The Pan-Pacific Entomologist

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OFFICERS FOR 1995

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THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly by the Pacific

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PAN-PACIFIC ENTOMOLOGIST

71(2): 75-77, (1995)

OBITUARY:

RICHARD K. ALLEN (1925-1992)

GEORGE F. EDMUNDS JR.! AND CHAD M. MurvoSH?

\Dept. of Biology, The University of Utah,

Salt Lake City, Utah 84112

2Dept. of Biological Sciences, University of Nevada Las Vegas,

Las Vegas, Nevada 89154 |

Richard Knapp “Dick” Allen, a leading mayfly systematist, died at his home

in Lake San Marcos, California on 7 Aug 1992. He was born in Salt Lake City,

Utah on 21 Apr 1925, to Knapp R. and Leora Allen. After serving in the U:S.

Army in the Korean War, he obtained his B.S., M.S. and Ph.D. degrees at the

University of Utah (M.S. in 1955, Ph.D. 1960). He then served a National Science

Foundation supported post-doctoral during which a revision of the then very large

mayfly genus Ephemerella (over 80 species) was completed for North America.

His solution of the Drunella grandis—spinifera complex (then in Ephemerella) was

a major accomplishment. He taught at the University of California Los Angeles

in 1963, replacing the late Dr. John N. Belkin who was on leave. In 1964, he

joined the faculty of California State University at Los Angeles where he taught

a variety of courses, including marine invertebrate biology, from 1964 to 1976.

He suffered at least 3 myocardial infarctions between 1973 and 1978 and under-

went angioplasty to open up heart arteries. In 1976 he was operated on for colon

cancer and as related by Dick, “‘while still recovering from anesthesia the physician

sat at my bedside and told me to get my affairs in order as I had about six months

to live.”’ At this time he took a medical retirement from the University and was

given intense chemotherapy treatments.

Even as an undergraduate, Dick was devoted to exercise and physical fitness,

going to a gymnasium almost daily. Part of his recovery from heart circulatory

problems and colon surgery included increasing long rides on his bicycle, until

he rode 30 miles or more each day. The doctor’s predictions were obviously a

misjudgment; Dick lived 16 more years during which time he produced 23 papers.

Although he suffered numerous other medical problems, he did not seek sympathy.

Both of his hips were replaced as a consequence of an inherited hip socket de-

terioration disorder. In 1991 he had coronary bypass surgery to supply blood to

his heart and lengthen his useful life. He died following a relatively short illness

with untreatable pre leukemia (not a precursor of leukemia).

He published a well illustrated book, Common Intertidal Invertebrates of

Southern California, that was a guide to marine invertebrates of Southern Cali-

fornia, and 66 papers on the systematics and distribution patterns of maryflies,

principally of the southwestern United States and Mexico. His research on mayflies

was supported by a series of Systematic Biology NSF grants and later by his own

funds.

He was sole author of 26 papers, and published the others with ten different

co-authors. These include George Edmunds (Major Professor, Univ. Utah), and

colleagues, Chad Murvosh (Univ. Nevada Las Vegas), W. L. Peters (Florida A&M

76 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

R. K. Allen (left) on his sailboat at Dana Point, California with C. M. Murvosh.

Univ.), Lewis Berner (Univ. of Florida), the late Selwyn Roback (Philadelphia

Academy of Sciences) as well as his students at California State University Los

Angeles, Richard Brusca, Janet I. Kilgore, Esther S. M. Chao, Sandra D. Cohen

and D. L. Collins.

Dick Allen was a very social man and had a host of devoted friends. He was

1995 EDMUNDS & MURVOSH: R. K. ALLEN OBITUARY 77

an amiable and hard working field companion. At times, some colleagues became

a target of his criticism although the cause was often trivial, perhaps a difference

of taxonomic opinion or procedure. One of us (Edmunds) knew him throughout

his career (although not closely since 1976) and the other (Murvosh) had a close

relationship with him since 1978. We believe his occasional difficulties with

colleagues related strongly to his many health problems. He was an opinionated

man who worked hard and played hard, but he was normally a good and fun

loving friend. His often brusque exterior kept many persons from knowing well

his warmth, compassion, and generosity.

Dick traveled and collected widely in the United States and Mexico in a motor

home, camping near streams or rivers. In Mexico where he traveled by himself

or with Murvosh, he carried cans of dog food which he used to enlist a wandering

“perro” as a watch dog. He gave the dog a can of food and a bowl of water at

night and one in the morning and thus always had a barking sentry if anyone

approached his motor home.

He married Mary Anne Ruzicka in 1951. They were subsequently divorced but

remained good friends and were later remarried. Richard is survived by his widow,

his daughter, Mrs. Jody Anne Varner (whom he honored with the name Lepto-

hyphes jodiannae) and two granddaughters.

Most of his studies were concentrated on the mayfly families Tricorythidae in

the Americas and Ephemerellidae on a world basis but he worked widely on other

North and Central American species. He concentrated on the mayfly fauna of

Mexico and the southwestern United States. He was the author or co-author of

151 species, 13 genera or subgenera, and 2 subtribes. The writers have prepared

a bibliography of Dick’s mayfly papers and an annotated list of the taxa he named.

Copies are available on request to either author. A major concern during the last

few days he was able to speak was that the specimens that he had borrowed were

to be returned to the collections which owned them, and that his own specimens

be given to the California Academy of Sciences in San Francisco. His wife and

daughter delivered his remaining collections to Chad Murvosh at the University

of Nevada Las Vegas. The authors will attempt to finish the job of distributing

them but any curator who believes Dr. Allen’s specimens included borrowed

material under their care should contact one of the writers.

Dick Allen “retired” from studying mayflies several times but actually contin-

ued his studies throughout his life. Much of his collection, including the specimens

collected in Mexico by Allen and Murvosh are already deposited at the California

Academy of Sciences.

PAN-PACIFIC ENTOMOLOGIST

71(2): 78-81, (1995)

A NEW GENUS OF HEBRIDAE FROM

CHIAPAS AMBER (HETEROPTERA)

J. T. POLHEMUS

University of Colorado Museum, Englewood, Colorado 80110

Abstract. —Stenohebrus glaesarius NEW GENUS, NEW SPECIES, the only recorded fossil he-

brid, is described from Chiapas Amber and compared to modern hebrids.

Key Words. —Insecta, Heteroptera, Hebridae, fossil

The rich collection of fossil insects in Chiapas Amber held by the University

of California has provided startling discoveries of previously unknown fossil taxa

(Petrunkevitch 1971), which now include the only known fossil hebrid described

below. A general account of the fossiliferous amber of Chiapas was given by Hurd

et al (1961). The specimen described here was mentioned by Andersen (1982:

252) and illustrated by Poinar (1992: 115) in his book on amber inclusions.

The unique well preserved specimen is enclosed in a small block of amber that

has been cut and polished on all sides except the slightly curved surface (designated

top) touching or immediately adjacent to the dorsal surface of the insect, parts of

which lie at or just beneath the surface at a slight angle (Fig. 1). The top surface

is Somewhat roughened, pocked and degraded, especially over the vertex of the

head and thoracic dorsum, probably because of the proximity of the insect. Ad-

ditionally there are air inclusions, especially on the dorsal surface of the specimen,

which obscure some fine features, in particular those of the abdominal dorsum.

These imperfections obliterate or obscure some of the more interesting features

of the insect. The left hemelytron is spread or separated from the abdomen, and

lies very close to the top surface of the amber block in a quite clear area, thus the

venation and the pattern of the membrane are visible. The sternum of the insect

(Fig. 2) has a few air inclusions, but most details are quite clearly visible from

one of the faceted sides, although the angle of the insect requires turning in various

directions to get a clear view of certain fine features (e.g., the bucculae).

The morphological terminology follows that of Andersen (1981, 1982). All

measurements are in mm unless otherwise noted.

STENOHEBRUS J. T. POLHEMUS, NEW GENUS

(Figs. 1-3)

Type-species: Stenohebrus glaesarius J. T. Polhemus, NEW SPECIES

Description. —Macropterous form: Small, elongate oval, pubescence not visible. Length, 1.83 mm,

width 0.67 mm. Head long, porrect, only slightly declivent but narrowed anteriorly, large ventral lobe

projecting anteriorly; antennal tubercles very large, produced laterally to outer eye level; dorsal tricho-

bothria not visible; vertex obscured. Eyes globose, exserted, very prominent, adjacent to pronotum,

ommatidia large, coarsely faceted, about 30 in number, ocular setae not visible; ocelli obscured.

Antennae approximately half as long as body; segment 1 stout, shorter than head; segment 2 more

slender, about 4 as long as 1; segments 3 and 4 long, flagelliform, set with scattered long setae, without

obvious spines. Venter of head with long carinate bucculae, strongly developed, produced posteriorly

(Fig. 3, arrow); labium long, reaching beyond posterior coxae. Thoracic dorsum mostly obscured.

Pronotum longest on midline, strongly bilobed, humeri prominent. Paired ventral thoracic carinae

1995 POLHEMUS: A NEW HEBRID GENUS ee

well developed, parallel throughout, continuing separately onto base of abdomen. Femora stout,

unarmed; tibia slender, unarmed except for spur of stiff setae distally; tarsi 2 segmented, first segment

shortest, second segment distally set with long setae. Claws long, very slender, with prominent basal

spur; arolia, parempodia not evident. Abdomen longer than broad, not depressed ventrally. Female

first gonocoxae large, prominent, typical of Hebrus spp.

Diagnosis. — This genus differs from known extant hebrids by the protuberant

eyes, strongly laterally produced antennal tubercles, long slender claws, and ap-

parent lack of arolia. Stenohebrus is most similar to Hebrometra Cobben, sharing

the basal spur on the claws, but in the latter the rostral cavity is closed posteriorly

and the bucculae are reduced, not produced posteriorly, whereas in Stenohebrus

the rostral cavity is open posteriorly, and the bucculae are well developed and

strongly produced posteriorly.

Discussion. —In Andersen’s (1981) key to the genera and subgenera of Hebridae,

Stenohebrus keys to couplet 9, where the laterally produced antennal tubercles

ally it with Hebrus subgenus Timasielliodes Poisson from the Ethiopian and Ori-

ental regions, but other features discussed above place it closer to Hebrometra

Cobben. The head is produced anteriorly as in Hyrcanus Distant.

Etymology.—The name Stenohebrus is derived from stenos, Gr. Narrow, and

the nominate genus Hebrus. Masculine.

Material Examined. —Stenohebrus glaesarius.

STENOHEBRUS GLAESARIUS J. T. POLHEMUS, NEW SPECIES

(Figs. 1-3)

Type. —Holotype, macropterous female, data: MEXICO. Chiapas Amber, U.

C. Mus. Paleo. No. 12894, Oligocene-Miocene boundary, in Paleontological Mu-

seum, University of California, Berkeley.

Description, Macropterous Female. —(See generic description; only additional details given here.):

Elongate, apparent ground color brownish black; legs, rostrum, antennae brownish. Eyes brown,

moderately large, strongly exserted, coarsely faceted. Structural characteristics: Head long, porrect,

only slightly declivent anteriorly; median length (from above) 0.30 mm, width across antennal tubercles

0.28 mm; width across eyes, 0.28 mm; vertex mostly obscured. Eyes moderately large, exserted, with

about 30 ommatidia; width of an eye: interocular space, 0.06: 0.16. Ocelli obscured. Antennae long,

slender, segment I stoutest, abruptly widened basally, then parallel sided; segment IJ more slender

basally, widening distally; segments IIIJ—V long, slender, with scattered long setae; length of segments

I-V: 0.18 mm: 0.14 mm: 0.26 mm: 0.11 mm: 0.26 mm (separation between IV—-V broad, indistinct).

