized them and produced viable offspring.

Material Examined—Gonatocerus triguttatus: MEXICO. NUEVO LEON, 6 km SE of Allende,

Sanatorio Naturista de Canoas, ex. parasitized host egg mass coll. 10 Apr 2000, L. G. Bezark and S.

V. Triapitsyn, 1 2 em. 14 Apr 2000 in UCR quarantine (ex. Oncometopia clarior (Walker) on orange

leaf). TAMAULIPAS: Gomez Farias, 8 Mar 2000, S. N. Myartseva, L. G. Bezark and S. V. Triapitsyn,

1 2 (dissected from a sharpshooter egg mass on orange leaf); nr. Santander Jiménez, Hotel Rancho

Mariposa, ex. parasitized host egg mass coll. 9 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 2 2 2

parasitoids em. in UCR quarantine 15 Mar 2000 (ex. Oncometopia sp. on hibiscus leaf); 5 km N of

Valle Hermoso, 10 Mar 2000, S. Triapitsyn, 1 ¢ (on hibiscus, by sweeping); same location, ex.

parasitized host egg masses coll. 13 Apr 2000, L. G. Bezark, 14 22, 15 do em. 20-27 Apr 2000 at

UCR quarantine (ex. Homalodisca coagulata (Say) on hibiscus leaves) [UATM, UCRC].

Paraulacizes thunber gi—MEXICO. NUEVO LEON, 6 km SE of Allende, Sanatorio Naturista de

Canoas, 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn (on Malva sp.). TAMAULIPAS: Gémez

Farias, 8 Mar 2000, S. N. Myartseva, L. G. Bezark and S. V. Triapitsyn, 1 ¢ (on orange); Municipio

Hidalgo, nr. Ejido Benito Juarez, garden in Hotel Hacienda Santa Engracia, 7 Mar 2000, S. N. Myart-

seva, L. G. Bezark and S. V. Triapitsyn (on Malva sp.) [CSCA].

Ufens sp. This species first emerged from a single sharpshooter egg mass laid

in an orange leaf collected at Llera de Canales, Tamaulipas. It was later reared

in Nuevo Leon from the eggs of O. clarior. It is a gregarious species: 76 wasps

emerged from a clutch of 15 sharpshooter eggs (3—7 emergences per host egg).

Morphologically, Ufens sp. is very similar to, but probably distant from, one taken

from unspecified leafhopper eggs on elm and jojoba in southern California; H.

lacerta is known to be a common associate to the latter plant there. It is also

different from the other two Ufens species known from Homalodisca eggs in

2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION 41

southern California. All species involved are almost certainly undescribed (J. D.

Pinto, personal communication).

An attempt was made to initiate a colony of the Mexican Ufens sp. at UCR

quarantine. In the first generation, wasps were offered egg masses of H. coagulata

laid in leaves of chrysanthemum (14 egg clutches), toyon (2), grape (2), and

orange (6). Wasps emerged from only one clutch, laid in an orange leaf, after 16

days, a longer developmental period than was observed for any of the three Gon-

atocerus species evaluated (D.J.W.M., unpublished data). The second generation

was offered host eggs on Verbascum sp. (4), crape myrtle (6), orange (17), and

chrysanthemum (8). No wasps emerged from these leaves. We suggest that a

combination of conditions and strong host plant preferences are responsible for

the failure to propagate this trichogrammatid species successfully.

Material Examined —MEXICO. NUEVO LEON, 6 km SE of Allende, Sanatorio Naturista de Canoas,

ex. parasitized host egg mass coll. 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn, 37 22,6 6d em.

17 Apr 2000 at UCR quarantine (ex. Oncometopia clarior (Walker) on orange leaf). TAMAULIPAS:

Llera de Canales, ex. parasitized sharpshooter egg mass collected 8 Mar 2000, L. G. Bezark and S. V.

Triapitsyn, 62 2°, 14 dd parasitoids em. in UCR quarantine 14-15 Mar 2000 [UATM, UCRC].

DISCUSSION

Considering the great diversity of the proconiine leafhoppers, which are among

the largest leafhoppers known, and whose eggs are laid in clusters in plant tissue

(Young 1968), it is surprising how little is generally known about their biology

and natural enemies beyond a few economically important species in the United

States. The work by Turner & Pollard (1959) had been practically the only one

available on this subject until recently, when the establishment of H. coagulata

in California prompted the interest in sharpshooter investigations, including stud-

ies of their egg parasitoids (Triapitsyn et al. 1998).

Most of the reported egg parasitoids of Cuerna, Homalodisca and Oncometo pia

species are members of Gonatocerus. All known North American species parasitic

on these leafhopper genera belong to the ater species group of Gonatocerus as

defined by Huber (1988), and we believe that this might be the case for all

proconiines in the New World. The amazing diversity of ater group species of

Gonatocerus in Malaise trap samples from Central and South America correlates

very well with the even greater diversity of the sharpshooters from these areas.

Further research is needed to demonstrate the validity of this apparent correlation.

It is unlikely that species of the ater group of Gonatocerus in the New World

are species-specific to their leafhopper hosts; rather, they may be narrowly to

broadly oligophagous, 1.e., able to parasitize species of a number of closely-related

host genera within the tribe Proconiini. Some species of parasitoids, however,

may display a preference for certain sharpshooter species. Most likely, however,

that they discriminate their hosts based on the size of hosts’ eggs and the habitat.

Some sharpshooter species, like H. coagulata, are able to feed upon many plant

species, but prefer certain ones for oviposition. To be successful in finding host

egg masses, female parasitoids must concentrate their searching activity on those

plants. As a result, different parasitoid species may have become more host plant

specific than insect host specific, like for instance the common North American

mymarid Anagrus nigriventris Girault (Al-Wahaibi & Walker 2000). Thus, for a

classical biological control program to be successful in California against H. coa-

42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

gulata, whose females oviposit on a great number of different plants, introduction

of many different species, as well as various biotypes from any given species,

may be warranted.

ACKNOWLEDGMENT

We thank Enrique Ruiz Cancino, Svetlana N. Myartseva and Vladimir A. Trja-

pitzin (Universidad Aut6énoma de Tamaulipas, Ciudad Victoria, Mexico) for fa-

cilitating our collecting efforts in Mexico; Walker A. Jones (USDA-ARS, Wes-

laco, Texas) for assistance in collecting and logistical support, Raymond J. Gill

(California Department of Food and Agriculture, Sacramento) for the sharpshooter

identifications, John D. Pinto (University of California, Riverside) for identifica-

tion of Ufens species, and John T. Huber (Canadian Forestry Service, Natural

Resources Canada, Ottawa) for valuable advice on the identity of Gonatocerus

species. Vladimir Berezovskiy (University of California, Riverside) point- and

slide-mounted the parasitoids and made line drawings. This study was sponsored

in part by a grant to Mark S. Hoddle and Serguei V. Triapitsyn (University of

California, Riverside) who gratefully acknowledge the California Department of

Food and Agriculture for financial support.

LITERATURE CITED

Al-Wahaibi, A. K. & G. P. Walker. 2000. Searching and oviposition behavior of a mymarid egg

parasitoid, Anagrus nigriventris, on five host plants species of its leafhopper host, Circulifer

tenellus. Entomol. Exp. Appl., 96: 9-25.

Coronado-Blanco, J. M., E. Ruiz-Cancino & S. V. Triapitsyn. 2000. Chicharritas de la tribu Proconiini

(Homoptera: Cicadellidae) asociadas a citricos en Tamaulipas, México. Acta Zool. Mex. (n. s.),

81: 133-134.

Girault, A. A. 1917. Descriptiones stellarum novarum. Privately printed, 22 pp.

Huber, J. T. 1988. The species groups of Gonatocerus Nees in North America with a revision of the

sulphuripes and ater groups (Hymenoptera: Mymaridae). Mem. Entomol. Soc. Canada, 141:

1-109.

Morgan, D. J. W., S. V. Triapitsyn, R. A. Redak, L. G. Bezark & M. S. Hoddle. 2000. Biological

control of the glassy-winged sharpshooter: current status and future potential. pp. 167-171. In

Hoddle, M. S. (ed.). [Proceedings] California Conference on Biological Control, held July 11-

12, 2000 at the historic Mission Inn, Riverside, California, 205 pp.

Triapitsyn, S. V., R. EK Mizell, Il, J. L. Bossart & C. E. Carlton. 1998. Egg parasitoids of Homalodisca

coagulata (Homoptera: Cicadellidae). Florida Entomol., 81 (2): 241-243.

Triapitsyn, S. V. & P. A. Phillips. 2000. First host record of Gonatocerus triguttatus (Hymenoptera:

Mymaridae) from eggs of Homalodisca coagulata (Homoptera: Cicadellidae), with notes on

the distribution of the host. Florida Entomol., 83 (2): 200-203.

Turner, W. E & H. N. Pollard. 1959. Life histories and behavior of five insect vectors of phony peach

disease. Tech. Bull. U.S. Dep. Agric. 1188, 28 pp.

Young, D. A. 1968. Taxonomic study of the Cicadellinae (Homoptera, Cicadellidae). Part 1. Proconiini.

United States Nat. Mus. Tech. Bull. 261, 287 pp.