Rostrum extending between hind coxae. Bucculae very prominent, produced posteriorly. Pronotum

long, length approximately 0.35 mm, constricted medially, dorsal sculpturing obscured. Collar prom-

inent, set off with a row of pits. Mesoscutellum short, length 0.05 mm. Metanotal elevation triangular,

rounded distally, shallowly notched distally; median carina not visible; length 0.16 mm, width 0.30

mm. Legs slender, covered with short setae plus tibia distally set with long stiff setae; distal extreme

of all tibia set with a short curved spur of closely packed setae; proportions as follows: femur, tibia,

tarsal 1, tarsal 2 of fore leg, 0.35: 0.37: 0.03: 0.12; of middle leg, 0.41: 0.41: 0.03: 0.14; of hind leg,

0.53: 0.53: 0.03: 0.15. Hemelytra long, reaching tip of abdomen; hind wings obscured; clavus lightly

tinged with brown; corium smoky, indistinct; membrane fumose, with lighter spot near distal angle

of closed proximal cell; costal margin set with outwardly directed stiff setae. Abdominal venter sparsely

clothed with short appressed setae. First gonocoxae large, plate-like, covering all but triangular proc-

tiger.

Diagnosis. —Genus monotypic, see that diagnosis.

Discussion.—The slender claws with a prominent basal spur are similar to

semiaquatic Heteroptera living on damp earth or hygropetric habitats, e.g., He-

80 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Figures 1-3. Stenohebrus glaesarius. Figure 1. Dorsal view. Figure 2. Ventral view. Figure 3. Ventral

view, detail; posteriorly produced bucculae, arrow.

brometra spp., Ochterus spp., thus it seems likely that this species ranged away

from the waters edge.

Etymology.—The name glaesarius, L., of amber, refers to the origin of the

species in Chiapas Amber.

Material Examined. —See type.

1995 POLHEMUS: A NEW HEBRID GENUS 81

ACKNOWLEDGMENT

Iam indebted to the late P. D. Hurd Jr. of the University of California, Berkeley,

and more recently the Smithsonian Museum, for the opportunity to study this

interesting fossil.

LITERATURE CITED

Andersen, N. M. 1981. Semiaquatic bugs: phylogeny and classification of the Hebridae (Heteroptera:

Gerromorpha) with revisions of Timasius, Neotimasius and Hyrcanus. Syst. Entomol., 6: 377—

412.

Andersen, N. M. 1982. The semiaquatic bugs (Hemiptera, Gerromorpha). Phylogeny, adaptations,

biogeography and classification. Entomonograph Vol. 3. Scandinavian Science Press, Klam-

penborg, Denmark.

Hurd, P. D. Jr., R. F. Smith & J. W. Durham. 1962. The fossiliferous amber of Chiapas, Mexico.

Ciencias, 21: 107-118.

Petrunkevitch, A. et al. 1971. Studies of fossiliferous amber arthropods of Chiapas, Mexico. Part

II. Univ. Calif. Publ. Entomol., 63.

Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press, Stanford, California.

PAN-PACIFIC ENTOMOLOGIST

71(2): 82-86, (1995)

THE GENUS LIOMETOPUM MAYR

(HYMENOPTERA: FORMICIDAE)

IN CALIFORNIA, WITH NOTES ON NEST

ARCHITECTURE AND STRUCTURAL IMPORTANCE*

HANIF GULMAHAMAD

Terminix International,! Rancho Cucamonga, California 91730

Abstract.—The genus Liometopum Mayr has two species that occur in California. Both species

are structural pests that can build elaborate nesting structures in human habitation. The nest

trabecula of Liometopum occidentale Emery is described and presented here. Two nesting in-

cidences within homes are reported.

Key Words.—Insecta, Hymenoptera, Formicidae. Liometopum occidentale, Liometopum luc-

tuosum, nest structure, structural importance

The genus Liometopum Mayr is comprised of four extinct and five living species.

Two of the living species are from the Old World and three are from western

North America (Cook 1953; Eckert & Mallis 1937; Gregg 1963a; Wheeler &

Wheeler 1973, 1986). Liometopum apiculatum Mayr ranges from Colorado through

Arizona, New Mexico and Texas into Mexico. It is reported from foothill areas

at elevations of 1.2 to 2.1 kilometers (Gregg 1963a, Smith 1979). Liometopum

occidentale Emery, commonly referred to as the velvety tree ant, ranges from

northern Oregon through California into Mexico. In California, this species is

most abundant at lower elevations from sea level to 1.2 kilometers (Cook 1953).

However, Mallis (1941) took this species at 1.8 kilometers elevation on Mt. Baldy,

Los Angeles County, California. I have also seen this species on several occasions

associated with human habitation on wooded lots at Mt. Baldy, California. On

17 Jul 1993, I observed it in association with oak and sycamore trees at Pilgrim

Pines campground, Yucaipa, California. This campground is at an elevation of

approximately 1.5 kilometers.

Liometopum luctuosum W. M. Wheeler was previously considered a subspecies

of L. occidentale. It was elevated to species level by Wheeler & Wheeler (1986).

This species ranges from Wyoming to western Texas thence to Nevada and Cal-

ifornia and into Mexico. It was reported as a montane species occurring at ele-

vations of from 1.2 to 2.4 kilometers (Gregg 1963a; Wheeler & Wheeler 1973,

1986). However, in Tuolumne and Calaveras Counties in California, L. luctuosum

can be found at 0.3 kilometer elevation and higher. It is common at 0.8 kilometer

level in Tuolumne county (J. Tassano, personal communication). I collected this

species once on the north shore of Big Bear Lake, Big Bear, San Bernardino County,

California. I observed this species on 17 Jul 1993 trailing along the base of a low

concrete wall at Pilgrim Pines campground in Yucaipa, California.

* Author page charges partially offset by a grant from the C. P. Alexander Fund, PCES.

19559 Center Avenue, Suite N.

1895 GULMAHAMAD: LIOMETOPUM IN CALIFORNIA 83

ECONOMIC IMPORTANCE

Liometopum occidentale is associated with structures (Bennett et al. 1988, Ebel-

ing 1975, Hedges 1992, Mallis 1990). It is an aggressive and pugnacious species

which feeds on insects and honeydew produced by homopterans. The workers

possess repugnatorial glands from which odoriferous volatile secretions are pro-

duced. The odor resembles that of butyric acid, which most people find disa-

greeable. Liometopum occidentale is commonly found in pronounced columns

going up and down trees such as oak, alder, elm, cottonwood, pine, and sycamore,

where they tend honeydew secreting insects. These ants occasionally invade homes

creating nuisance problems. Homes on properties with trees are more prone to

invasion than those without trees especially when limbs from nearby trees are

touching and/or resting on the structure. On two occasions, I observed these ants

walking on power lines to gain access to homes. Eckert & Mallis (1937) reported

that their trails may extend 60 meters or more from the nest.

Although these ants cannot sting, they are very aggressive and readily swarm

onto any individual that disturbs them. They bite and then spray an irritating

fluid into the wound. Liometopum occidentale is a bane of picnickers in camp-

grounds, parks, and other outdoor recreational areas in southern California (Ebel-

ing 1975).

Pest control operators in California are not familiar with this ant. Because of

its size, its single node, and the concave dorsum of its thorax in profile, it is

commonly mistaken for carpenter ants, especially Camponotus clarithorax Emery

which is about the same size and exhibits similar coloration. Camponotus clari-

thorax is the most common carpenter ant species in the lower elevations of urban

southern California.

I have found infestations of L. occidentale in structures in the cities of Clare-

mont, Glendora, Montclair, Ontario, Pomona, Mt. Baldy, Redlands, and Riv-

erside, in southern California. Most of these infestations were associated with oak

trees, one with pine, and two with sycamores.

LIOMETOPUM LUCTUOSUM

Wheeler (1905) found this species to be closely associated with pine trees. Gregg

(1963a) found that it is not limited to Pinus spp. This is probably valid as ants

of this type are opportunistic species which exploit whatever resources are readily

available to them at a given time. However, observations over many years indicate

that L. luctuosum exhibits a strong preference for pine trees (J. Tassano, personal

communication). It has excavated nests in styrofoam insulation and caused dam-

age to sheetrock in structures (Tassano 1987). In structures, it throws out chewed

up insulation or wood which form piles below the infested area (Tassano 1987).

Wheeler & Wheeler (1986) listed five records of this species in houses in Nevada.

In two of these they were producing piles of fine sawdust; in the third they were

catching insects attracted to lights; and in the fourth they were damaging plas-

terboard.

Liometopum luctuosum is eaten in Mexico. Immature stages of the reproduc-

tives are locally known as “‘escamoles’’. They are regarded as a delicacy and are

widely utilized as food. This ant has been raised and cultured by peasants in

Mexico for many years (Elorduy de Conconi et al. 1986).

84 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Figure 1. Carton nest structure of Liometopum occidentale Emery taken from a nest in a wall void

of a house in Redlands, California.

LIOMETOPUM OCCIDENTALE

Figure 1 shows a portion of the architectural work of L. occidentale taken from

an interstud wall void of an interior partition wall abutting a stall shower of a

home in Redlands, California. The subfloor beneath this stall shower was damaged

by moisture and fungus. Upon examination, it became evident that the fungus

damage extended into the wall abutting the shower. While gaining access to this

wall, a large colony of L. occidentale was found in the lower portions of two

interstud voids. Large numbers of worker ants came out from this area. A re-

pugnant, disagreeable odor immediately became evident. After the ants were killed

with a pyrethrin aerosol, the wall was opened revealing a mass of loose, leaflike,

papery material that is apparently masticated cellulose. This material covered a

spongiform mass of chambers and galleries composed of a carton type material

made by the workers (Fig. 1). Two buckets of nesting materials were removed

from the interstud voids of this house. The interior sides of the drywall were

scarified. No holes going completely through either side of the drywall were ev-

ident. The owner claimed that he was never bothered by these ants indoors.

Around the house were several large oak trees with branches resting on the tile

roof of the house. The property is a typical wooded lot of this area of Redlands,

California and it presented an ideal habitat for L. occidentale.

A similar carton trabecula is presented in Gregg (1963a: 792, plate XXIV) taken

from a nest of L. apiculatum in Colorado Springs, Colorado. Gregg (1963b)

described another nest of L. apiculatum which was taken from the ponderosa

pine-covered foothills near Boulder, Colorado. The trabeculae of this nest were

1995 GULMAHAMAD: LIOMETOPUM IN CALIFORNIA 85

composed of particles of sand and clay cemented together and incorporating

fragments of what appeared to be mica crystals. This nest structure was brittle

and crumbled upon handling. Elorduy de Conconi et al. (1986) described a car-

tonlike interlacing and anastomosing nest structure for L. /uctuosum in Mexico

except that the nests of this species do not always have trabeculae. From these

reports it would appear that all of the New World species of the genus Liometopum

construct elaborate nests. The materials used in constructing a nest depend on

what is available and whether or not the nest is located in the soil or above ground.

Another incident of L. occidentale nesting in a structure occurred at a single

family residential property in Riverside, California. The residents of this property

were experiencing a persistent ant problem. These ants were initially thought to

be carpenter ants and they were treated as such but the problem persisted. On

the second visit, the attic above the bathroom was inspected as the ants were

previously seen there. Upon lifting up the rolled insulation, large numbers of L.

occidentale were found inside and below the insulation. Subsequent remodeling

work in the bathroom entailed removing a large wall mirror which revealed several

holes about 2.5 mm in the drywall. These holes were made by workers which

were found behind the mirror and in the wall void behind the drywall. This wall

was an exterior insulated wall. Upon opening this wall, large numbers of L.

occidentale workers came out. Their typical dolichoderine odor was evident. Hol-

low cavities were found in the insulation for about 1.8 meters of wall void. Large

amounts of pine needles were present in the wall. No particular nest structure

was found in this wall and no trabeculae were evident.

The exterior area adjacent to the infested wall had three pine trees whose

branches were touching the structure thus providing access for the ants to invade

the house. Inspection of the outside areas revealed long trails of L. occidentale

on the wooden fencing bordering the horse trails. I have also seen L. occidentale

on many occasions trailing on wooden fences.

Wheeler (1905) stated that all American species of Liometopum nest in the soil.

Other nesting areas reported and/or suggested in the literature are: in or under

decaying logs, the interior of hollow trees, under rocks, under boulders, in duff,

under wood, under bark, in crevices in trees, under stones, in hollow places in

trees, beneath fallen timbers, etc. (Cook 1953, Gregg 1963a, b, Eckert & Mallis

1937, Mallis 1941, Wheeler & Wheeler 1973, 1986). Many of these nesting areas

are not available to these ants in urban situations. The nesting of these ants in

human habitation is an example of an opportunistic species exploiting a resource

provided by man.

ACKNOWLEDGMENT

I thank Roy R. Snelling of Los Angeles County Natural History Museum for

confirming the determination of L. occidentale.

LITERATURE CITED

Bennett, G. W., J. M. Owens & R. M. Corrigan. 1988. Truman’s scientific guide to pest control

operations. Edgell Communications, Duluth, Minnesota.

Cook, T. W. 1953. The ants of California. Pacific Books, Palo Alto, California.

Ebeling, W. 1975. Urban entomology. Univ. Calif. Div. Agri. Sci., Berkeley, California.

Eckert, J. E. & A. Mallis. 1937. Ants and their control in California. Calif. Agr. Expt. Sta. Circ.,

342: 1-39.

86 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Elorduy de Conconi, J. R., B. Darchen, A. F. Robles, E. S. Castro & S. C. Correa. 1986. Estructura

del nido de Liometopum occidentale var. luctuosum. Manejo y cuidado de estos en los nucleos

rurales de Mexico de las especies productoras de escamoles (L. apiculatum M. y L. occidentale

var. luctuosum W.). (Hymenoptera- Formicidae). An. Inst. Biol; Univ. Nac. Auton. Mex. (Zool.),

57: 333-342.

Gregg, R. E. 1963a. The ants of Colorado. Univ. Colo. Press, Boulder, Colorado.

Gregg, R. E. 1963b. The nest of Liometopum apiculatum Mayr (Hymenoptera: Formicidae). Univ.

Colo. Stud. Biol. Ser., 11: 1-6.

Hedges, S. A. 1992. PCT field guide for the management of structure-infesting ants. Franzak and

Foster Co; Cleveland, Ohio.

Mallis, A. 1941. A list of the ants of California with notes on their habits and distribution. Bull. S.

Cal. Acad. Sci., 40: 61-100.

Mallis, A. 1990. Handbook of pest control. Franzak and Foster Co., Cleveland, Ohio.

Tassano, J. A. 1987. The pine tree ant—a wood destroying ant. Voice of PCOC. Summer 1987.

p. 13.

Smith, D. R. 1979. Superfamily Formicoidea. Tribe Tapinomini. Genus Liometopum Mayr. In

Krombein, K. V., P. D. Hurd, D. R. Smith and B. D. Burks (eds.). Catalog of Hymenoptera

in America north of Mexico. Volume 2. Smithsonian Institution Press, Washington. DC.

Wheeler, W. M. 1905. The north American ants of the genus Liometopum. Bull. Amer. Mus. of

Nat. Hist., 21: 321-333.

Wheeler, G. C. & J. Wheeler. 1973. Ants of deep canyon. Univ. Calif.

Wheeler, G. C. & J. N. Wheeler. 1986. The ants of Nevada. Nat. Hist. Mus. Los Angeles County,

Los Angeles, California.

PAN-PACIFIC ENTOMOLOGIST

71(2): 87-91, (1995)

DACNE PICTA CROTCH: A RECENTLY INTRODUCED

PEST OF STORED, DRIED SHIITTAKE MUSHROOMS

(COLEOPTERA: EROTYLIDAE)

WARREN E. SAVARY

U.S. Food and Drug Administration,! Alameda, California 94502

Abstract.—A major infestation of the dacnine erotylid beetle Dacne picta Crotch was found in

dried shiitake mushrooms (Lentinula edodes (Berkeley) Pegler) stored in a warehouse in San

Francisco, California. The mushrooms were imported from the People’s Republic of China, via

Hong Kong, in June 1990, six months prior to the discovery of the infestation. The spread of

the infestation within the warehouse was documented.

Key Words. —Insecta, Coleoptera, Erotylidae, stored products, introduced pests, mushrooms

In November 1990, investigators from the San Francisco District, U.S. Food

and Drug Administration (FDA), inspected a 400-square-foot warehouse located

on the ground floor of a large, old, concrete-and-wood building in San Francisco,

California. The inspection was conducted to ascertain whether a shipment of 103

cartons of dried shiitake mushrooms (Lentinula edodes (Berkeley) Pegler), which

had been imported from the People’s Republic of China, via Hong Kong, in

October 1990, was present. The mushrooms had been refused entry into commerce

in the United States and were to be re-exported. The inspection revealed that 40

of the 103 cartons were missing and that 40 cartons of previously imported

mushrooms had been substituted for them. This substitution revealed the capacity

of dacnine erotylid beetles to produce major infestations in stored dried mush-

rooms.

The substituted cartons came from two lots of mushrooms imported from Hong

Kong in June 1990. One lot, which originally contained 38 cartons, was repre-

sented in the warehouse by 19 cartons. The other lot, which originally contained

34 cartons, was represented by 21 cartons. Both living and dead beetles were

observed in and on the substituted 19 carton lot and on the warehouse floor (Figs.

1, 2). Neither the other substituted lot nor the remnants of the original lot appeared

to harbor beetles. Samples were collected from the substituted lots and were

examined by the FDA San Francisco District Microanalytical Laboratory. This

examination revealed that the substituted 19 carton lot was infested with the

dacnine erotylid beetle Dacne picta Crotch, which is known to occur in Japan and

mainland Asia (John Lawrence, personal communication). As a result, all 103

cartons in the warehouse were placed under embargo by the California Department

of Food and Agriculture (CDFA), which was cooperating in the investigation.

A follow-up inspection was conducted in December 1990, at which time the

embargoed material was still in the warehouse. Re-inspection revealed that beetles

were still in and on the first substituted lot (Fig. 3) and that the second substituted

1 1431 Harbor Bay Parkway.

88 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Figure 1. Larvae of Dacne picta along seam of carton containing shiitake mushrooms. Carton is

form lot imported from the People’s Republic of China, via Hong Kong, to the United States in June

1990. Photographed in November 1990.

Figure 2. Adults of Dacne picta on floor of warehouse, near cartons containing shiitake mushrooms.

Photographed in November 1990.

1995 SAVARY: DACNE PICTA IN DRIED MUSHROOMS 89

Figure 3. Adults of Dacne picta on corner of carton containing shiitake mushrooms. Carton is

from same lot as carton shown in Figure 1. Photographed in December 1990.

Figure 4. Adults and larvae of Dacne picta on carton containing shiitake mushrooms. Carton is

from substituted lot that appeared to be uninfested in November 1990. Photographed in December

1990.

90 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

lot had become heavily infested with D. picta since the previous inspection (Fig.

4). One beetle was found on a plastic bag in a carton belonging to the original lot,

but none were observed in the enclosed mushrooms. At least 400 live beetles

were observed on the warehouse floor. Both infested lots were voluntarily de-

stroyed by the warehouse owner during inspections. The destructions were wit-

nessed by representatives of the CDFA. The remaining 63 cartons were re-ex-

ported to Hong Kong.

Several aspects of this infestation suggest that D. picta is a potentially serious

pest of stored dried shiitake mushrooms. Dacne picta is able to maintain viable

populations in stored mushrooms, as evidenced by the discovery of the infestation

six months after the affected mushrooms were imported. Presumably, the beetle

arrived in the country along with the mushrooms. The infestation was well-

established and active when discovered, with numerous live adults and larvae

present. Further, D. picta can colonize and infest mushrooms while in storage.

During the month following the initial discovery of the infestation, a second lot

of mushrooms, previously inspected, sampled and found to be pest-free, became

infested. A number of beetles were observed on cartons belonging to a third lot,

and one had entered the lot, but none had penetrated the packaging. The presence

of many beetles on the cartons and warehouse floor suggests that D. picta readily

disperses.

Lawrence (1988) reported D. picta from stored dried mushrooms imported to

Australia from Asia. Dr. Lawrence (personal communication) told me that this

report was based on two separate collections of D. picta from dried shiitake

mushrooms purchased in Canberra. The mushrooms had been imported from

the People’s Republic of China. The first collection was made on 3 Mar 1981 by

R. A. Barret and consisted of about 10 adults and several larvae. The second

collection was made on 30 Jul 1981 by R. Farrow and consisted of seven adults.

The FDA Los Angeles District Microanalytical Laboratory has found D. picta

in dried mushrooms imported to Los Angeles from Korea in four instances. In

three of these, the beetles were in shiitake mushrooms (L. edodes): a single live

adult was collected on 4 Dec 1986 by Michael L. Zimmerman, a single dead adult

was collected on 8 Dec 1986 by James J. Madenjian, and 19 dead adults and 2

live adults were collected on 28 Nov 1988 by James J. Madenjian. In one instance

(10 Nov 1987), two dead D. picta adults were collected from straw mushrooms

(Volvariella volvacea (Bulliard ex Fries) Singer). Recently, Dr. Paul Johnson of

the University of Wisconsin (personal communication) told me of a collection of

three D. picta adults that were found inside a cellophane bag of dried shiitake

mushrooms from Japan. The mushrooms were purchased at an Oriental foods

store in Corvallis, Oregon on 2 Feb 1982.

Dacne picta is not the only member of the genus recorded from stored dried

mushrooms. Two dead D. japonica Crotch adults were collected on 14 Apr 1987

by James J. Madenjian from shiitake mushrooms imported to Los Angeles, Cal-

ifornia, from the People’s Republic of China. The multiple introductions of Dacne

spp. into the United States and elsewhere, particularly the three known intro-

ductions of live D. picta into California, and the demonstrated ability of D. picta

to produce major and persistent infestations in stored dried mushrooms raise

questions as to whether these erotylids will become established pests outside of

their natural range.

1995 SAVARY: DACNE PICTA IN DRIED MUSHROOMS 91

ACKNOWLEDGMENT

FDA Investigators Kathryn D. Macropol and Darla R. Bracy discovered the

San Francisco infestation of D. picta and collected the samples which documented

its spread. Dr. John F. Lawrence of the Commonwealth Scientific and Industrial

Research Organization (CSIRO), Canberra City, Australia, identified the beetles

and provided useful information regarding the Australian records of D. picta. Dr.

Paul J. Johnson of the University of Wisconsin-Madison informed me of the

Oregon record of D. picta and pertinent data. Kathryn D. Macropol, Alan R.

Olsen, and James B. Orr of the United States Food and Drug Adminstration, San

Francisco, California, and James J. Madenjian of the United States Food and

Drug Adminstration, Los Angeles, California, reviewed drafts of this manuscript

and made many helpful suggestions. I thank them all for their contributions.

LITERATURE CITED

Lawrence, John F. 1988. Notes on the classification of some Australian Cucujoidea (Coleoptera). J.

Aust. Entomol. Soc., 37: 53-54.

PAN-PACIFIC ENTOMOLOGIST

71(2): 92-104, (1995)

REGIONAL DIFFERENCES IN THE DISTRIBUTION

OF THE PYRALID MOTH URESIPHITA REVERSALIS

(GUENEE) ON FRENCH BROOM,

AN INTRODUCED WEED

C. B. MontTLior,! E. A. BERNAYS,” J. HAMAI AND M. GRAHAM

Division of Biological Control, University of California,

Berkeley California 94720

Abstract.—The factors influencing the distribution of the moth Uresiphita reversalis (Guenée)

on French broom, Genista monspessulana (L.), its major host plant in California, were inves-

tigated. Within an intensively searched 7000 km? area around the San Francisco Bay, larvae

were abundant on broom in the region east of the Bay, but consistently absent northwest of the

Bay. In field experiments there were no differences in egg mortality, larval development, pupal

weight, predation, plant chemistry, average ambient temperatures or solar radiation between the

two regions which might account for the distribution of the insect. Survivorship of experimentally

placed larvae was dramatically reduced in field sites northwest of the Bay compared to sites east

of the Bay during a rainy winter season. Winter precipitation was consistently twice as high

northwest of the Bay compared to east of the Bay, and large losses of larvae were associated

with rainfall. After three years of drought in California, larvae were found in a site northwest of

the Bay for the first time. It is postulated that heavy rain is a major source of mortality for these

insects, which, along with other unknown factors, may preclude establishment in particular

microclimates.

Key Words. —Insect distribution, Uresiphita, Pyralidae, Genista, Cytisus, precipitation

French broom, Genista (= Cytisus) monspessulana (L.), is a naturalized legu-

minous shrub, native to the Mediterranean, and is well adapted to the climate of

coastal California, where it was introduced about 100 years ago. It has since

become an important weed in parts of the state. Its only major herbivore in

California, Uresiphita reversalis (Guenée) (Lepidoptera: Pyralidae), was first re-

ported in large numbers in California on Genista and “other brooms” (Keifer

1931) in Ventura and San Diego Counties in the southern part of the state.