Received § December 2000; Accepted 13 September 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 43-55, (2002)

COPULATION DURATION IN THREE SPECIES OF

ANTHOCORIS (HETEROPTERA: ANTHOCORIDAE) AT

DIFFERENT TEMPERATURES AND EFFECTS ON

INSEMINATION AND OVARIAN DEVELOPMENT

Davip R. HoRTON, TAMERA M. LEwIs, & TONYA HINOJOSA

USDA-ARS, 5230 Konnowac Pass Road, Wapato, Washington 98951

Abstract—We compared duration of copulation among three species of predatory bugs: Antho-

coris tomentosus Péricart, A. whitei Reuter, and A. nemoralis (Fabricius). Copulation duration at

room temperatures (22—24° C) was longest in A. whitei (x = 89.3 min), of intermediate length

in A. tomentosus (x = 40.0 min), and shortest in A. nemoralis (x = 12.7 min). By interrupting

mating pairs we showed that long duration copulations were more likely than short copulations

to result in insemination and in ovarian maturation in the female. Probability of ovarian devel-

opment following a shortened copulation was often lower than probability of insemination, sug-

gesting that insemination alone was not always enough to prompt ovarian development. Size of

the sperm reservoir in the female increased with increasing duration of copulation. Males of all

species transferred seminal products for most of the copulation period, suggesting that the longer

copulations noted for A. whitei and A. tomentosus (relative to duration in A. nemoralis) were

not due to post-insemination mate-guarding by A. whitei or A. tomentosus. Copulation duration

was longer at 15° C (range: 25-181 min) than at 25° C (range: 13-102 min). We interrupted

mating pairs at both temperatures in all species to determine how the combined factors of

temperature and copulation duration affect insemination rates and probability of oocyte matu-

ration. For a given copulation duration, probability of insemination and oocyte maturation were

higher at 25° C than at 15° C. Species’ differences in copulation duration may reflect differences

among species in how rapidly males transfer sperm, and that lower temperatures may make it

physically more difficult for the male to force seminal products through the aedeagus.

Key Words.—Insecta, Anthocoridae, mating behavior, sperm transfer, copulation.

Predatory bugs in the genus Anthocoris are important sources of biological

control in natural and agricultural ecosystems (Lattin 1999). Despite their impor-

tance, however, these species are poorly studied in certain aspects of basic biology.

Our laboratory has recently begun to study the reproductive biology of several

North American species of Anthocoris, with goals ultimately to improve our un-

derstanding of mating behavior and reproductive biology in these insects.

In the present study, we explored how copulation duration varies among three

species of Anthocoris that inhabit the Pacific northwest region of the United

States: A. tomentosus Péricart, A. whitei Reuter, and A. nemoralis (Fabricius).

Copulation duration varies extensively among insect taxa (Thornhill & Alcock

1983), to the extent that even closely related species may exhibit large divergence

in this trait (Thornhill & Alcock 1983, Cordero 1990, Lachmann 1997). There

are potentially a variety of costs and benefits associated with short- or long-

duration couplings (Alcock 1994). Here, we address whether artificially shortened

copulations in these species affect probability of insemination, probability of ovar-

ian development, and amount of seminal material transferred to the female. The

first hypothesis tested here is that females that experience artificially shortened

copulations would be less likely than undisturbed females to be inseminated, as

shown in other insect species (Farias et al. 1972, Lew & Ball 1980). We also

determined whether these artificially shortened copulations resulted in decreased

44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

probability of ovarian development in the female. Maturation of ovaries in An-

thocoris requires mating (Anderson 1962, Shimizu 1967). However, it is not

known whether the mating act itself is sufficient to prompt ovarian development,

or whether insemination is required. If the mating act alone is enough to prompt

ovarian development, then we expect that severely shortened copulations, 1.e.,

those resulting in a lack of insemination, would still prompt ovarian development.

Our second major objective was to determine whether the longer copulations

of A. whitei and A. tomentosus (relative to copulation duration in A. nemoralis

[see below]), was due to mate-guarding by males of these two species. Many

insect species prolong copulation beyond that necessary to complete insemination,

as a male strategy to delay remating by the female and to ensure paternity (Alcock

1994). Females of A. tomentosus and A. whitei will mate multiple times under

laboratory conditions (unpublished data). Here, we interrupted copulating pairs at

several time intervals between intromission and the end of copulation, and esti-

mated quantity of seminal products that had been transferred by the male at each

duration. If males prolonged copulation beyond the time required to inseminate

the female (1.e., if the male engaged in mate-guarding behavior), we would expect

that the amount of seminal material transferred in artificially shortened copulations

would not differ from the amount transferred in uninterrupted matings.

Finally, we determined whether temperature mediates the interaction between

copulation duration and probability of insemination. In Anthocoris spp., sperm

are transferred by the male through a thin membranous tube in the female (cop-

ulatory tube; Carayon 1953) directly into her sperm pouch. Elsewhere, we sug-

gested that males in certain species of Anthocoris experience physical difficulties

in forcing seminal materials through the aedeagus and female copulatory tube

(Horton et al. 2001). Here, we hypothesized that these difficulties would be am-

plified with decreasing temperatures, resulting in longer copulations at a lower

temperature than a higher temperature. Moreover, we hypothesized that probabil-

ity of insemination would decrease with decreasing temperature for a copulation

of a given duration. This last hypothesis was tested by interrupting mating pairs

at specified time intervals for matings conducted at different temperatures.

MATERIALS AND METHODS

Source of Insects.—Laboratory cultures of the three species were begun from

field-collected insects. Nymphs and adults of A. tomentosus were collected from

Salix sp. growing west of Tieton, Washington. Anthocoris whitei was collected

from antelope bitterbrush, Purshia tridentata (Pursh), growing in rangeland west

of Tieton. Assays with A. tomentosus and A. whitei were done using offspring of

field-collected insects. The third species, A. nemoralis, is native to Europe but

was released in western North America to control pear psylla, Cacopsylla pyricola

(Foerster) (McMullen 1971). Nymphs and adults of A. nemoralis were collected

in Richmond, California from Acacia longifolia Willdenow, where it occurs in

association with a psyllid pest (Dreistadt and Hagen 1994), and from an uniden-

tified shrub species located in Golden Gate Park, San Francisco. Insects from the

two sites were combined into a single culture. Assays used a mix of first-gener-

ation (offspring of field-collected insects) and second-generation insects.

Insects were reared on pear seedlings infested with pear psylla at 22° C under

long-day conditions (16:8 h [L:D]). Offspring from the parental cultures were

2002 HORTON ET AL.: ANTHOCORIS COPULATION 45

collected as older nymphs and placed singly in glass petri dishes lined with filter

paper. Psylla-infested leaves were added daily to each dish. Petri dishes were

checked daily for eclosion of new adults. Date of eclosion and sex of each bug

were recorded. All assays used previously unmated insects | to 6 days of age.

Voucher specimens of each species have been deposited in the M.T. James

Entomology Museum at Washington State University, Department of Entomology,

Pullman.

Study 1. Copulation Duration, Insemination, and Oocyte Maturation.—Copu-

lation duration was quantified for the interval beginning with intromission and

ending with the male’s withdrawal of the aedeagus. Withdrawal from the female

was always followed immediately by physical separation of the two sexes in all

species. Matings were done at room temperature (22—24° C) under fluorescent

lighting. Plastic petri dishes (60 mm in diameter) were used as mating arenas.

Preliminary trials were conducted to determine copulation duration in pairs al-

lowed to mate without interference. Based upon these trials, we established a

range of copulation durations to be tested. Pairs were manually separated after

the appropriate time interval by gently prodding the insects with a small paint

brush. The test durations are (in min from intromission to manual interruption):

A. tomentosus (O [virgin females], 10, 20, 25, 30, 40, and uninterrupted); A. whitei

(O, 10, 20, 40, 60, 90, and uninterrupted); and A. nemoralis (O, 2, 4, 8, 12, and

uninterrupted).

Probability of sperm transfer and ovarian maturation were recorded as a func-

tion of copulation duration for the manipulated and uninterrupted pairs. Imme-

diately after copulation, females were either dissected to determine whether in-

semination had occurred, or set aside to allow oocyte development (see below).

In Anthocoris, sperm are transferred by the male directly into the female’s mem-

branous sperm pouch (Carayon 1953); there is no spermatophore. To obtain an

indication about the amount of seminal material transferred by the male at dif-

ferent copulation durations, we measured size of the sperm pouch in females

allowed to copulate for specified time intervals. After mating, the female was

killed by crushing her head and thorax. The abdomen was dissected away from

the rest of the body in a drop of saline by using two insect pins. The sperm pouch

was carefully removed from the other tissues and submerged in a drop of saline

on a microscope slide. The pouch was situated on the slide so that the entrance

of the copulatory tube was at the base of the pouch (Fig. 1). The pouch was then

measured immediately at 50X under a dissecting microscope equipped with an

ocular micrometer. Two measurements were recorded (Fig. 1): maximum length—

measured from the base of the membranous pouch to the top of the pouch; max-

imum width—measured perpendicular to the length measurement. Virgin females

were included as controls.

After measuring the pouch, we checked for the presence of sperm by placing

a cover slip over the pouch and pressing on the slip until the pouch ruptured.

Pouch contents were then examined for the presence of sperm at 100—400 under

a compound microscope.

Females that were not dissected were set aside to record whether ovaries ma-

tured. Following copulation in both interrupted and uninterrupted pairs, females

were separated from the male and put singly on pear seedlings infested with pear

psylla. Seedlings and insects were placed in environmental chambers at 22° C and

46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

/ *— Copulatory tube

Figure 1. Methods used for estimating maximum length and width of sperm pouch in virgin and

mated females of A. tomentosus, A. whitei, and A. nemoralis. Pouch shapes are highly variable both

within and between species; illustration provides more-or-less typical shape for A. tomentosus.

a 16:8 h photoperiod. Females were then dissected 8—12 (A. tomentosus) or 4—6

(A. nemoralis and A. whitei) days later; the preoviposition period (1.e., the number

of days between mating and onset of egglaying) in these species is ca. 8, 4—5,

and 3-4 days in A. tomentosus, A. whitei, and A. nemoralis, respectively (unpub-

lished data). Dissected females were scored as reproductive or as non-reproduc-

tive. To be scored as reproductive, the female had to have at least one ovariole

containing a mature egg. Appearance of the ovaries is quite different in repro-

ductive and non-reproductive females of these species. The mature terminal oo-

cyte in a reproductive female is approximately 1.5 (A. nemoralis), 2.0 (A. whitei),

and 4.0 (A. tomentosus) times as long as the terminal oocyte in an unmated female

of the same age (unpublished data), and, moreover, has the characteristic appear-

ance of the deposited egg.

Probability that the female contained at least one mature oocyte was modeled

as a logistic function of copulation duration (PROC CATMOD; SAS Institute

1987). We tested whether copulation duration affected probability of insemination

using x? tests. To compare frequency of females scored as reproductive to fre-

quency of females that were successfully inseminated, we conducted 2 X 2 con-

tingency tests. To determine whether size of the sperm pouch depended upon

2002 HORTON ET AL.: ANTHOCORIS COPULATION 47

copulation duration, we first conducted a principal components analysis (PROC

PRINCOMP; SAS Institute 1987) on the width and length measures to create a

single size variable. Analysis of variance was then used to compare mean principal

component scores among different copulation durations. Sample sizes are provid-

ed in the RESULTS.

Study 2. Effects of Temperature.—Two temperatures were compared for effects

on mating and insemination: 15° C and 25° C. Assays were conducted under

fluorescent lighting in controlled environmental rooms set to the appropriate tem-

perature. Petri dishes containing the adult insects were placed in the rooms 1 h

before the assay was conducted to allow the insects to acclimate. After 1 h, pairs

(1 male and 1 female) were moved to mating arenas (plastic petri dishes 60 mm

in diameter) and allowed to mate. Pairs that had not initiated copulation within

30 min of being placed in the arenas were discarded.

Copulation duration was manipulated at both temperatures by separating pairs

at two specified time intervals, producing for each species copulations of three

durations (in min): A. tomentosus—10, 30, and uninterrupted controls; A. whitei—

20, 40, and uninterrupted controls; A. nemoralis—2, 8, and uninterrupted controls.

The female from each pair was collected at the end of the mating bout. One-half

of the females were then dissected to determine whether insemination had oc-

curred. Females that were not dissected were set aside to monitor ovarian matu-

ration, using the methods described above. Sample sizes are provided in the RE-

SULTS.

Mean copulation duration in uninterrupted pairs was compared between tem-

peratures with a two-sample t-test (PROC TTEST; SAS Institute 1987). Contin-

gency tests were used to test whether probability of insemination or probability

of oocyte maturation in interrupted matings were affected by temperature and

copulation duration. For each species, a 2 X 2 X 2 (temperature [15° C versus

25° C] X duration [short interrupted versus long interrupted] < status [inseminated

versus not inseminated]) model was fitted to the insemination data and analyzed

using log-linear methods (PROC CATMOD; SAS Institute 1987). The analyses

were repeated replacing frequency of insemination with frequency of oocyte mat-

uration (.e., female containing at least one mature oocyte versus no mature oo-

cytes).

RESULTS

Study 1. Copulation Duration, Insemination, and Oocyte Maturation.—Mean

copulation duration in uninterrupted pairs differed among the three species (Fig.

2: solid circles having standard error bars). There was also substantial variation

within species, as indicated by the range of values noted in uninterrupted pairs:

A. tomentosus, range = 7—64 min, x = 40.0 min, n = 17; A. whitei, range = 39—

138 min, X = 89.3 min, n = 13; A. nemoralis, range = 7.7—20.9 min, x = 12.7

min, n = 20. Probability of oocyte maturation in females from interrupted pairs

increased with increasing duration of copulation (Fig. 2; solid circles and solid

lines). Probability of ovarian maturation was 82—100% in uninterrupted females

(solid circles with error bars in Fig. 2). In some interrupted females that were

scored as reproductive, oocyte maturation was limited to a subset of the ovarioles,

rather than in all ovarioles as seen in females from undisturbed matings.

Probability of insemination increased with increasing copulation duration in all

ore)

THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

O 10 20 30 40 50

Uninterrupted

A. white!

O 20 40 60 80 100 120

A. MemOrals O

(©) WNEdS YPN SAJEWS JO %

Uninterrupted

— Model

@ Observed

% of Females Having at Least One Mature Oocyte (@)

0 4 8 12— 16

Copulation Duration (Minutes)

Figure 2. Effects of copulation duration on probability that the female had at least one mature

oocyte at dissection (solid circles) and on probability of insemination (open circles; data not collected

for A. tomentosus at duration = 25 min). Solid circles having standard error bars are results for females

2002 HORTON ET AL.: ANTHOCORIS COPULATION 49

three species (Fig. 2, open circles; by y? tests, P < 0.01 in all three species).

Percentage of females having sperm was very high in the uninterrupted pairs: A.

tomentosus, 100% (n = 12); A. whitei, 91.7% (n = 12); A. nemoralis, 100% (n

= 17). Probability of insemination was often higher than probability of oocyte

maturation, particularly at the shorter copulation durations (Fig. 2: compare paired

open and filled circles), which may indicate that presence of sperm in the female

was not always sufficient to prompt oocyte maturation.

Size of the sperm pouch increased linearly with increasing duration of copu-

lation in all three species (Fig. 3; P < 0.005 for all three species); quadratic

effects were non-significant.

Study 2. Effects of Temperature.—Mean copulation duration in uninterrupted

pairs was significantly longer at 15° C than 25° C for all three species (Fig. 4; P

< 0.01 for all species [by f-tests]). Copulations averaged 79, 32, and 12 min

longer at 15° C than 25° C for A. whitei, A. tomentosus, and A. nemoralis, re-

spectively (Fig. 4).

Percentage of females that were inseminated or were characterized as having

at least one mature oocyte increased with increasing temperature and duration of

copulation (Fig. 5; statistical tests summarized in caption). For A. tomentosus,

females that were paired at 15° C invariably failed to mature their ovaries if

interrupted before finishing copulation (Fig. 5). Uninterrupted copulations tended

to result in insemination and oocyte maturation at both temperatures for all three

species.

DISCUSSION

While it is apparent that copulation duration is highly variable in the Insecta

(Thornhill & Alcock 1983, Eberhard 1996), the significance of this variation is

not well understood (Cook 1994, Eberhard 1996). If the function of copulation

were no more than a means to deliver sperm to the female, then one would expect

selection to favor brevity (Cook 1994, Eberhard 1996). That is, time spent in

copulation is not available for other activities, including functions such as feeding,

egglaying, or search for additional mates (Eberhard 1996). However, the fact that

copulation in many species occurs for time periods longer than necessary to trans-

fer sperm (Cordero 1990, Alcock 1994) suggests that sperm transfer is not the

sole function of copulation. Long-duration copulation may be favored for any of

a number of reasons, including as a means to increase certainty of paternity, to

from uninterrupted pairs. Curves are logistic regressions fitted to ovarian development data, excluding

uninterrupted pairs. Regressions are of the form: Prob. of ovarian development = exp(B) +

8,[duration])/(1 + exp(B,) + 8 ,[duration])), where By and B, are intercept and slope coefficients, re-

spectively. All three regressions indicated that probability of oocyte maturation varied with copulation

duration (by x?-tests; P < 0.01): By = —5.14, —2.01, and —1.58 for A. tomentosus, A. whitei, and A.

nemoralis, respectively; B, = 0.16, 0.043, and 0.40 for A. tomentosus, A. whitei, and A. nemoralis,

respectively. x?-tests also indicated that probability of insemination was significantly (P < 0.05) higher

than probability of ovarian maturation for A. tomentosus (at durations of 10, 20, and 30 min), for A.

whitei (at durations of 10 and 40 min), and for A. nemoralis (at duration of 4 min). Sample sizes 12—

20 females for each point except for virgin A. whitei (n = 4).

50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

@ Interrupted matings

oe °

A. tomentosus

First principal component

0 20 40 60 80 ~=100

0 2 4 6 8 10 12

Copulation duration (min)

Figure 3. Effects of copulation duration on mean (+ SEM) sperm pouch size (first principal

component obtained from principal components analysis conducted on pouch length and width).

2002 HORTON ET AL.: ANTHOCORIS COPULATION 51

reduce remating opportunities for females, or to transfer nutrients to offspring

(see discussions in Alcock 1994, Eberhard 1996).

In the present study, there was a 7-fold difference in mean copulation duration

between the species having the shortest mean duration (A. nemoralis) and the

species having the longest copulation (A. whitei). Too little is known about the

reproductive biology of Anthocoris spp. to do more than speculate about the

selective factors that may influence copulation duration in these three species.

However, some of the variation among species appears to be due to how rapidly

the male was able to transfer sperm and seminal products to the female once

intromission had occurred. For example, at 10 minutes following intromission,

the percentage of females having sperm in the sperm pouch was approximately

40%, 30%, and > 80% in A. tomentosus, A. whitei, and A. nemoralis, respectively

(Fig. 2). To reach the point at which 75% or more of females contained sperm

in the sperm pouch required copulation of approximately 4, 20, and 40 min in

duration for A. nemoralis, A. tomentosus, and A. whitei, respectively. Moreover,

measurements of sperm pouch size suggested that males transferred seminal prod-

ucts for most of the time that the insects were paired. That is, for all 3 species,

there was a significant linear increase in sperm pouch size with copulation du-

ration for inseminated females (Fig. 3). This result suggests that the longer cop-

ulations of A. whitei and A. tomentosus relative to duration in A. nemoralis were

not due to mate-guarding activities by male A. whitei and A. tomentosus. Instead,

species’ differences in duration apparently are due to differences in how rapidly

males of each species could transfer sperm, or in species’ differences in the

amount of seminal materials transferred.