According to California Department of Food and Agriculture (CDFA) records,

in 1958 the distribution of U. reversalis remained limited to the southern counties.

The earliest records of U. reversalis from northern California are from the early

1980s (J. Powell, pers. comm.). The larvae are aposematic and restricted to a few

genera of the Leguminosae containing quinolizidine alkaloids, e.g., Genista, Lu-

pinus and Laburnum in California (Bernays & Montllor 1989). Larvae are known

to have been deliberately released (S. Bunnell, pers. comm) in the San Francisco

Bay area, but the insect has become established only in selected regions. For

example, in 1986, it was conspicuously rare northwest of the Bay, where the plant

is very abundant and rapidly spreading, in spite of multiple releases dating back

to 1975. In nearby areas across the Bay to the east, however, U. reversalis was

commonly found defoliating French broom, completing several generations per

year on this host (Bernays & Montllor 1989).

! Present address: 4316 La Cresta Ave., Oakland California 94602.

2 Present address: Department of Entomology, University of Arizona, Tucson Arizona 85721.

1995 MONTLLOR ET AL.: DISTRIBUTION OF URESIPHITA REVERSALIS 93

In this study, we have chosen a fairly recent association between an introduced

weed and its primary herbivore to study factors which might account for the

highly disjunct distribution of the herbivore on a relatively small scale (within a

7000 km? area). The factors examined in areas where insects were present or

absent were 1) egg and larval mortality with or without predators excluded, 2)

larval development and pupal weight, 3) weather, specifically temperature, solar

radiation, and precipitation, and 4) plant characteristics, specifically nutrients and

relevant secondary compounds, which are known to influence the feeding of this

species (Montllor et al. 1990).

METHODS AND MATERIALS

Surveys. —Surveys were undertaken around the San Francisco Bay Area during

the summer and fall of 1986-1988, to quantify the apparent differences in dis-

tribution of G. monspessulana and U. reversalis. The insects are multivoltine and

non-diapausing. They build up during the first half of the year and reach peak

abundance in September—October. On visits to different regions, stops were made

on roads and paths, concentrating on sites where there was a possibility of finding

the plant. Overall coverage of representative parts of approximately 7000 km?

around the San Francisco Bay was attempted, excluding dense urban or industrial

regions. Areas of up to 10,000 m2? at each of 90 sites around the Bay area were

thoroughly searched for plants. The plant is most common along roadsides and

is easily identified in transit, but absence in sites was checked by additional

searches. Where the plant was found, thorough searches for insects or insect

damage was made on at least 50 plants/site unless fewer plants were present.

Development of a Natural Population of U. reversalis.—A population of U.

reversalis was followed over a year to better understand its phenology. A 500 m

roadside transect of broom in Oakland, Alameda Co., was surveyed weekly (or

sometimes bimonthly in winter) from 14 Dec 1988 to 8 Feb 1990. Each week the

same 100 bushes, usually 2 m apart, were examined for eggs and larvae, and the

numbers and instars were noted. Plants were approximately 1-2 m high at the

start of the survey.

Releases.—In order to obtain empirical evidence relating to the suitability of

San Francisco Bay area regions for U. reversalis, larvae were released in specific

sites over several years, and sites were subsequently searched for establishment

(Fig. 1). Prior to our experiments, larvae had been released in the Bay Area in

1975 in Alameda Co., east of the Bay (referred to hereafter as EBay) (two sites,

approximately 50 larvae per site) and in Marin Co., northwest of the Bay (hereafter

referred to as NWBay) (two sites, approximately 50 and 1000 larvae) (S. Bunnell,

pers. comm.). In 1985-86, no wild populations of U. reversalis were known from

NWBay sites, while insects were abundant throughout the EBay. Subsequently,

we released larvae in the NWBay at Sky Oaks in September 1985 (100 late instars);

April 1986 (1000-1300 late instars); and July 1986 (400-500 late instars). In

April, June, July, September and November 1988, between 200 and 375 eggs or

first instar larvae were released on each occasion in three NWBay sites as part of

our field experiments (see below). One of these field sites (Olema) was checked

for evidence of larvae monthly from June—October 1989. The other two (Sky

Oaks and Muir Woods) were intensively searched for larvae in October 1989. At

94 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

o®

° ° e

ie) €

oO ° @

7-o ° @

o Bo

40 e fe)

1 Vee

OP | ders

N oe | eee tan Se

psi , )

r a 8

Figure 1. Results of surveys around the San Francisco Bay Area for the presence of Genista

monspessulana and the pyralid folivore Uresiphita reversalis. Solid circles = plant and insect both

present; small dots = both absent; open circles = plant present, insect absent. Experimental (E) and

weather measurement (W) sites in Northwest Bay and East Bay regions: NWBAY 1, Muir Woods (E,

W); NWBAY 2, Sky Oaks (E, W); NWBAY 3, Olema (E); NWBAY 4, Kentfield (W); EBAY 5,

Claremont (E); EBAY 6, LHS (E); EBAY 7, Mt. View (E); EBAY 8, Gill Tract (W); EBAY 9, Berkeley

(E, W); EBAY 10, Oakland Airport (W); EBAY 11, Oakland—natural population development.

each site, an area of approximately 5000 m? around the place of release was

intensively searched.

Field Experiments. —Field sites were chosen in Alameda and Marin Counties,

east (EBay) and northwest (NWBay) of the San Francisco Bay, respectively. Larvae

of U. reversalis had previously been observed feeding on French broom at or near

all the EBay sites, but at none of the NWBay sites; these sites represented areas

where this species was abundant, and rare or absent, respectively (Fig. 1). A total

of four different sites were used over the course of experiments in the EBay, and

three in the NWBay.

Egg Survival.—A possible difference in field survival of eggs in the two regions

1995 MONTLLOR ET AL.: DISTRIBUTION OF URESIPHITA REVERSALIS 95

was experimentaly investigated. Egg masses of U. reversalis, deposited on broom

in a laboratory culture (Bernays & Montllor 1989), were used for examination of

pre-hatching mortality. The eggs were counted and placed in the field by pinning

the leaves on which the eggs had been deposited onto 3-5 plants at each of three

EBay and NWBay sites, on five different occasions. Each plant received one or

more egg masses, consisting of approximately 30 eggs (range = 13 to 68). A total

of 71 and 72 egg masses were placed at NWBay and EBay sites, respectively, over

a year: in June, July and August 1988, and April and June 1989.

Pinned leaves were collected four to five days after eggs were put out, and

examined under a dissecting microsope for signs of non-hatching, hatching, and/

or predation of eggs. Predation was assumed to have occurred when eggs and their

shells were gone from the leaf surface, or when a portion of the leaf containing

eggs had been chewed off. No egg parasites are known for this species in California.

Larval Survival and Development.—To examine possible differences in larval

survival in the two regions, eggs (Experiment 1) or first instars (Experiments 2-

5) from a laboratory culture, were laid or placed on single leaves of broom in

groups of approximately 25 (range = 21-35), and pinned to bushes in the field.

Three to four plants 1-2 m tall at each of three EBay and NWBay sites were

selected. Two groups of neonate larvae, or ready-to-hatch eggs, were placed on

each plant, one enclosed in a white nylon mesh bag to exclude natural enemies,

and the other left uncovered. Positions on plants were representative of those

found naturally. Insects were put out on five occasions, starting in April, June,

July, September, and November 1988. Larvae were counted at intervals until all

insects in bags had pupated and all larvae in the open had disappeared (larvae

leave the plant to pupate). Because large larvae will move to adjacent branches

on the same or a neighboring plant to feed, isolated bushes were used if possible.

Otherwise, the neighboring bushes were always searched for missing larvae. Sur-

vivorship in the open and in bags was estimated by counting remaining larvae

twice during the experiments. For Experiment 5 (November) no larvae were

enclosed in bags. Pupae from bags from Experiments 3 and 4 were counted and

weighed in the laboratory.

Because late fall and winter mortality appeared to be important, additional

simplified experiments were carried out in the winter of 1989-90 at one EBay

and one NWBay site. In these, 10 to 15 second instar larvae were placed on each

of 7-10 plants per site in November (Experiment 6, starting n = 110 larvae) and

in January (Experiment 7, starting n = 100), and remaining larvae were counted

weekly.

Two measurements of development were made during four of the experiments

in which larvae were placed in the field. The distribution of instars in bagged and

unbagged (open) cohorts at all sites at a given sampling date was determined.

Sampling dates ranged from 13 to 24 days after the beginning of a given exper-

iment. Sampling of all sites was always accomplished within one or two days.

Plant Chemistry. —The possibility that regional differences (genetic or environ-

mental) in plant chemistry might affect insect abundance was also examined. Plant

material was collected from experimental plants at all sites during the second

(June 1988) larval survival experiment.

Material was collected twice, early and late in the experiment, dried in an oven

at 70° C, and analyzed for total alkaloids (Montllor et al. 1990), Kjeldhal nitrogen,

96 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

and soluble carbohydrates (anthrone method). Nitrogen and carbohydrates were

selected as the major important nutrients. Quinolizidine alkaloids were selected

as the characteristic secondary components of the host plant. These are phago-

stimulants for U. reversalis larvae, and are sequestered by them (Montllor et al.

1990). We have also found that the sequestered alkaloids are effective deterrents

to hymenopteran predators of these larvae (Montllor et al. 1991).

Weather. — Because there appear to be marked regional differences in the weath-

er around the San Francisco Bay Area, the possibility that climatic factors may

be correlated with insect abundance was examined. Temperatures, precipitation

and solar radiation were collected with a LICOR datalogger, hygrothermographs,

and/or from National Oceanographic and Atmospheric Administration (NOAA)

records. Data were collected for three EBay sites and three NWBay sites, at or

near the field sites (see Fig. 1). NWBay weather sites were Muir Woods (at a field

site), Sky Oaks (at a field site), and Kentfield (4.5 km from Sky Oaks field site).

EBay sites were Berkeley (1-2 km from LHS, Claremont, Dwight-Derby field

sites), Gill Tract (5-8 km from all sites excluding Mt. View), and Oakland (4.5

km from Mt. View field site).

Laboratory experiments. — Daily temperatures fluctuated more widely in NWBay

sites than in EBay sites, so we examined the effects of temperature fluctuations

on larval mortality. Some larval Lepidoptera apparently survive better under

fluctuating, rather than constant, temperatures in the laboratory (McDonald 1990).

However, infectious agents may be more virulent at alternating than at constant

temperatures, e.g., Bacillus popillae in certain scarab beetles (Milner et al. 1980).

Insects were reared at two temperature regimes in environmental chambers in

the laboratory, in which the mean temperature was the same, but one fluctuated

and the other did not. Five groups of 10 first or second instar larvae were placed

on cut broom at temperatures of either 32°:8° C (12L:12D) or at 21° C (+19) (12L:

12D). Food was changed and larvae were counted every other day until larvae

began to pupate. This experiment was replicated three times for a total of 150

larvae per treatment.

RESULTS

Surveys. —General surveys for G. monspessulana around the San Francisco Bay

area demonstrated that the plant was abundant northwest of the Bay, and was

also common east of the Bay. Fewer plants occured on the San Francisco peninsula,

and south of San Francisco natural vegetation predominated, with less apparent

invasion of this weed (Fig. 1). Populations of U. reversalis were common on broom

east of the Bay but mostly absent on the west side of the Bay. Most sampling was

done at times when insect populations would be at or near their peaks. The scarcity

of this insect northwest of the Bay, and its absence on the northwest coast, where

broom populations were large and spreading, was striking.

In a subsequent survey of NWBay field sites, larvae of U. reversalis were found

at Sky Oaks in October 1989. Searches of Olema and Muir Woods, our two other

NWBay sites found no U. reversalis in 1989. The last releases at all these sites

had been in November 1988 (Experiment 5), therefore populations apparently

successfully overwintered at Sky Oaks in 1988-89 for the first time.