By manipulating copulation duration, we showed that shortened copulations

had major effects on fitness. Probability of insemination and ovarian maturation

decreased with decreasing duration of copulation (Fig. 2), and quantity of seminal

material transferred by the male was reduced at the shorter durations (Fig. 3).

Other studies have monitored the effects of artificially shortened copulations on

fitness measures in insects. In several species, interrupted copulations have been

shown to result in reduced probability of insemination (Farias et al. 1972, Lew

& Ball 1980, Lachmann 1997), results similar to those reported here. Studies have

also shown that longer copulations may result in more sperm being transferred

(Yamagishi & Tsubaki 1990). We infer, based upon measurements of sperm pouch

size (Fig. 3), that number of sperm transferred to the female by males of Antho-

coris increased with increasing duration of copulation.

Mating is necessary to prompt oocyte maturation in Anthocoridae, and unmated

females in this family deposit few or no eggs (Anderson 1962, Shimizu 1967,

Carayon 1970). Almost no research has been done to determine what factors

Sample sizes are (see also Fig. 2): 9-14 (A. tomentosus), 9-12 (A. nemoralis), and 12-18 (A. whitei).

Sample sizes tend to be smaller than those provided in Fig. 2 because damage to the sperm pouch

during dissection occasionally prevented measurement of the pouch. Open symbols are results for

uninterrupted controls. Effects of copulation duration (by ANOVA; excluding uninterrupted controls):

A. tomentosus (F = 5.5; df = 4,48; P = 0.001 [linear: P = 0.004; quadratic: P = 0.13]); A. whitei

(F = 8.8; df = 5,82; P < 0.001 [linear: P < 0.001; quadratic: P = 0.18]); A. nemoralis (F = 16.2;

df = 4,51; P < 0.001 [linear: P < 0.001; quadratic: P = 0.11)).

52 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

A. nemoralis 15°C

y, mean = 25.4 min

A. tomentosus

15°C

mean = 72 min

A. whitei 15°C

mean = 181 min

Percentage of Pairs

> oy

Copulation Duration (min)

Figure 4. Frequency histograms showing distribution of copulation durations for A. tomentosus,

A. whitei, and A. nemoralis mating at two temperatures. Number of pairs per temperature: A. tomen-

tosus (50), A. whitei (30), and A. nemoralis (50).

associated with mating prompt ovarian development in these insects. In the pre-

sent study, probability of ovarian maturation was often lower than probability of

insemination at a given duration of copulation (Fig. 2). This result indicates that

intromission alone was not sufficient to prompt ovarian development, and strongly

suggests also that insemination was not always enough to lead to oocyte matu-

100 + A. tomentosus, P 100 | A whitei 100 4 A. nemoralis

: 25

~ oO

xo)

)

& 100

c Mam 15 Celsius Wmgg 15 Celsius |7

° 75 ZZZA 25 Celsius 25 Celsius

oO 50

10 min 30 min Uninterrupted 20 min 40 min_ Uninterrupted 2min 8min Uninterrupted

Copulation Duration Copulation Duration Copulation Duration

Figure 5. Percentage of dissected females containing sperm (upper panels) or having at least one

mature oocyte (bottom panels) as a function of copulation duration and temperature. Missing bars for

A. tomentosus indicate 0%. Number of females dissected (per bar): A. tomentosus (25), A. whitei (15),

and A. nemoralis (25). Insemination: A. tomentosus (temperature, y? = 12.2, P < 0.001; duration, y?

= 22.5, P < 0.001); A. whitei (temperature, x? = 3.4, P = 0.066; duration, x? = 10.2, P = 0.001);

A. nemoralis (temperature, y? = 7.5, P = 0.006; duration, yx? = 28.6, P < 0.001). Mature oocytes: A.

tomentosus data not analyzed due to 0% values; A. whitei (temperature, x? = 8.0, P = 0.005; duration,

x? = 2.0, P = 0.16); A. nemoralis (temperature, x? = 6.4, P = 0.01; duration, xy? = 25.3, P < 0.001).

Data analyzed as a 2 X 2 X 2 categorical model (analysis excluded uninterrupted controls; thus,

duration effects refer to comparison of short and intermediate copulations).

2002 HORTON ET AL.: ANTHOCORIS COPULATION a3

ration. The anthocorid egg lacks a true micropyle, and fertilization of the egg

takes place in the ovaries before the chorion is formed (Cobben 1968). Sperm

must move from the sperm pouch into a bridge of conductive tissue and then

eventually to the base of the ovaries (Carayon 1953). In the phylogenetically

related Cimicidae, insemination prompts activation of the corpus allatum, which

in turn controls maturation of the eggs in the female (Davis 1964, 1965). Sperm

must reach the base of the ovaries to result in activation. Sperm are often visible

at the base of the terminal oocyte in newly reproductive Anthocoris females (un-

published data), but whether that presence is sufficient by itself to prompt egg

development is unknown. Davis (1964) showed that seminal fluid was necessary

to prompt migration of sperm to the ovaries in Cimicidae, and it is possible that

by interrupting mating pairs of A. tomentosus, A. whitei, and A. nemoralis, that

we prevented the male from transferring sufficient amounts of seminal products

to prompt sperm migration in many females, even in those females that had been

inseminated.

Variation in copulation duration may have both genetic and environmental com-

ponents (Ward & Simmons 1991, Mihlhauser et al. 1996). One environmental

factor shown to affect copulation duration is temperature (Cook 1994). In this

study, copulation duration in Anthocoris spp. was substantially longer in pairs

mated at 15° C than those mated at 25° C (Fig. 4). Additionally, the magnitude

of fitness costs associated with shortened copulations was shown to differ between

matings done at the two temperatures. For example, percentage of females that

were inseminated following a copulation of short duration (20 min in A. whitei,

10 min in A. tomentosus, and 2 min in A. nemoralis) increased from 31 to 53%

(A. whitei), from 4 to 48% (A. tomentosus), and from 8 to 46% (A. nemoralis) if

mating was conducted at 25° C rather than 15° C (Fig. 5). In sum, for a copulation

of a given duration, probability of insemination and, hence, oocyte maturation

increased with increasing temperature.

Why should copulation duration be longer at 15° C than 25° C? The simplest

explanation would seem to be that males had physical difficulties inseminating

the female at the lower temperature. As noted above, males in the genus Antho-

coris transfer seminal products to the female’s sperm pouch via a thin copulatory

tube (Carayon 1953). The aedeagus of the male is fully extended through the

female’s tube during copulation. At 15° C, it may have been difficult for the male

to force seminal products through the aedeagus and copulatory tube. All three

species may inhabit geographic areas that experience quite cool temperatures (Kel-

ton 1978), particularly during spring and fall, and it seems almost certain that

these insects encounter daytime temperatures similar to the 15° C temperature

monitored here. If the insects mate in the field at this temperature, which remains

to be determined, then copulation duration in the field could vary substantially

within each species even under field conditions.

Finally, results of this study prompt some interesting questions regarding mat-

ing activity of these species under field conditions. Our assays indicate that pre-

mature termination of copulation in these species had fairly dramatic effects on

fitness. External factors that prompt premature break-up of a copulating pair may

result in the female failing to mature any eggs from that copulation. Such factors

prompting early termination could include the activities of natural enemies, activ-

ities of competing males, or decisions by the female to end the copulation (Eber-

54 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

hard 1996: 125). It remains to be determined whether any of these factors are

important in affecting the mating activities of these species under field conditions.

ACKNOWLEDGMENT

The comments of Carrol Calkins, Pete Landolt, Tom Unruh, and Tom Weissling

on an early draft of the manuscript are appreciated. We thank Dr. Donald Dahlsten

(U.C. Berkeley) for providing us with A. nemoralis several years ago; behavioral

observations made on those insects prompted the current study. The assistance of

Deb Broers, Carrie Colby, Richard Lewis, and Brian Reed in collecting and rear-

ing insects is greatly appreciated. The Winter Pear Control Committee provided

partial financial support.

LITERATURE CITED

Alcock, J. 1994. Postinsemination associations between males and females in insects: the mate-guard-

ing hypothesis. Ann. Rev. Entomol., 39: 1-21.

Anderson, N. H. 1962. Growth and fecundity of Anthocoris spp. reared on various prey (Heteroptera:

Anthocoridae). Entomol. Exp. Appl., 5: 40-52.

Carayon, J. 1953. Existence d’un double orifice génital et d’un tissu conducteur des spermatozdides

chez les Anthocorinae (Hemipt. Anthocoridae). C.R. Acad. Sci. Paris, 236: 1206-1208.

Carayon, J. 1970. Action du sperme sur la maturation des ovaires chez les Hémiptéres a insémination

traumatique. Coll. Internat. Centre National Recherche Scientifique, 189: 215-247.

Cobben, R. H. 1968. Evolutionary trends in Heteroptera. Part 1. Eggs, architecture of the shell, gross

embryology and eclosion. Centre Agric. Publ]. Documentation, Wageningen, The Netherlands.

Cook, D. E 1994. Influence of temperature on copula duration and mating propensity in Lucilia cuprina

Wiedemann (Diptera: Calliphoridae). J. Aust. Entomol. Soc., 33: 5-8.

Cordero, A. 1990. The adaptive significance of the prolonged copulations of the damselfly, Ischnura

graellsii (Odonata: Coenagrionidae). Animal Behav., 40: 43-48.

Davis, N. T. 1964. Studies on the reproductive physiology of the Cimicidae (Hemiptera)—I. Fecun-

dation and egg maturation. J. Insect Physiol., 10: 947-963.

Davis, N. T. 1965. Studies on the reproductive physiology of Cimicidae (Hemiptera)—II. Artificial

insemination and the function of the seminal fluid. J. Insect Physiol., 11: 355-366.

Dreistadt, S. H. & K. S. Hagen. 1994. Classical biological control of the acacia psyllid, Acizzia

uncatoides (Homoptera: Psyllidae), and predator-prey-plant interactions in the San Francisco

Bay area. Biol. Control, 4: 319-327.

Eberhard, W. G. 1996. Female control: sexual selection by cryptic female choice. Princeton University

Press, Princeton, New Jersey.

Farias, G. J., R. T. Cunningham & S. Nakagawa. 1972. Reproduction in the Mediterranean fruit fly:

abundance of stored sperm affected by duration of copulation and affecting egg hatch. J. Econ.

Entomol., 65: 914-915.

Horton, D. R., T. M. Lewis & T. Hinojosa. 2001. Copulation duration and probability of insemination

in Anthocoris whitei (Hemiptera: Anthocoridae) as a function of male body size. Can. Entomol.,

133: 109-117.

Kelton, L. A. 1978. The insects and arachnids of Canada. Part 4. The Antocoridae of Canada and

Alaska (Heteroptera: Anthocoridae). Canada Dept. Agric. Publ. No. 1639. 101 pp.

Lachmann, A. D. 1997. Sperm transfer during copulation in five Coproica species (Diptera: Sphaer-

oceridae). European J. Entomol., 94: 271-286.

Lattin, J. D. 1999. Bionomics of the Anthocoridae. Annu. Rev. Entomol. 44: 207-231.

Lew, A.C. & H. J. Ball. 1980. Effect of copulation time on spermatozoan transfer of Diabrotica

virgifera (Coleoptera: Chrysomelidae). Ann. Entomol. Soc. Am., 73: 360-361.

McMullen, R. D. 1971. Psylla pyricola Forster, pear psylla (Hemiptera: Psyllidae). Common. Inst.

Biol. Cont. Tech. Comm., 4: 33-38.

Miihlhauser, C., W. U. Blanckenhorn & P. I. Ward. 1996. The genetic component of copula duration

in the yellow dung fly. Anim. Behav., 51: 1401-1407.

2002 HORTON ET AL.: ANTHOCORIS COPULATION nf,

SAS Institute. 1987. SAS/STAT guide for personal computers (Version 6). SAS Institute, Cary, North

Carolina.

Shimizu, J. T. 1967. A biology of Anthocoris antevolens White, a Predator of Pear Psylla (Hemiptera:

Anthocoridae). M.S. Thesis, University of California, Berkeley.

Thornhill, R. & J. Alcock. 1983. The evolution of insect mating systems. Harvard University Press,

Cambridge, Massachusetts.

Ward, P. I. & L. W. Simmons. 1991. Copula duration and testes size in the yellow dung fly, Scatophaga

stercoraria (L.): the effects of diet, body size, and mating history. Behav. Ecol. Sociobiol., 29:

77-85.

Yamagishi, M. & Y. Tsubaki. 1990. Copulation duration and sperm transfer in the melon fly, Dacus

cucurbitae Coquillett (Diptera: Tephritidae). Appl. Entomol. Zool., 25: 517-519.

Received 24 November 2000; Accepted 20 September 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 56-61, (2002)

DIAERETUS ESSIGELLAE (HYMENOPTERA:

BRACONIDAE), A NEW SPECIES PARASITIC ON

ESSIGELLA PINE APHIDS (HOMOPTERA: APHIDIDAE)

FROM CALIFORNIA

P. STARY! & R. L. ZUPARKO?

Institute of Entomology, Academy of Sciences of the Czech Republic,

BraniSovska 31, 370 05 Ceské Budéjovice, Czech Republic

?Essig Museum of Entomology, University of California,

Berkeley, California 94720

Abstract.—Diaeretus essigellae n.sp., an aphidiine parasitoid reared from Essigella californica

(Essig) on Pinus spp. from California, is described. The new species is distinguished from other

Nearctic aphidiines by its reduced fore wing venation and the absence of notauli, propodeal

carinae and prongs on the female hypopygium. Diaeretus essigellae may be a useful biocontrol

agent against FE. californica in other countries.

Key Words.—Insecta, Hymenoptera, Braconidae, Diaeretus essigellae, Essigella californica, tax-

onomy, biocontrol.

In the Essig Museum of Entomology (University of California, Berkeley), the

junior author found specimens of an aphidiine species (Hymenoptera: Braconidae)

collected from California, which did not appear to match any other Nearctic gen-

era. After consulting with the senior author, we determined that it was a new

species very close to the monotypic Diaeretus leucopterus (Haliday), a Palearctic

species. We later found additional specimens of the new species in the Entomol-

ogy Research Museum at the University of California, Riverside, determined by

C. EF W. Muesebeck as “Diaeretus n. sp.’’. In this paper, we describe the new

species and indicate its potential as a biological control agent.

MATERIALS AND METHODS

Characters and terms follow those in Wharton et al. (1997). Except for two

females and one male which were remounted on microscope slides, all specimens

were point-mounted and examined through a stereomicroscope; measurements

were made with an ocular micrometer. All illustrations were made by the senior

author, using a camera lucida.

Depositories.—Type specimens will be deposited in the following institutions:

Australian National Insect Collection, CSIRO, Canberra, ACT, Australia (ANIC);

Bohart Museum, University of California, Davis, California (UCD); California

Academy of Sciences, San Francisco, California (CAS); Entomology Research

Museum, University of California, Riverside, California (UCR); Essig Museum

of Entomology, University of California, Berkeley, California (EME); National

Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM);

The Natural History Museum, London, United Kingdom (BNHM), and the per-

sonal collection of P. Stary, Institute of Entomology, Academy of Sciences of the

Czech Republic, BraniSovska, Czech Republic (PS).

2002 STARY & ZUPARKO: A NEW PINE APHID PARASITOID 57

DIAERETUS ESSIGELLAE STARY & ZUPARKO, NEW SPECIES

Types.—Holotype, female deposited USNM, data: USA. CALIFORNIA. RIV-

ERSIDE Co., Riverside, 21 Mar 1960, E.I. Schlinger, reared from Essigella “‘pini”’

on Pinus canariensis, (60-3-21a). Allotype, male deposited UCR, data: USA.

CALIFORNIA. LOS ANGELES Co. Lake Hughes, 22 May 1959, E.I. Schlinger,

reared from Essigella ‘“‘pini’”’ on Pinus ponderosae, (59-5-23k). Paratypes: USA.

CALIFORNIA. LOS ANGELES Co. same data as allotype, 3 females, 1 male.

MARIN Co. San Rafael, 4 Jun 1991, R.L. Zuparko, foliage of Tilia sp., 1 female.

MONO Co. White Mountains, Crooked Creek Lab., 10,100’ elev. 16-21 Aug

1984, R.F Gill, 1 female. RIVERSIDE Co. Lake Hemet, 18 May 1959, E.I. Schlin-

ger, from Essigella sp. on Pinus ponderosae? (59-5-18b), 1 male; Riverside, 30

Jan — 4 Feb 1925, P. Timberlake, from Essigella sp. on pine, 1 female, 2 males;

same data as holotype, 35 females, 25 males; 12 Apr 1960, E. I. Schlinger & J.

C. Hall, reared from E. californica on P. canariensis (60-4-12a), 8 females and

3 males; Riverside, California Experiment Station (= U.C. Riverside campus) 3

Mar 1960, E.I. Schlinger, reared from Essigella “‘pini”’ on Pinus canariensis (60-

3-3a), 5 females, 5 males; same data except 9 Mar 1960 (60-3-9a), 10 females,

9 males, same data except 23 Mar 1960 (60-3-23a), 17 females, 5 males; same

data except 27 Mar 1960 (60-3-27a), same data except 30 Mar 1960, E.I. Schlin-

ger & J.C. Hall (60-3-30m), 57 females and 61 males; Riverside, Citrus Experi-

ment Station (= U.C. Riverside campus), FG. Andrews, 30 Mar 1966, from Es-

sigella on Pinus canariensis (F66-3-30a), 4 females and 1 male; deposited ANIC,

CAS, BNHM, EME, PS, UCD, UCR, USNM.

Description—Female—Length 1.4—1.9 mm (n = 134). Head transverse, smooth, shiny and sparse-

ly haired, slightly wider than thorax at tegulae. Occipital carina distinct and complete. Ocelli form

isosceles triangle, ocell-ocular distance about 0.5X eye height, anterior-posterior ocellar distance about

2.0X ocellar width. Eye hairless, oval, slightly convergent ventrally, medium-sized, height 1.2 width.

Frons width about 0.5X eye width; temple width about 0.7 eye width; tentorio-ocular line about

0.25 eye width; malar space equal to tentorio-ocular line; intertentorial line 2.0 tentorio-ocular

line; genal length about equal width of mandible base. Mandible prominent, bidentate. Maxillary

palpus 4 segmented, labial palpus 2 segmented. Antenna inserted about midway up eye height, distance

from eye about 1.0X torulus width; 14 segmented, filiform, only slightly thickened to the apex,

reaching mid-metasoma; length Ist flagellomere (Fl) almost 4.0X width, with numerous short ad-

pressed setae, and sparse semi-erect setae which are about 0.75X Fl width, no placodes (Fig. 1); F2

subequal to Fl, 1 placode (Fig. 2); remaining flagellomeres very little wider than F2.