Development of a Natural Population of U. reversalis.—In 1989, at the Oakland

(Alameda Co., EBay) site selected to follow a natural population, larvae were

1995 MONTLLOR ET AL.: DISTRIBUTION OF URESIPHITA REVERSALIS 97

No. Larvae

100

1989-1990

1/3 4/10 7/19 10/27 1/3

Figure 2. Number of second instar larvae of U. reversalis on 100 plants of G. monspessulana (site

11) at given sample dates over an entire year (1989).

found in all months, but numbers were very low in January through March. Over

four years, adults were caught at light traps in neighboring Berkeley in all months

except January (J. Powell, pers. comm.), and the first new generation of caterpillars

appeared at our site in April. Numbers built up slowly through the summer,

reaching a peak in October.

Numbers of second instars peaked at intervals of 47 to 77 days, beginning in

May (Fig. 2) (eggs and first instars are inconspicuous, therefore their presence/

absence is not reliably recorded). The first peak occurred on 5 May, a second 51

days later (23 Jun), another 47 days later (10 Aug), and the last and biggest 77

days later (26 Oct). Itis not certain that each new peak represents a new generation,

since in laboratory studies, insects reared at an average temperature of 17° C (a

realistic temperature for this site in spring/summer; see Table 3) took a mean of

76 days to develop from egg to adult (Montllor et al., unpub.). However, depending

on insolation these caterpillars may increase their body temperature by 3° C or

more at this air temperature (Bernays & Barbehenn, unpublished data). Adult

emergence in the spring may be somewhat staggered depending on pupation

microsites, but it is likely that there are 3-4 generations during the year.

Field Experiments. —Field experiments, in which eggs or young larvae were

placed at selected sites, covered the period from April through November, months

during which young larvae were found in our natural field population (Fig. 2).

Instances of egg predation were more frequent in NWBay sites on four out of

five occasions. A total of 34% of the egg masses showed some evidence of predation

in NWBay sites (n = 71), compared to 19% in EBay sites (n = 72), but in spite

98 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Table 1. Survivorship of larvae when placed as neonates (or eggs) on open branches of broom in

EBay and NWBay sites (1988). Starting numbers of larvae. were between 200 and 375 for each

experiment, placed on 3—5 plants at each of three sites in each region.

% survivorship

Exp. (start) Days from start Instar “EBay ~=NWBay— P*

1 (April) 11-12 2-3 36 42 >0.1

22-23 3-5 10 12 >0.4

2 (June) 7-8 2-3 50 40 <0.03

17-19 2-5 21 20 >0.8

3 (July) 8 3-4 37 42 >0.1

23 3-5 24 21 >0.5

4 (Sept.) 12-13 2-3 20 21 >0.8

21-22 3-4 9 10 >0.7

5 (Nov.) Liste 2 35 20 <0.001

20-21 2 23 12 <0.001

* Numbers of living and dead (or missing) larvae compared between regions by Chi-square test.

of this trend, there was considerable variation and the difference between regions

was not significant (P > 0.05, G-test). There were no differences in hatching rate,

and no parasitism.

Numbers of larvae remaining on the bush on which they had been originally

placed (or on an adjacent bush) was used as our measure of larval survivorship.

Survival of larvae up to approximately the fourth instar on open (unbagged)

branches of broom was significantly greater in EBay sites in two of the five

experiments; in the experiment conducted during the 1988-89 winter rainy season

this difference was large and sustained (Table 1). However, during the winter of

1989-90, no difference in survivorship of larvae put out as second instars (Ex-

periments 6 and 7) was found between a NWBay site (Sky Oaks) and an EBay

site (Mt. View). This may be, in part, because little rain fell in either area during

those experiments (see Weather below).

The number of insects surviving to pupate on bagged branches (1.e., predators

excluded) was significantly greater in EBay sites in two of the four experiments

(June and July) in which larvae were bagged (P < 0.05, Mann-Whitney U-tests);

there was no significant difference in the other two experiments. Overall, the mean

survival to pupation was 28% in EBay sites compared to 17% in the NWBay sites

(P < 0.03, ANOVA) over the four experiments. Survival in bags was usually

lower in NWBay compared to EBay sites, though the pattern over the four field

experiments was inconsistent (Fig. 3). The greater mortality in bags at NWBay

sites is evidently responsible for the difference in survival to the pupal stage (only

measured in bags) between regions. The reason for this difference is not known.

Survival of larvae in bags was greatly increased compared to that of unprotected

larvae in both areas. In NWBay sites, survival in the open was 56% of that in

bags; in EBay sites survival in the open was 48% of that in bags. Although larvae

of U. reversalis are not preferred by many arthropod predators in laboratory studies

(Montllor et al. 1991, and unpublished), they are subject to predation by Hemip-

tera (Anthocoridae) and Hymenoptera (Vespidae) in the field (Bernays & Montllor

1989). However, predation pressure, measured as the difference between survival

1995 MONTLLOR ET AL.: DISTRIBUTION OF URESIPHITA REVERSALIS 99

% Survival

EBay a said NWBay

ZZ ILL

A]AM@aaZ7”Z7”Z,

Lzz77-z*zzztttttttbb

Lz ob&&&pzb&xApzg

L777

AMZZZZ77

AZZ

AMAaAaAAH?B#

E =F E. L E, LL, JF

April May July Sept

Figure 3. Percent survival of U. reversalis cohorts bagged on G. monspessulana in four experiments

in 1988 at EBay and NWBay sites (pooled). Insects were censused three times: E = early (instars 2-

3); L = late (instars 3-5); P = pupae. *Numbers of pupae significantly different between regions (P <

0.05, Mann-Whitney U-test); ns, not significant.

on open compared to bagged branches, was not demonstrably different between

the two areas under study here.

Development. — Distributions of larval instars were significantly different, but

inconsistently so, in EBay sites compared to NWBay sites in all experiments.

Larvae put out in April and June (Experiments 1 and 2) developed more quickly

in the EBay sites. Although a greater proportion of larvae put out in July (Ex-

periment 3) had reached the fourth instar after two weeks in NWBay compared

to EBay sites, there were also more early instars in NWBay sites (on open branches,

11% were still second instars, compared to none in EBay sites). In contrast to the

earlier season results, larvae put out in September were ahead in NWBay sites

after 22 days (Fig. 4). Age structure of larvae in bags and in the open showed

similar differences, indicating that where differences occur between regions, they

are not likely to be due to differential loss of early instars to predation, or late

instars to disease. In sum, during our experiments, U. reversalis larvae appeared

to have an early season advantage in EBay sites, and a late season advantage in

NWBay sites, with no net overall effect of region on development rates.

In two experiments, fresh weights of pupae collected in the two regions were

compared. In Experiment 3 the mean weights of pupae in EBay sites compared

to NWBay sites were 87.8 and 92.5 mg, respectively (P > 0.6, nested ANOVA).

Similarly, in Experiment 4 mean pupal weights were 91.1 and 98.4 mg in EBay

compared to NWBay sites (P > 0.6, nested ANOVA). In both experiments, there

100 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

% in Instar

Apr-24 d Jun-17 d Jul-17 d Sep-22 d

ZEEE EEE

‘

]a]acp#™

AZZ

CA AZZ

ié¥)

nstar

ié¥)

=— Ol

MXM East Bay MH NWest Bay

Figure 4. Percent of larvae which were given instars on open branches during four experiments,

beginning in given months. Sampling dates were given number of days (d) after putting out 1st instar

larvae (or eggs, in April). All distributions were significantly different between regions (P < .001, Chi-

square).

was significant variation in pupal weights between sites within regions but not

between regions.

Plant Chemistry.—No differences in soluble carbohydates, total nitrogen, or

total quinolizidine alkaloids were found between plants at the NWBay and EBay

sites (Table 2).

Weather. — Rainfall was consistently higher in the NWBay in 1985-1989 (Fig.

5). Data collected from other sites and from NOAA corroborates this difference

both during years of drought (1986-91) and of normal rainfall. NOAA reports

“normal rainfall’? to be more than twice as great in Kentfield (NWBay) as in

Berkeley (EBay) in a typical November, December, January and February. For

Table 2. Mean levels (+ SD) of carbohydrates (CHO), Kjeldahl nitrogen (N) and quinolizidine

alkaloids (QA) in broom samples (% DW).

Region

(no. plants) CHO N QA

EBay 8.4 + 2.3 ice US 0.4 + 0.2

(12)

NWBay 6.7 + 0.8 2i2-2 OS 0.3 + 0.3

(8-9)

P >0.05 >0.05 >0.05

(t-test)

1995 MONTLLOR ET AL.: DISTRIBUTION OF URESIPHITA REVERSALIS 101

Precipitation (mm)

Berkeley(EBay) ME sky Oaks(NWBay)

No &

2 CAA Z_—_—_ZZZ|.

O CXqZL--LLZD

OO CXL

Aaae'—C "$7,

ws _—|—A/|]Myj;]e—0nqn7

\ Y \

\ \ \ \ \

D N D N N J

85-86 86-87 87-88 88-89

Figure 5. Precipitation during November, December and January in four years at Berkeley (EBay)

and Sky Oaks (NWBay).

example, total “normal rainfall’? at these locations for these four months are

reported as 85 mm and 40 mm, respectively.

Total numbers of larvae at our monitored natural field site in the EBay fell

dramatically at the onset of the late fall rains. The relationship between larval

mortality and rainfall was examined further by regressing larval losses against

precipitation during the first two weeks of Experiments 5, 6 and 7, conducted

during the 1988-89 and 1989-90 rainy seasons. This regression showed a signif-

icant correlation (r = 0.67, P < 0.04).

Mean daily temperatures were not significantly different in the two regions in

any of the four months sampled in 1988 (Table 3). Although mean daily fluctu-

ations in temperature appeared to be slightly larger in NWBay sites, the differences

only approached significance in one month (July; P = 0.06).

Solar radiation was compared at Sky Oaks (NWBay) and Berkeley (EBay) over

72 days in May through August 1988. All days for which daily totals for both

sites were available were used in the sample. Summer fog can be very common

in these areas, but may differ significantly on a microgeographic scale. We found

mean total daily radiation at the two sites to be very similar: 7158 Wm~? at Sky

Oaks and 7087 Wm? at Berkeley (P > 0.3, Wilcoxon rank sum).

Laboratory Experiments: Temperature Fluctuation.—When the three experi-

ments (each with 50 insects per treatment) were pooled, mortality was not sig-

nificantly higher for larvae reared in an environment in which temperatures fluc-

tuated widely than at a constant temperature. At 8-32° C, 23% of the larvae died,

whereas 15% died when reared at 21—22° C (Chi-square, P > 0.1).

102 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Table 3. Temperature measurements (°C, mean + SD) from three EBay and three NWBay sites

during four months in 1988.

May Jul Sep Dec

A. Monthly means of daily median temperatures (high + low/2)

EB 14.7 + 3.0 1$.2 £.2.6 16.6 + 1.8 9.5 + 3.7

NWB 14.6 + 3.6 19.9 + 3.5 s+ 21 9.8 + 3.9

Pe 0.78 0.39 0.57 0.71

B. Monthly means of daily fluctuation (high-low)

EB 10.5 + 4.7 11.2 + 4.3 11.1 + 5.2 8.3 + 3.4

NWB 13.9 + 5.1 17.1 + 4.9 15.6 + 5.5 8.8 + 3.7

cea 0.23 0.06 0.31 0.78

* Nested ANOVA.

DISCUSSION

Our field experiments showed minimal differences in egg mortality, larval de-

velopment, pupal weight, predation, plant chemistry, ambient temperatures or

solar radiation between the regions where U. reversalis was abundant or absent.

Early winter mortality was significantly higher for larvae in NWBay sites, and

other evidence also suggests that winter precipitaion may be at least one important

factor in limiting the distribution of this insect. Other sources of mortality certainly

exist, but most remain unexplained (e.g., differences between EBay and NWBay

mortality in May and July 1988).

Moran & Whitham (1988) have suggested that weather may have a large and

direct effect on insect populations on which predation and parasitism are relatively

low. Populations of a leaf mining sawfly, Schizocerella pilicornis (Holmgren), are

rarely attacked by natural enemies and appear to be limited in distribution in

northern California by climatic factors such as temperature and solar radiation

(Wrubel 1990). Although predation may account for some losses of U. reversalis

larvae in summer months (Bernays & Montllor 1989), we did not find differences

in predation between areas where U. reversalis was common or absent. Larvae

are probably protected from many vertebrate and invertebrate predators by virtue

of the alkaloids which they sequester from Genista (Montllor et al. 1990, 1991).