Mesoscutum smooth, shiny, notauli completely effaced, traced with sparse setae on disc, anterior

portion almost perpendicular to pronotum. Epicnemial carina present anterior-ventrally on mesopleu-

ron, extending to about 0.5X between mid- and hind coxae. Sternaulus absent. Transscutal articulation

deep and smooth. Scutellum smooth and rounded. Propodeum shiny, smooth, sparsely haired, without

carinae (Fig. 3).

Fore wing hyaline with reduced venation (Fig. 5): m-cu and r-m completely absent, RS short, not

reaching apex of stigma; stigma triangular, length about 3.0X width; R1 length about 0.5X stigma

length; Rs short, pigmented section not exceeding apex of stigma; M, M+Cu, 1A and Icu-a present

and pigmented, 1CU only faintly indicated basally. Disc completely covered with setae, somewhat

denser distad of M; length of apical-posterior marginal setae about 3.0 length discal setae. Hind

wing hyaline with complete basal cell; basal cell bare except few setae anteriorly, remainder of wing

uniformly covered with setae; posterior marginal setae about 0.33 wing width at hamuli.

Metasoma lanceolate, length about 1.2 length head and mesosoma. Petiole smooth, shiny with

sparse setae laterally on apical half (Fig. 4); almost parallel-sided, but markedly narrowed basad of

spiracles, almost 3.0X as long as broad at spiracles; dorsum slightly impressed laterally just posterior

to spiracles. Remaining segments smooth, shiny with sparse setae. Genitalia as figured (Fig. 6); ovi-

58 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

Placode

. /

Spiracle

longitudinal channel

Figures 1-7. Female Diaeretus essigellae. Figure 1. First flagellar segment. Figure 2. Second

flagellar segment. Figure 3. Propodeum. Figure 4. Petiole. Figure 5. Forewing. Figure 6. Female

genitalia. Figure 7. Ovipositor sheath.

positor sheath spatulate, apical setae with tubiform base, with an internal longitudinal darkened channel

(Fig. 7); ovipositor curved down.

Color medium brown; scape yellow, pedicel light brown but yellow ventrally, Fl light brown but

with narrow yellow ring basally, remaining segments successively darker towards the apex. Clypeus

slightly lighter than rest of face, mandibles and palpi yellow. Legs generally yellow, except: mid- and

hind coxa and hind femur distad of trochantellus and all Sth tarsal segments brown; apices of tibiae,

2002 STARY & ZUPARKO: A NEW PINE APHID PARASITOID 59

mid-femur and all 4th tarsal segments light brown. Apices of petiole and following segment with

slightly lighter areas; terminal metasomal segments slightly darker.

Male—Length 1.2—1.8 mm (n = 113). Generally as female, except: antenna with 15-16 segments,

length Fl 2.5-3.0X width, length F2 3.0X width; metasomal petiole tends to be slightly more con-

stricted basal of spiracles. Color of antenna and legs generally darker: scape and pedicel light brown,

base of Fl yellow, remainder of flagellum dark brown; hind trochanter and trochantellus yellow brown,

mid tibia and femur yellow brown except for lighter apices, all tarsal segments (except basitarsi)

yellow brown or at least darkly tinged.

Variations.—Out of 108 females with intact antennae, 107 were 14-segmented

and one was 15-segmented. Of 81 males with intact antennae, 22 were 15-seg-

mented, and 59 were 16-segmented. Smaller specimens (especially in the males)

tended to have a markedly greater constriction basad of the petiole spiracles, thus

giving the petiole more of an hourglass shape. Some individuals had a greater

degree of darkening on legs. The two individuals from Mono and Marin Counties

were markedly darker: body almost black; scape light brown, pedicel dark brown

and flagellum almost black; mandibles yellow brown; base of forecoxa, all of

mid- and hindcoxae concolorous with body, foretibia slightly darkened, midtibia

and femur brown, hindtibia and femur dark brown.

Diagnosis.—The new species is readily separated from all other Nearctic Aphi-

diinae by the following combination of characters: reduced fore wing venation

(m-cu and r-m completely absent, RS short, not reaching apex of stigma), me-

sonotum without notauli, propodeum without carinae, female hypopygium without

prongs. The species keys to couplet 14 in van Achterberg (1997), where Diaertus

Foerster is separated from Adialytus Foerster and Diaeretiella Stary. The new

species can be separated from D. leucopterus by the absence of the propodeal

carina, and from Adialytus and Diaeretiella by the absence of notauli.

Distribution.—Known only from California (Los Angeles, Marin, Mono, Riv-

erside, and Ventura Counties).

Host.—Essigella californica (Essig) (Homoptera: Aphididae) on Pinus canar-

iensis C. Smith, Pinus quadrifolia Parlatore ex. Sudworth, ?P. ponderosae Doug-

las and ?Pinus radiata D. Don.

Material Examined.—See Types. Two additional specimens (1 female, 1 male) in the senior author’s

collection have the data: USA. CALIFORNIA. VENTURA Co. Cuyama Valley, 22 May 1959, E.I.

Schlinger, from Essigella “‘pini” on ‘“‘Pinus cembroides parryana’’.

DISCUSSION

Mackauer & Stary (1967) reported an undescribed Diaeretus sp. in western

North America, undoubtedly based on the same series of specimens from the

Riverside collection which we have used to describe D. essigellae. The host of

these specimens was recorded to be Essigella pini Wilson. However Sorensen

(1994) demonstrated that E. pini is known only from eastern North America, and

that earlier records of this species from western North America refer to any of

several other Essigella spp. Unfortunately, an examination of the mummies from

which the new species emerged could not identify the host aphid more specifically

than “Essigella sp.”’ (J.T. Sorensen, personal communication). However, one se-

ries of specimens (60-4-12a) we examined were correctly labelled as being reared

from FE. californica, and others were from the same collection series (60-3-9a)

which were identified as E. californica (Essig) by Sorensen (1994), collected on

60 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

Pinus canariensis in Riverside. E. californica is the most polyphagous species of

the genus, and the only Essigella species reportedly reared from P. canariensis

in California (Sorensen 1994),

The two specimens from Ventura County were also recorded from E'ssigella

“pini’’, but on “‘P. cembroides parryana Voss’’, which is a synonym for P. guad-

rifolia (Munz & Keck 1968). Sorensen (1994) reported that, other than a single

record of E. fusca voegtlini Sorensen, the only Essigella species recorded from

this pine was E. hoerneri Gillete & Palmer (known from Ventura County), and

specifically noted that E. californica was not found on it. E. californica and E.

hoerneri are very closely related, difficult to distinguish from each other, and have

overlapping geographical distributions, although they have different favored host

plants (Sorensen 1994). We therefore believe it likely that D. essigellae will attack

both species.

In northern California, parasitized E. californica have been noted on Pinus

radiata (D.L. Dahlsten, personal communication). However, the adult aphidiines

responsible have yet to be collected. To date, we have seen only a single D.

essigellae from northern California (Marin County), with no host records.

Several species have been previously placed within the genus Diaeretus, but

Stary (1960) determined that it was monotypic, containing only D. leucopterus.

Based on this, one of the characters used to distinguish the genus was a propo-

deum with a carinate areola (Stary 1960, 1970; van Achterberg 1997). The ab-

sence of this areola in D. essigella initially led us to believe that this species

represented a new genus. However, in the closely related genus Pauesia Quilis

Pérez, there is a considerable variation in the completeness of the longitudinal

carinae on the propodeum, suggesting that the presence of propodeal carina is of

doubtful generic value. We expect many Nearctic aphidiine species still remain

to be discovered, leading to possible future nomenclatural changes, and felt it

better to adopt a conservative approach and keep the new species in an established

genus. It might be added that both Diaeretus species attack closely related hosts

(in the subtribe Eulachina) in similar ecological niches: D. leucopterus is a par-

asitoid of Eulachnus (= Protolachnus) spp. on conifers (Mackauer & Stary 1967,

Mackauer 1968, Stary 1970), while D. essigella attacks Essigella californica on

Pinus spp.

Pinus radiata has been exported throughout the world, with large plantings in

Australia, New Zealand, Chile, South Africa and Spain (Ohmart 1981). Recently,

E. californica has been reported in Spain and France (Tizado & Nutiiez-Pérez

1996, Turpeau & Remaudiére 1990), New Zealand, and as a major pest of P.

radiata in Australia (Carver & Kent 2000). In contrast, E. californica is not

considered a pest in California, though some sooty mold is associated with its

honeydew production (Burke 1937, Ohmart 1981) and some Essigella species

occasionally damage Christmas tree plantations (Sorensen 1994). Although the

efficacy of D. essigellae in suppressing E. californica populations has not yet

been determined, the relatively innocuous nature of the aphid in California sug-

gests it may be under effective biological control here. If so, D. essigellae may

prove to be valuable as an importable biocontrol agent.

ACKNOWLEDGMENT

We wish to thank John Sorensen for providing the generic identification of the

mummies of the host aphids, and Steve Heydon and Serguei Triapitsyn for the

2002 STARY & ZUPARKO: A NEW PINE APHID PARASITOID 61

loan of specimens. We express thanks to two anonymous reviewers who provided

suggestions for improving the manuscript. We are also grateful to Evert Schlinger

for his help in pinpointing the original collection site in Riverside. This work was

partially supported by the Academy of Sciences of the Czech Republic, Ento-

mology Institute Project Z 5/007/907.

LITERATURE CITED

Burke, H. E. 1937. Important insect enemies of the Monterey pine. Proc. Western Shade Tree Con-

ference, 4: 21-30.