In our experience with field collected larvae, parasitism is also relatively rare,

with the exception of a tachinid pupal parasite. Therefore there is a potential for

large direct or indirect effects of weather on populations of this insect.

Although we have not found differences in temperature or solar radiation in

our two regions under study, precipitation, which is much greater northwest of

the Bay than east, may play an important role in larval and pupal mortality. The

major causes of pupal mortality are not known. However, large losses of larvae

were clearly related to rainfall during our experiments and in the natural popu-

lation of U. reversalis which we sampled regularly for over a year. Observations

of other wild populations of U. reversalis in the East Bay showed that on four

occasions during 1985-86, population losses of 80% or more were recorded in

association with heavy rains. In one case, 14 larvae were counted the day before

a heavy storm; the following day, 2 larvae remained on the same bushes (Bernays,

unpublished).

1995 MONTLLOR ET AL.: DISTRIBUTION OF URESIPHITA REVERSALIS 103

Although U. reversalis larvae were released at Sky Oaks in 1985-1988, only in

the fall of 1989 were new generations found there; in previous years, insects

presumably failed to overwinter, as new larvae were never found. That winter

(1988-89) was the third of below-normal rainfall in northern California. No larvae

were found in fall 1989 at the other NWBay sites where field experiments had

been carried out the previous two years.

The probability of establishment of a field population even under the best

environmental circumstances is not known. Probability of establishment may be

quite difficult to measure accurately (Harrison 1989) and experiments were not

carried out which might have addressed this issue. However, one of the EBay

sites, LHS, did not have an endemic population of U. reversalis when we began

our experiments in 1986, but by the next year, there were large populations there,

whereas there were none in NWBay sites after introductions of similar numbers

of larvae. This gives some evidence that the numbers of larvae put out in 1986

were large enough to allow for establishment under the proper conditions, although

we cannot rule out the possibility that LHS populations were at least partially the

result of migrants from nearby areas.

Weather may also affect mortality of larval Lepidoptera indirectly through

incidence of disease or of predation/parasitism. Both rainfall and high temperature

maxima have been implicated in increased mortality from granulosis virus of

Pieris rapae L. (Lepidoptera) larvae (Harcourt 1966). Bacillus popillae was shown

to be more infectious in certain scarab beetles under fluctuating compared to

constant temperatures (Milner et al. 1980).

Although the incidence of disease was not quantified in this study, many flaccid

dead larvae were found at all sites in the first three experiments. Such dead larvae

collected on 13 Jul 1988 at two EBay sites contained granulosis virus and Bacillus

cereus Frankland & Frankland, both common disease agents of larval Lepidoptera.

We were unable to significantly increase larval mortality in the laboratory by

fluctuating rearing temperature. In a recent review, Benz (1987) discusses the

problem of environmental effects on disease outbreaks. There are no general

patterns, and even with baculoviruses high temperatures or low temperatures can

either induce expression of particular diseases or suppress them.

It appears that we have witnessed a persistent decimation of populations of U.

reversalis in winter, and subsequent populations which are too small to persist,

in a particular region of northern California. Four years of drought, coupled with

repeated introductions (the last in January 1990) may have finally allowed pop-

ulations to take a foothold northwest of the San Francisco Bay. A large population

was found at Sky Oaks in October 1990. In the spring of 1990, larvae were also

found in Ross, Marin Co., 5 km from our nearest field site (Sky Oaks). Whether

populations will persist when winter precipitation levels return to normal has yet

to be determined.

The possibility that such small-scale weather patterns might dictate this insect’s

distribution is plausible, and is analogous to altitudinal restrictions, which may

be very sharply delimited. Conditions for adult survival in winter, and factors

affecting female fecundity in spring are not known for this species. Price et al.

(1990) have recently pointed out the importance of natality on insect population

dynamics. Although we have emphasized larval mortality factors in our study,

104 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

we recognize the potential importance of factors affecting successful adult eclosion,

mating and oviposition, which we have not measured here.

ACKNOWLEDGMENT

Thanks to J. A. Powell for discussion and unpublished data, to R. V. Barbehenn

for technical support, to G. M. Thomas for identification of larval pathogens, to

the Golden Gate National Recreation Area, to Muir Woods National Forest, and

to Casey May and the Marin Municipal Water District for cooperation throughout

this study. This work was supported in part by USDA grants 87-CRSR-2-3113

and 88-38300-3628.

LITERATURE CITED

Bernays, E. A. & C. B. Montllor. 1989. Aposematism of Uresiphita reversalis larvae (Pyralidae). J.

Lep. Soc., 43: 261-273.

Benz, G. 1987. Environment. pp. 177-214. In Fuxa, J. R. & Y. Tanada (eds.). EEO of insect

diseases. John Wiley & Sons, New York.

Harcourt, D. G. 1966. Major factors in survival of the immature stages of Pieris rapae (L.). Can.

Entomol. 98: 653-662.

Harrison, S. 1989. Long distance dispersal and colonization in the Bay checkerspot butterfly, Eu-

phydryas editha bayensis. Ecology, 70: 1236-1243.

Keifer, H. H. 1931. Notes on some California Lepidoptera of economic interest. Monthly Bulletin,

Calif. Dept. Agric., 20: 613-626.

McDonald, G. 1990. Simulation model for the phenological development of Mythimna convecta

(Walker) (Lepidoptera: Noctuidae). Aust. J. Zool., 48: 649-663.

Milner, R. J., J.T. Wood & E. R. Williams. 1980. The development of milky disease under laboratory

and field temperature regimes. J. Invert. Path., 36: 203-210.

Moran, N. A. & T. G. Whitham. 1988. Population fluctuations in complex life cycles: an example

from Pemphigus aphids. Ecology, 69: 1214-1218.

Montllor, C. B., E. A. Bernays & R. V. Barbehenn. 1990. Importance of quinolizidine alkaloids in

the relationship between larvae of Uresiphita reversalis (Lepidoptera: Pyralidae) and a host

plant, Genista monspessulana. J. Chem. Ecol., 16: 1853-1865.

Montllor, C. B., E. A. Bernays & M. L. Cornelius. 1991. Responses of two hymenopteran predators

to surface chemistry of their prey: significance for an alkaloid-sequestering caterpillar. J. Chem.

Ecol., 17: 391-399.

Price, P. W., N. Cobb, T. P. Craig, G. Wilson Fernandes, J. K. Itami, S. Mopper & R. W. Preszler.

1990. Insect herbivore population dynamics on trees and shrubs: new approaches relevant to

latent and eruptive species and life table development. pp. 1-38. Jn Bernays, E. A. (ed.). Insect-

plant interactions, Vol. II. CRC Press, Boca Raton.

Wrubel, R. P. 1990. Studies on discrimination in choice of oviposition site and the influence of

climate on the distribution and abundance of Schizocerella pilicornis Holmgren (Hymenoptera:

Argidae) in California. Ph.D. Dissertation, University of California, Berkeley.

PAN-PACIFIC ENTOMOLOGIST

71(2): 105-109, (1995)

THE DESERT DAMPWOOD TERMITE

(ISOPTERA: KALOTERMITIDAE) AS A

STRUCTURAL PEST IN THE COLORADO

DESERT OF SOUTHERN CALIFORNIA*

HANIF GULMAHAMAD

Terminix International,! Rancho Cucamonga, California 91730

Abstract.—The desert dampwood termite, Paraneotermes simplicicornis (Banks) is found in the

Colorado and Mojave deserts of California. This termite is often found associated with dead,

dying, or living vegetation within its geographic range. It was not considered a pest of structures

in the past. I report an incident where this termite caused significant damage to a home in

Cathedral City, California. With urbanization increasingly encroaching on both the high and

low deserts of California, the desert dampwood termite can be expected to be found with

increasing frequency attacking structures when the right conditions exist for its survival.

Key Words. —Insecta, Isoptera, Kalotermitidae, Paraneotermes simplicicornis, structural pest

The desert dampwood termite, Paraneotermes simplicicornis (Banks) was first

described in 1920 from specimens taken in 1917 at Laguna, Uvalde County,

Texas (Banks & Snyder 1920). The first California record of this species was in

1926 when specimens were taken near Indio, Riverside County, California (Light

1937). The known range of this species is from the 99th meridian to the 117th

meridian. The east—west territory traverses about 544 kilometers. The north—

south range extends from the 36th parallel (Las Vegas, Nevada) to the 26th parallel

(San Blas de Sinaloa, Mexico). This distance is approximately 380 kilometers

(Light 1937). In California, P. simplicicornis is confined to the Mohave and Col-

orado deserts where its westernmost records are a few kilometers east of Barstow

and a few kilometers east of Cabazon in the lower reaches of San Gorgonio Pass

respectively (Light 1937).

The geographic territory inhabited by this termite is arid or semi arid with

sparse xerophytic vegetation. In these areas, the desert dampwood termite is found

associated with a number of dead, dying, or living native plants (Light 1934,

1937).

Light (1937) reported that although this termite is found in xerophytic situations

it has a high moisture requirement. It obtains this moisture either by a subter-

ranean mode of existence, essentially confining its attacks to wood in or on the

ground, or by feeding on living plants. The desert dampwood termite has been

reported to attack living hedge plants, young citrus trees, and grape vines (Light

1934, 1937; Ebeling 1975; Weesner 1965, 1970). It has caused damage to untreated

poles and posts within its range (Light 1934).

In addressing the question as to whether or not the desert dampwood termite

is able to attack sound wood, Light (1937) found that: (1) Foragers are unable to

attack sound wood except when such wood is in or on the ground or when they

* Author page charges partially offset by a grant from the C. P. Alexander Fund, PCES.

19559 Center Avenue, Suite N.

106 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

have access to decayed wood, and (2) Even when foragers have moist soil con-

nections, they are unable to survive for long except on decayed wood. Both Light

(1931) and Nutting (1966) found that foragers of this termite present in surface

wood are the work force of a colony found elsewhere in the soil.

This paper reports on one case where this termite was found attacking sound,

dry, structural wood which was over five meters away from the soil.

CASE HISTORY

On 28 Apr 1993, an inspection was made of a single family, residential, slab-

on-grade constructed building located in Cathedral City, California. The owner

had called complaining of droppings in a cabinet which is against an exterior

perimeter wall. This cabinet abuts the ceiling of the house above the washer and

dryer which is adjacent to the kitchen area. Inspection of the upper areas of the

cabinet revealed what appeared to be a mud tube about 5.5 centimeters long on

the drywall extending downwards from the ceiling. The bottom end of this mud

tube was open and pellets were being pushed out from it. These pellets were

scattered on the upper shelf of the cabinet. Closer examination using a hand lens

revealed that what appeared to be a mud tube was really a similar type of con-

struction made entirely of pellets which were glued together. This evidence sug-

gested that the attic area above should be inspected to determine the source of

this infestation.

The attic inspection revealed substantial damage to several ceiling joists. A

total of about ten to fifteen ceiling joists exhibited some damage. Vertical and

horizontal pellet tubes of varying lengths were found on the ceiling joists. Many

of the vertical pellet tubes had rings of pellets around them which had been expelled

by the termites. The expelled pellets were loose, friable, and dark brown in color.

The damage caused by this termite in construction grade lumber is similar to that

caused by subterranean termites in that the tunnels followed the grain of the wood.

Pellets were readily evident in these tunnels. Most of these pellets were glued

together and thus were not as loose and friable as those produced by termites of

the genus Jncisitermes. The joists were eaten on the inside but the termites seldom

broke through to the exterior. They left a thin layer of wood on the outside which

offered them protection from the elements and their natural enemies. When the

rolled insulation between the damaged ceiling joists was lifted, damage to the

paper backing of the insulation was evident in several areas. Piles of pellets were

also found adjacent to. the ceiling joists near the insulation.

No moisture problem could be found in this area of the attic. There was no

evidence of plumbing or roof leaks in the entire attic. The damaged ceiling joists

were completely dry wood. In this area of the eastern Colorado desert, it is common

to find daytime attic air temperatures ranging from 50° C to 55° C or more during

the hot summer months.

Probing of the infested joists revealed all stages of nymphs and several soldiers.

The damage to the ceiling joists was so severe that the termite inspector expressed

great reluctance to step on them because he feared that he might fall through the

ceiling. Portions of some of these ceiling joists were so damaged that large chunks

of wood could be removed with one’s bare hands. Figure 1 shows three pieces of

wood which were taken from damaged ceiling joists of the infested property at

Cathedral City. What appeared to be mud tubes in these samples are really pellets

1995 GULMAHAMAD: DESERT DAMPWOOD TERMITE IN S. CALIFORNIA 107

Figure 1. Portions of ceiling joists showing desert dampwood termite damage with fecal pellets

glued together in the tunnels.

which are glued together. The termites confined their attack to the ceiling joists

of this property and they did not go any higher in the attic. An interior inspection

of the living areas revealed no clues as to the origin of this attic infestation of the

desert dampwood termite. However, a leaky spigot was found on the exterior of

the home about 1.2 meters from where the pellets were found in the cabinet. This

water leak can best be described as a fast drip which created a small, standing

puddle of water adjacent to the exterior foundation of the structure. This water

spigot had been leaking for some time because it sustained a growth of lush, green

vegetation in the immediate area.

The exterior walls of this structure were covered with stucco which extended

below the exterior grade level. It appears that P. simplicicornis was able to bridge

the gap between the soil and wood members in the wall by constructing pellet

tubes behind the stucco. Although a recommendation was made to open the

exterior wall adjacent to the interior infested area for further inspection, the

homeowner was not receptive to this suggestion. Based on the extent of the damage

found in the ceiling joists, it is my belief that possible hidden damage exists in

the adjacent wall.

SUMMARY AND CONCLUSIONS

I have found the desert dampwood termite on two other occasions attacking

structures in the Palm Desert and Cathedral City areas. On one of these occasions,

damage was done to a window sill and framing. Swarmers emerged inside the

108 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

house adjacent to the damaged area and they alerted the homeowner to the

problem. On another occasion, this termite was found causing damage to a door

frame and its molding. I had earlier ignored these incidences attributing them to

stochastic factors. I have also received specimens of the desert dampwood termite

from Las Vegas, Nevada where they were reported as having been found damaging

construction grade lumber buried in the ground at the base of a mobile home

skirting.

All of these records suggest that this termite is now being found more often in

structures. In 1934 and 1937, the desert dampwood termite essentially confined

its attack to wood below ground (Light 1934, 1937). Weesner (1965) referring to

this termite stated that: ““These termites apparently have somewhat higher mois-

ture requirements than many species of the Kalotermitidae and are known to

damage living trees. This latter habit is probably the chief cause for economic

concern.”’ Ebeling (1975) also showed the lack of structural importance of this

species when he devoted only two sentences to it in his book on urban entomology.

Substantial urbanization of portions of the Colorado and Mojave deserts have

caused large areas of land to be cleared of native vegetation. This is depriving

endemic termites of their natural food sources thus forcing them to seek food in

and around human habitation. Increased urbanization in the deserts of California

is usually accompanied by ample irrigation of the landscape. Former city dwellers,

who are now encroaching on the deserts for economic and other reasons, are

determined to create an aesthetically pleasing greenery to enhance the quality of

their surroundings. In doing so, they are creating conditions which are conducive

to the proliferation of the desert dampwood termite. As a result, this termite will

likely become a pest of human habitation more frequently in the future.

Pest control operators (PCOs) within the geographic range of the desert damp-

wood termite are not familiar with this species. Many of them simply call this

termite a subterranean termite. PCOs might succeed in controlling the desert

dampwood termite using the same techniques, methods, and materials used in

controlling subterranean termites because P. simplicicornis is a drywood termite

with subterranean habits. I believe that the current common name of P. simpli-

cicornis (the desert dampwood termite) is a misnomer. It could more aptly be

described as the drywood subterranean termite.

At this time, I am not sure that the infestation of the desert dampwood termite

described in this paper can be controlled by simply applying a termiticide to the

soil. If this property is ever treated to control this infestation, it would be inter-

esting to cut these termites off at soil level with a termiticide soil barrier and then

monitor their survival in the ceiling joists.

LITERATURE CITED

Banks, N. & T. E. Snyder. 1920. A revision of the Nearctic termites (Banks) with notes on biology

and geographic distribution (Snyder). U.S. Nat. Mus. Bull., 108: 1-228.

Ebeling, W. 1975. Urban entomology. University of California Press, Berkeley, California.

Light, S. F. 1931. The termites of Nevada. Pan-Pac. Entomol, 8: 5-9.

Light, S. F. 1934. The desert dampwood termite. Paraneotermes simplicicornis. pp. 311-313. In

Kofoid, C. A. (ed.), Termites and termite control. (2nd ed). University of California Press,

Berkeley, California.

Light, S. F. 1937. Contributions to the biology and taxonomy of Kalotermes (Paraneotermes) sim-

plicicornis Banks (Isoptera). Univ. of California Publ. Ent., 6: 423-464.

1995 GULMAHAMAD: DESERT DAMPWOOD TERMITE IN S. CALIFORNIA 109

Nutting, W. L. 1966. Colonizing flights and associated activities of termites. I. The desert damp-

wood termite Paraneotermes simplicicornis (Kalotermitidae). Psyche, 73: 131-149.

Weesner, F. M. 1965. The termites of the United States: A handbook. National Pest Control As-

sociation. New Jersey.

Weesner, F.M. 1970. Termites of the Nearctic Region. pp. 477-525. In Krishna K. & F. M. Weesner

(eds.), Biology of termites (Vol. 2). Academic Press, New York.

PAN-PACIFIC ENTOMOLOGIST

71(2): 110-112, (1995)

AN ATYPICAL NEW SPECIES OF CORTICARINA

FROM THE CLARION ISLANDS, MEXICO

(COLEOPTERA: LATHRIDITDAE: CORTICARIINI)

FRED G. ANDREWS

Insect Biosystematics, Plant Pest Diagnostics Center,!

California Department of Food and Agriculture,

Sacramento, California 95832-1448

Abstract. —Corticarina keiferi NEW SPECIES is described from the Clarion Islands (Mexico).

Key Words.—Insecta, Coleoptera, Lathridiidae, Mexico

Clarion Island, Mexico, lies some 450 miles to the southwest of the tip of Baja

California and is part of the Revillagigedos Island group. The Island volcanically

originated along the Clarion Ridge from 1 to 2.5 million years in ago (Duncan &

Clauge 1985). No other Lathridiidae is known from this island.

CORTICARINA KEIFERI NEW SPECIES

Type Material. —Holotype. Male. MEXICO. Clarion Islands. 30 Apr 1925, H.

H. Keifer, collector. Deposited in the California Academy of Sciences. Paratypes

(40), same data. Deposited in the California Academy of Sciences and the Cali-

fornia Department of Food and Agriculture.

Description. — Length 1.3-1.6 mm. Width 0.60-0.70 mm. Pronotal length 0.30-0.42 mm. Pronotal

width 0.37-0.46 mm (n = 39). Unicolorous light red-brown. Antennae 0.30-0.40 mm long, segments

3-7 cylindrical, elongate, length 2x width, segment 8 expanded apically, slightly longer than wide,

segments 9-11 forming club, segments 9, 11 longer than wide, segment 10 as wide as long. Pronotum

1.0-1.20 x as wide as long, lateral margins rounded, widest at anterior one-third, dorsal surface evenly,

flatly rounded, no depressed areas, lateral margins smooth, without serrations. Pronotal surface smooth,

shiny, punctures small, shallow, not visible at 60 magnifications, setae short, decumbent, not visible

at 60 magnifications. Elytra elongate oval, laterally smooth, rounded, 1.4-1.6x longer than wide.

Elytron with 8 regular rows of small indistinct striae, interstriae with small indistinct rows of setae,

setae shorter than punctures. Fully winged. Eyes fully developed, 103 facets in single eye examined.

Prosterna with distinct transverse fovea anterior and lateral to procoxa, first tarsal segment produced

beyond second segment, subequal in length to third segment. Metasterna, abdominal segments regularly

clothed with short distinct decumbent setae. Male: protibia toothed on inner surface, tooth four-tenths

distance from apex. Male genitalia as in Figs. 3, 4.

Diagnosis. —Corticarina keiferi shares with other Corticarina species a similar

overall shape, form of the male genitalia, form of the male protibia, shape of the

mentum and generalized relationship between the tarsal segments. It differs, how-

ever, from them in sharing, with Melanophthalma (Cortilena), the smooth shiny

dorsal surface with small inconspicuous punctures and setae of the pronotum and

elytra. Additionally, the shape of the pronotum when viewed dorsally bears a

strong resemblance to M. (Cortilena). This species, can be differentiated from all

other Corticarina species by the characters mentioned above in conjunction with

the unique shape of the male genitalia.

13294 Meadowview Road.

1995 ANDREWS: A NEW CORTICARINA 111

Figures 1-4. Corticarina keiferi. Figure 1. Dorsal habitus. Figure 2. Ventral habitus. Figure 3.

Dorsal view male aedeagus. Figure 4. Lateral view male aedeagus.

112 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

LITERATURE CITED

Duncan, R. A. & D. A. Clauge. 1985. Pacific plate motion recorded by linear volcanic chains. Pp.

89-121. In The ocean basins and margins. Plenum Press, New York.

PAN-PACIFIC ENTOMOLOGIST

71(2): 113-120, (1995)

THE TAXONOMIC STATUS OF BOMBUS ALBOANALIS

FRANKLIN AND ITS RELATIONSHIP WITH OTHER

TAXA OF THE SUBGENUS PYROBOMBUS FROM

NORTH AMERICA AND EUROPE

(HYMENOPTERA: APIDAE)

A. SCHOLL,! R. W. THorp,”? J. A. BisHop,? AND E. OBRECHT*

1Universitat Bern, Zoologischés Institut, Baltzerstr. 3,

CH-3012 Bern, Switzerland

?University of California, Department of Entomology, Davis, California 95616

3University of Alaska, Department of Biology and Wildlife,

Fairbanks, Alaska 99775

4Naturhistorisches Museum Bern, Bernastr. 15, CH-3005 Bern, Switzerland

Abstract.— Bombus alboanalis Franklin, 1913, has either been regarded as a distinct species or

has been synonymized with B. frigidus Smith 1854, from which it differs in coat color of the

abdomen. However, both taxa are sympatric in Alaska and there is no evidence for intergradation

from coat color variation. We surveyed 18 enzymes by vertical starch-gel electrophoresis and

compared B. alboanalis with B. frigidus and other species of the subgenus Pyrobombus. We

found that B. alboanalis differs from B. frigidus at two enzyme loci suggesting that these taxa

are genetically separated. According to the enzyme survey B. alboanalis belongs to a group of

very closely related Pyrobombus species that includes B. frigidus, B. cingulatus Wahlberg and B.

jonellus (Kirby), and more distant B. sitkensis Nylander, B. mixtus Cresson, B. pratorum (L.),

and B. pyrenaeus (Pérez). We present evidence that B. alboanalis may be conspecific with B.

Jonellus.

Key Words.—Insecta, Apidae, enzyme electrophoresis, bumble bee genetic relationships, bio-

geography

Bombus alboanalis is a bumble bee from Alaska that was described as a species

by Franklin (1913). This taxon is poorly understood. Frison (1929) listed B.

alboanalis as a distinct species, but Burks (1951), who based his compilation

largely on manuscript notes by the late T. H. Frison, treated B. alboanalis as a

variety of B. frigidus Smith. Hurd (1979) followed Burks (1951) in listing B.

alboanalis as a variety of B. frigidus. A published revision, however, is not avail-

able.

Bombus alboanalis and B. frigidus are structurally identical but they differ in

their coat color of the abdomen, which is white tailed in B. alboanalis and red

tailed in B. frigidus. Polymorphism in coat color is very frequent among bumble

bees. Several polytypic species of the New World [e.g., B. occidentalis Greene]

and the Old World in particular [e.g., B. terrestris auct., B. soroeensis (Fabr.)] are

red tailed in one subspecies and white tailed in another, and a great variety of

intermediates can be found where subspecies overlap in their distribution. In

contrast, we have seen many specimens of B. alboanalis and B. frigidus from

sympatric populations in Alaska (unpublished data), but there is no evidence for

intergradation. This suggests to us that both taxa are distinct species. We have

therefore used data from enzyme electrophoresis as additional information to

investigate whether these taxa are genetically separated in areas of sympatry.