Carver, M. & D. S. Kent. 2000. Essigella californica (Essig) and Eulachnus thunbergii Wilson (He-

miptera: Aphididae: Lachninae) on Pinus in southeastern Australia. Australian J. Entomology,

39: 62-69.

Mackauer, M. 1968. Aphidiidae. Pars 3. Hymenopterorum Catalogus (n. ed.). Ferriére, C. & J. van

der Vecht (eds.). Dr. W. Junk, The Hague.

Mackauer, M. & P. Stary. 1967. Hym. Ichneumonoidea, World Aphidiidae. Jn Delucchi, V. & G.

Remaudieére (eds.). Index of entomophagous insects. Le Francois, Paris.

Munz, P. A. & D. D. Keck. 1968. A California flora. University of California Press, Berkeley, Cali-

fornia.

Ohmart, C. P. 1981. An annotated list of insects associated with Pinus radiata D. Don in California.

CSIRO, Division of Forest Research, Divisional Report 8.

Sorensen, J. T. 1994. A revision of the aphid genus E'ssigella (Homoptera: Aphididae: Lachninae): its

ecological associations with, and evolution on, Pinaceae hosts. Pan-Pac. Ent., 70: 1-102.

Stary, P. 1960. The generic classification of the family Aphidiidae (Hymenoptera). Acta Soc. Ent.

Cechosl., 57: 238-252.

Stary, P. 1970. Biology of aphid parasites (Hymenoptera: Aphidiidae) with respect to integrated con-

trol. Dr. W. Junk, The Hague.

Tizado, E. J. & E. Niifiez-Pérez. 1996. Identificada Essigella californica (Hom.: Aphididae) sobre

repoblaciones de Pinus radiata en Espafia. p. 170. In VII Congreso Ibérico de Entomologia.

Santiago de Compostela 19—23 de Septiembre 1996.

Turpeau, E. & G. Remaudiére. 1990. Decouverte en France d’un puceron des pins americains du genre

Essigella. C.R. Acad. Agric. France, 76: 131-132.

van Achterberg, C. 1997. Subfamily Aphidinae. pp. 119-131. In Wharton, R. A., PR. M. Marsh & M.

J. Sharkey (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera).

Intern. Soc. Hymen., Special Publ. 1.

Wharton, R. A., P- M. Marsh & M.J. Sharkey (eds.). 1997. Manual of the New World genera of the

family Braconidae (Hymenoptera). Intern. Soc. Hymen., Special Publ. 1.

Received 11 May 2001; Accepted 29 October 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 62, (2002)

Scientific Note

THE IMMIGRANT LEPTOPODID, PATAPIUS SPINOSUS

(ROSSD, IN OREGON (HEMIPTERA: HETEROPTERA:

LEPTOPODIDAE)

Usinger (1941. Bull. Brooklyn Entomol. Soc., 36: 164-165) first reported this

non-indigenous species from North America, based on specimens from California.

Brothers (1979. Great Basin Nat., 39: 195-196) published on its occurrence in

Idaho and Nevada and included a drawing of the adult. Recently, Zack et al.

(2001. Pan-Pac. Entomol., 77: 47-50) added Washington as a locality and pro-

vided observations on the habitat and habits of this remarkable bug in eastern

Washington. The collections of the Oregon Department of Agriculture, Salem,

Oregon, contain a specimen of Patapius spinosus (Rossi) with the following in-

formation on it: OR, Rogue River, 10 III 1959, under pine bark, K. Goeden coll.

Also included in their collection are specimens bearing the following information:

CA, Glenn Co., Willows, 12 XII 1962, under boards, Haig Dudley coll. Previ-

ously, I have seen many specimens from a xeric site in California where speci-

mens were collected beneath the loose bark of a dead, fallen tree—far from any

water. Lindskog (1995. pp. 137-140. Jn Aukema & Rieger (eds.). Cat. Heter. Pal.

Reg. 222 pp.) indicated that species of Patapius Horvath may be found under the

bark of trees well removed from water: species of other genera of Leptopodidae

are known to live in habitats much like that of species of Saldidae—close to

water. The movement of grape stock in the earlier days of California grape cul-

tivation was a possible source of individuals of this predatory bug. Subsequent

movement of grape stock and other vegetative and road building materials (see

Zack et al. 2001) likely contributed its further distribution in western North Amer-

ica. It may be expected in other disturbed sites throughout the region.

Acknowledgment——My thanks to J. LaBonte, O.D.A., for the information of

the Oregon specimen.

J. D. Lattin, Department of Entomology, Oregon State University, Corvallis,

OR, 97331-2907.

Received 10 May 2001; Accepted 20 August 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 63-65, (2002)

Scientific Note

METOPOPLAX DITOMOIDES (COSTA), A SPECIES OF

OXYCARENIDAE NEW TO NORTH AMERICA

(LYGAEOIDEA: HEMIPTERA: HETEROPTERA)

Metopoplax ditomoides (Costa) is reported for the first time as an established

species in the United States. The only other record was its interception in Mas-

sachusetts in excelsior in a shipment of goods from Spain (USDA, Plant Pest

Contr. Div., Agric. Res. Ser. 1958. Coop. Econ. Ins. Rep., 8(42): 907). This was

the sole citation of M. ditomoides from North America cited by Slater (1964.

Univ. Conn., Storrs, Conn. U.S.A. 778 pp.), and it was not listed from the United

States by Ashlock and A. Slater (1988. Cat. Het. pp. 167—245) or by Slater and

O’Donnel (1995. N.Y. Entomol. Soc., New York. 410 pp.) Both males and females

of this distinctive lygaeoid were beaten from commercially grown hazelnut trees

(Corylus avellana (L.)) grown on a farm near Monmouth, Oregon during August

1998 by K. Wetherill. These specimens were part of a survey of insects on cul-

tivated and native species of Corylus in western Oregon by Wetherill (2000. M.S.

Thesis, Oreg. St. Univ., Corvallis. 123 pp.). Only adults were taken on these trees.

Additional specimens were brought in for identification in February 2000. These

specimens included a specimen recovered from a Lindgren Funnel Trap placed

by the Oregon Department of Agriculture at Lowell Road, vicinity Jasper, Lane

County, Oregon, 28 July 1999 (ex. J. LaBonte); adult individuals swarming on

cars and a mailbox near Corvallis, Benton County, Oregon, 7 February 2000 (ex.

L. Royce); hundreds of adults swarming in a farmhouse near Gervais, Marion

County, Oregon, 11 February 2000 (ex. D. McGrath) and adults swarming in a

home in Amity, Polk County, Oregon, 14 February 2000 (ex. L. Royce). These

records indicate the species is now well established in Oregon and likely to be

found elsewhere.

Slater (1964) provided a bibliography of M. ditomoides from its description in

1843 and the reader is referred to that publication for complete coverage. This

species 1s widespread in parts of southern Europe and northern Africa. The initial

specimen found in England was taken at Hounslow Heath, Middlesex, on a “‘rub-

bish tip up”’ (Woodroffe, 1953a. The Entomologist, 86: 34—34a, 1953b. 86: 224—

225) suggesting a recent introduction as the fauna of that country has been long

known. Woodroffe (1953b) included a drawing of the adult and Southwood and

Leston (1959. Land & water bugs of the British Isles. 436 pp.) discussed the

species and included an illustration. Metopoplax ditomoides resembles species of

Crophius but is more slender in shape. (See Slater & O’Donnel, 1995, p. 74 for

discussion of status of Crophius) and see Hoberlandt (1987. Acta Entomol. Mus.

Prague, 42: 11—30) about possible synonymy of Crophius with Anomaloptera

Amyot & Serville. The small size (2.8—3.4 mm); swollen, spatulate clypeus; dis-

tinctive black head, pronotum, and scutellum, together with the pale, almost white

forewings with distinctive dark veins make the recognition easy (Fig. 1.) Wood-

roffe (1953b) indicated that Matricaria maritima (L.) and M. chamomilla (L.)

were common on the site in England. Both species of plants are known intro-

64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1)

Figure 1. Metopoplax ditomoides (Costa) (after Woodroffe, 1953b).

ductions into the Pacific Northwest and are contained in the Oregon State Uni-

versity herbarium from ballast dumps and other localities in Oregon (Scott Sun-

berg, personal communication, July 1999). This suggests the possibility of a pri-

mary host to be a forb with movement onto Corylus later in the season. Meto-

poplax Fieber belongs to the Lygaeoidea family Oxycarenidae, a group of seed

bugs containing important pests of cotton and other crops (Samy. 1969. Trans.

Royal Entomol. Soc. London, 121: 79-165). Knowing this, one should be aware

of the possible pest status of Metopoplax ditomoides. Crophius Stal and Dycod-

erus Uhler are the only native genera of the Oxycarinidae found in the United

2002 SCIENTIFIC NOTES 65

States and Canada (Ashlock & A. Slater, 1988). We have taken at least one species

of Crophius on pine trees in Oregon (J.D.L.), indicating that related taxa also

may be found on trees.

The discovery of this insect is a reminder that introductions still occur and may

include species of potentially economic importance. Asquith & Lattin (1991. Pan-

Pacif. Entomol., 67: 258—271) reviewed the species of Lygaeoidea that had been

introduced into the Pacific Northwest when Metopoplax ditomoides had not yet

been recovered. Wheeler & Stoops (1999. Pan-Pacif. Entomol., 75: 52—54) report

Chilacis typhae (Perrin) (Lygaeoidea: Artheneidae) from Oregon and Washington.

This brings the number of introduced seed bugs in the Pacific Northwest to six.

Henry & Adamski (1999. J.N.Y. Entomol. Soc., 90: 275-301) reported on Rhy-

parochromus satunis (Rossi) (Lygaeoidea: Rhyparochromidae) from California,

adding yet another introduced species to the Pacific Coast region.

The Pacific Northwest species of Lygaeoidea may be arranged as follows: Ar-

theneidae: Chilacis typhae (Perrin); Oxycarenidae: Metopoplax ditomoides (Cos-

ta); Rhyparochromidae: Plinthisus brevipennis (Latreille); Megalonotus sabulicola

(Thomas); Stygnocoris rusticus (Fallén), and Stygnocoris sabulosus (Schilling)

(Henry. 1997. Ann. Entomol. Soc. Amer., 90: 271-301). Specimens of M. dito-

moides have been deposited at Oregon State University and at the Oregon De-

partment of Agriculture, Salem.

Acknowled gment.—We thank the Oregon Filbert Commission for support of

this project; M.T. AliNiazee for interest and guidance of the larger project; S.

Sunberg for helpful information on introduced plants now found in Oregon; to

D. McGrath, J. LaBonte, and L. Royce who provided additional specimens of this

bug from Oregon localities; two reviewers for their efforts and comments; and L.

Parks for assistance in the preparation of this paper.

This scientific note is dedicated to my long-time friend and colleague, Dr. James

A. Slater, whose extensive scientific efforts on the Lygaeoidea have provided us

with a superb foundation of all future work on this major taxon.

John D. Lattin,! Department of Entomology, Oregon State University, Corvallis,

Oregon 97331-2907 and Karen Wetherill,? University of New Mexico, Sevilleta

Long-Term Ecological Site, Albuquerque, New Mexico 87131-1091.

Received 28 November 2000; Accepted 5 April 2001.

PAN-PACIFIC ENTOMOLOGIST

78(1): 66-67, (2002)

Scientific Note

NEW HOST PLANT AND DISTRIBUTIONAL RECORDS

FOR SOME EBURIA LEPELETIER & AUDINET-SERVILLE

(COLEOPTERA: CERAMBYCIDAE) IN NORTH AMERICA

INCLUDING MEXICO

Cerambycids of the genus Eburia Lepeletier & Audinet-Serville are most in

evidence in the American tropics (Monné, M. A. 1993. Catalogue of the Cer-

ambycidae (Coleoptera) of the Western Hemisphere, Part II. Soc. Brasil. Entomol.,

Sao Paulo, Brasil). Only 10% of nominal Eburia species occur in the Nearctic

region, where they breed in trees belonging to several angiosperm families (Lin-

sley, E. G. & J. A. Chemsak. 1997. The Cerambycidae of North America, Part

VUI. Univ. Calif. Publ. Entomol., 117). This note provides unknown or unpub-

lished larval hosts and distributional records for 4 species of Eburia from the

southern United States and Mexico. Data come from field studies in southeastern

Texas and specimens in the TAMU collection. Plant nomenclature follows S. D.

Jones et al. (1997. Vascular plants of Texas. Univ. Texas Press, Austin, Texas);

collection codens follow R. H. Arnett, Jr. et al. (1993. The insect and spider

collections of the world (2nd ed.). Sandhill Crane Press, Gainesville, Florida).

Eburia haldemani LeConte. One female was reared from Ulmus crassifolia Nuttall

(Ulmaceae): TEXAS. GUADALUPE Co.: 10 km E of Seguin, 9 Jun 1988 (TAMU).

G. B. Vogt (1949. Pan-Pacif. Entomol., 25: 137-144) recorded this tree as a putative

larval host for E. haldemani. E. G. Linsley and J. A. Chemsak (1997) list E. hal-

demani as feeding in Morus sp. (Moraceae) and Celtis sp. (Ulmaceae).

Eburia linsleyi Lacey. Several adults were collected in ARIZONA. GRAHAM

Co.: Turkey Creek, 22 Jun 1976, S. McCleve (TAMU), and NEW MEXICO.

HIDALGO Co.: Indian Creek, Animas Mts, 1737 m, 5—6 Aug 1976, S. McCleve

(TAMU); Godfrey Place, Animas Mts, 1706 m, 7 Jul 1980, S. & J. Dobrott

(TAMU). These localities represent the easternmost records for the known distri-

bution of E. linsleyi (Monné 1993).

Eburia mutica LeConte. Adults of this Tamaulipan species were taken from

beneath loose bark and one eclosed from a pupa cut from dead trunk of young

Acer negundo L. var. negundo (Aceraceae): TEXAS. HARRIS Co.: 29.76° N,

95.44° W, 29 Jul 2000, P. Szafranski, 2 (MCZC). One male specimen of E. mutica

was also reared from Celtis lindheimeri Engelmann ex K. Koch: TEXAS. HI-

DALGO Co.: Bentsen-Rio Grande Valley State Park, 11—17 Jun 1977, R. Turnbow

(TAMU). Previously, E. mutica has been found developing in Havardia sp., Leu-

caena sp., Prosopis sp., Sesbania sp., Sophora sp. (Fabaceae), Citrus spp., Zan-

thoxylum sp. (Rutaceae), Celtis laevigata Willdenow and Ulmus sp. (Ulmaceae)

(Linsley & Chemsak 1997). The locality of E. mutica-A. negundo association

marks the northeastern range of EF. mutica (Fig. 1) and the southwestern edge of

the continuous, natural range of A. negundo. Other records that determine the

northern range of EF. mutica include: TEXAS. BRAZOS Co.: College Station, Jun

1937 & 7 Jun 1977, J. R. Ables (TAMU); GALVESTON Co.: Dickinson, 21 May

2002 SCIENTIFIC NOTES 67

00°

Figure 1. Updated geographical distribution of Eburia mutica (open triangles) and E. stigmatica

(filled triangles, S—state record). Lower Rio Grande valley symbols represent several nearby locales.

& 27 Jun 1932, J. N. Roney (TAMU); TRAVIS Co.: Austin, Jul 1923 (TAMU);

VAL VERDE Co.: Langtry, 21 Jun 1990, E. Riley & C. Wolfe (TAMU).

Eburia stigmatica Chevrolat. A female specimen was chopped from Chloro-

leucon ebano (Berlandier) Rico (= Pithecellobium flexicaule (Bentham) Coulter)

(Fabaceae): TEXAS. CAMERON Co.: 5 km SW of Olmito, 24 May 1994, D. J.

Heffern (TAMU). C. laevigata is the only other known host of E. stigmatica

(Linsley & Chemsak 1997). This subtropical species was also collected at several

localities that extend its range from the lower Rio Grande valley, Texas and

northeastern Mexico (Monné 1993) to the Pacific coast of southern Mexico and

southeastwards through Yucatan peninsula (Fig. 1): MEXICO. OAXACA: 19.3 km

W of Jalapa del Marques, 12 Jul 1971, Clark et al. (TAMU); 16.7 km N of

Niltepec, 15 Jul 1971, Clark et al. (TAMU). QUINTANA ROO: Apr 1963

(TAMU). Y UCATAN: Chichen-Itza, 19 Aug 1965, N. J. Dickey (TAMU); same

data except 10-11 Jun 1983, E. Riley (TAMU).

Acknowled gment.—Thanks are extended to Ed Riley for helpful assistance and

access to the cerambycid materials at the Texas A&M University Insect Collection.

Przemyslaw Szafranski. Developmental Biology Section, Department of Pa-

thology, Baylor College of Medicine, Houston, Texas 77030.

Received 5 May 2001; Accepted 17 October 2001.

PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

See volume 74: 248-255, October 1997, 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 INDOUBLE 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. 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.

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 | 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. FIGURE 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 pennitted 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 78 January 2002 Number 1

Contents

LANDOLT, P. J——Survival and development of Lacanobia subjuncta (Grote & Robinson)

(Lepidoptera: Noctuidae) larvae on common weeds and crop plants of eastern Washing-

ton state

CABRERA, B. J., P. M. MARSH, V. R. LEWIS, & S. J. SEYBOLD—A new species of

Heterospilus (Hymenoptera: Braconidae) associated with the deathwatch beetle, Hezi-

coelus gibbicollis (Leconte) (Coleoptera: Anobiidae)

AREEKUL, B. & D. L. J. QUICKE—A new species of Yelicones Cameron (Hymenoptera: Bra-

conidae) from Thailand

VETTER, R. S. & T. R. PRENTICE—tThe spider fauna associated with litter under woodrat

middens in Southern California (Arachnida: Araneae)

TRIAPITSYN, S. V., L. G. BEZARK, & D. J. MORGAN—Redescription of Gonatocerus atri-

clavus Girault (Hymenoptera: Mymaridae), with notes on other egg parasitoids of sharp-

shooters (Homoptera: Cicadellidae: Proconiini) in northeastern Mexico

HORTON, D. R., T. M. LEWIS, & T. HINOJOSA—Copulation duration in three species of

Anthocoris (Heteroptera: Anthocoridae) at different temperatures and effects on insemi-

nation and ovarian development

STARY, P & R. L. ZUPARKO—Diaeretus essigellae (Hymenoptera: Braconidae), a new

species parasitic on Essigella pine aphids (Homoptera: Aphididae) from California

SCIENTIFIC NOTES

LATTIN, J. D—The immigrant leptopodid, Patapius spinosus (Rossi), in Oregon (Hemiptera:

Heteroptera: Leptopodidae)

LATTIN, J. D. and K. WETHERILL—Metopoplax ditomoides (Costa), a species of Oxycareni-

dae new to North America (Lygaeoidea: Hemiptera: Heteroptera)

SZAFRANSKI, P.—New host plant and distributional records for some Eburia Lepeletier &

Audinet-Serville (Coleoptera: Cerambycidae) in North America including Mexico

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 78 April 2002 Number 2

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)