114 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Table 1. Numbers of specimens of Bombus jonellus species group studied electrophoretically and

origin of material.

British

California Columbia Alberta Alaska Norway Switzerland Total

B. alboanalis 8 8

B. cingulatus 3 3

B. frigidus 10 5 15

B. jonellus 6 14 20

B. mixtus 6 3 3 12

B. pratorum 14 14

B. pyrenaeus 10 10

B. sitkensis 2 4 6

MATERIAL AND METHODS

Bombus alboanalis and B. frigidus were collected from sympatric populations

near Fairbanks, Alaska in 1985, 1991 and 1992. Additional specimens of B.

frigidus are from three sites in southern Alberta where B. alboanalis has not been

recorded. We include for comparison electrophoretic data of Scholl et al. (1988)

on Pyrobombus species and additional specimens of six species of the B. jonellus

species group as summarized in Table 1. The specimens were shipped on dry ice

and stored at — 80°C until used for electrophoresis. In addition, specimens of both

taxa occurring sympatrically at several sites in Alaska were studied morpholog-

ically.

Electrophoresis. —Depending on the enzyme to be analysed either thoracic mus-

cle (T) or abdominal tissue (A) and one of the following three buffers were used

for electrophoresis: TBE = Tris-borate-EDTA, pH 9.3; TC = Tris-citrate, pH 7.3;

AC = citrate-N-(3-aminopropyl)-morpholine, pH 6.2. The tissue was homoge-

nized in ten volumes of Tris buffer, 0.1 M, pH 8.0. 20 ul of supernatant fractions

of homogenates were applied to vertical starch gels in Buchler instruments, using

a slotformer for 15 slots. The starch gels (13% w/v) were prepared from a mixture

of 25 g of Connaught starch and 35 g starch from the Institut des Sciences de

Evolution (Montpellier, France). The electrophoresis was run at 4° C for 15-16

h, voltage applied was 8 V cm~! (TBE buffer), 4 V cm™! (TC buffer) and 3 V

cm! (AC buffer).

The gels were sliced once or twice to provide two or three slices, each of which

was stained for a different enzyme. The enzymes scored are (tissue and buffer

system used in brackets): aconitase (A, TC), 2 loci: ACO-1 and ACO-2; arginine

kinase (A, TBE), APK; hydroxybutyric dehydrogenase (A, TC), BDH; a-glycer-

ophosphate dehydrogenase (T, TC), 2 loci: GPD-2 and GPD-3; glutamic-oxalo-

acetic transaminase (A, AC), GOT-2; glutamic-pyruvic transaminase (T, AC),

GPT; hexokinase (T, TC), HK -3; isocitrate dehydrogenase (T, AC), IDH; leucine

aminopeptidase (T, AC), LAP; malate dehydrogenase (T, AC), 2 loci: MDH-1

and MDH-2; malic enzyme (T, AC), MOD; peptidase (A, TC), PEP; phosphoglu-

cose isomerase (A, TBE), PGI; phosphoglucomutase (A, TC), PGM; superoxide

dismutase (A, TBE), SOD. Total: 18 loci.

In this survey the enzymes scored were selected by quality of resolution rather

than enzyme function. The enzyme staining followed standard procedures of our

1995 SCHOLL ET AL.: TAXONOMY OF BOMBUS 115

laboratory (Scholl et al. 1978, Bulnheim & Scholl 1981, Geiger & Scholl 1985);

most of these are originally from Ayala et al. (1972) and Harris & Hopkinson

(1976) and were slightly modified for optimal staining. Agar overlays were used

to detect the following enzymes: ACON, APK, GPT, HK, PEP, PGI and PGM.

All zymograms were photographed (Polaroid) for reference. The designation of

electromorphs is based on mobilities (in mm) relative to the electromorph of B.

lucorum (= index 100) which was used as a reference as in previous electrophoretic

studies on bumble bees (Scholl et al. 1990, 1992). A phenogram of the genetic

relationships of the species investigated was constructed by average linkage cluster

analysis (UPGMA) (Nei 1987) using Nei’s (1972) genetic distance (D).

Specimens Analyzed.—Bombus alboanalis: ALASKA. FAIRBANKS NORTH

STAR BOROUGH: nr. Fairbanks, 25 May 1990 & 25 Jun 1992.

Bombus frigidus: ALASKA. FAIRBANKS NORTH STAR BOROUGH: nr.

Fairbanks, 25 Jun 1992. CANADA. ALBERTA: Ya-Ha Tinda Ranch, E boundary

of Banff National Park, 51°43’ N, 115°30’ W, 28 May 1989; Barrier Lake, Kan-

anaskis River Valley, 51°02’ N, 115°02' W, 27 May 1989; Stettler, 8 km W, 22

May 1989.

In addition to the B. alboanalis and B. frigidus analyzed electrophoretically, we

have examined morphologically specimens of both taxa occurring sympatrically

at several other sites: ALASKA. FAIRBANKS NORTH STAR BOROUGH: Fair-

banks, 5-7 Jun 1990; MATANUSKA-SUSITNA BOROUGH: Denali Highway,

Mile 30, 15 & 21 Jul 1993, and Mile 82, 20 Jul 1993; SOUTHEAST FAIRBANKS

BOROUGH: Walker Fork Campground, Klondike Loop D-94.7, Taylor Highway,

7 Jul 1978: VALDEZ CORDOVA BOROUGH: Paxson, 14 & 20 Jul 1993; YU-

KON-KOYUKUK BOROUGH: Camp Denali, McKinley Park, 12 Jun—2 Aug

1985; Kathul Mt., Yukon River 1 May-—1 Jul 1991.

RESULTS AND DISCUSSION

Among the sympatric specimens of B. alboanalis and B. frigidus we examined

morphologically, we found no intergradation in color pattern between the two

taxa.

Allele frequencies at eleven loci which showed polymorphism and/or interspe-

cific variation are listed in Fig. 1. As in previous studies of Hymenoptera in

general (Packer & Owen, 1992) and other bumble bee species in particular (Obrecht

& Scholl 1981; Scholl et al. 1990, 1992) the level of polymorphism was very low

in the present study. In fact, most samples were monomorphic in all loci, except

B. frigidus from Alberta, where one specimen was heterozygous at the ACO-1

locus and two specimens from different sites were heterozygous at the HK-3 locus;

one specimen of B. pratorum was heterozygous at the ACO-1 locus. A rather

unusual situation, however, was observed in B. jonellus, where samples from both

Norway and Switzerland were highly polymorphic at the HK-3 locus, three alles

were Observed in similar frequencies in the populations from both countries, as

shown in Fig. 1. In addition, two specimens of B. jonellus from Switzerland were

heterozygous at the PEP locus. Seven enzymes were invariant in all specimens

surveyed. These enzymes (APK, BDH, GPD-2, GPD-3, MDH-2, PGI and SOD)

are not shown in Fig 1.

The B. alboanalis and B. frigidus from sympatric populations in Alaska showed

a consistent difference in HK-3 and PGM, resulting in the allele frequencies that

ACO-1 GOT-2 GPT HK-3 IDH LAP MDH-1 MOD PEP PGM

90 96 100}/100 102 104] 98 1057100 107) 89 93 96 100 105}100 105 113} 94 96 | 93 101} 98 102} 82 92 | 87 94

B. frigidus Alberta .00} 4. 0.90 0.10

B. cingulatus 00] 1. .00] 1.00

B. jonellus Norway 00] 1. 0.59 0.33

B. jonellus — switzerland 00] 1. ; 0.50 0.44

B. sitkensis 00 .00| 1.00

B. mixtus : é .00] 1.00

B. pratorum

B. pyrenaeus

*No variation: APK, BDH, GPD-2, GPD-3, MDH-2, PGI, SOD

Figure. 1. Allele frequencies at eleven enzyme loci that showed polymorphism and/or interspecific variation.

OTT

LSIDOTOWO.LNA OIIDVd-NVd AHL

(Z)IL TOA

1995 SCHOLL ET AL.: TAXONOMY OF BOMBUS 117

are shown in Fig. 1; all other enzymes were identical. The data on HK-3 and

PGM are of particular interest because they show that sympatric B. alboanalis

and B. frigidus have separate gene pools. The comparison of B. frigidus from

Alaska and Alberta showed minor differences, two rare alleles which were found

at the ACO-1 and HK-3 locus in Alberta were not observed in Alaska (Fig. 1).

The B. cingulatus specimens differed from B. frigidus at the HK-3 locus, having

an allele that was otherwise found in B. sitkensis, B. mixtus and B. pyrenaeus

respectively (Fig. 1), although B. jonellus was different from B. frigidus in IDH,

in addition to the HK-3 polymorphism which was mentioned above.

Clearly, more differences were observed between B. sitkensis, B. mixtus, B.

pratorum and B. pyrenaeus (Fig. 1) and in comparisons of these with the other

listed taxa. These differences need not be presented in detail here. We summarize

the genetic differentiation among the taxa surveyed in a phenogram (Fig. 2), that

was generated by average linkage cluster analysis (UPGMA), using Nei’s standard

genetic distance that was calculated from all 18 loci surveyed. This phenogram

suggests that B. alboanalis, B. frigidus, B. cingulatus and B. jonellus are very

closely related, but B. sitkensis, B. mixtus, B. pratorum and B. pyrenaeus are more

distant.

Regarding the relationships of B. alboanalis to other bumble bees, Franklin

(1913: 387) wrote: “I have seen a few workers of B. montanus Lep., determined

by Gerstaecker, and that species seems to be very closely allied to alboanalis, and

it is possible that they should be considered as variations of the same species.”

Bombus montanus (Lepeletier), 1836, is now a synonym of B. (Thoracobombus)

ruderarius (Miller), 1776, and is regarded as a subspecies (cf. Rasmont, 1983).

However, Gerstaecker (1869) apparently confused other species with B. montanus

Lep.; his B. montanus is now both in the synonymy of B. (Melanobombus) sicheli

Kriechbaumer, 1873 (Dalla Torre 1896) and B. (Pyrobombus) pyrenaeus Pérez,

1879 (Tkalcu 1969). Considering these facts, it is not possible to decide what

Franklin really had for comparison. Bombus ruderarius montanus, B. sicheli al-

ticola and B. pyrenaeus clearly differ from B. alboanalis in the coloration of the

abdomen which is red tailed in the former three taxa and white tailed in B.

alboanalis. However, Franklin (1913: 387) must have had white tailed specimens

for comparison, as he wrote: “‘the only difference in coloration of pile is that the

black interalar band of montanus is wider than that of alboanalis.”

The close genetic relationships of the nearctic B. alboanalis and B. frigidus and

the palaearctic B. cingulatus and B. jonellus that are revealed in this study, and

in particular the low level of genetic differentiation between B. alboanalis and B.

jonellus are of interest. In North America, B. alboanalis has been recorded from

Alaska and Manitoba, Canada (Hurd 1979). One of us (RWT) has identified

additional specimens from British Columbia and Yukon Territory. Franklin (1913)

mentions Bering Island and Copper Island as Asian records. Bombus frigidus has

a wider range. It is most common in Alaska and the Northwest Territories and

is sparsely distributed along higher elevations of the continental divide, apparently

as far south as Colorado (Stephen 1957, Hurd 1979). In contrast, B. cingulatus

and B. jonellus have a very wide range in the palaearctic region. Bombus cingulatus

is recorded from northern Scandinavia, east to Kamchatka, and to the Gulf of

Anadyr (Reinig 1936, 1939). Bombus jonellus has a similar range in the north

but extends west to Iceland (Prys Jones et al. 1981) and south to the Pyrenees

118 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

alboanalis

frigidus (Alaska)

frigidus (Alberta)

cingulatus

jonellus (N)

jonellus (CH)

sitkensis

mixtus

pratorum

pyrenaeus

0 0.1 0.2 0.3 0.4

Genetic distance (Nei) D

Figure. 2. Phenogram showing the genetic relationship between B. alboanalis and other taxa of

the subgenus Pyrobombus, as revealed by the electrophoretic data.

and the Balkan (Rasmont 1983) and, in Asia, has even been recorded from the

western Aleutian Islands (Panfilov 1982). Thus, B. jonellus and B. alboanalis are

parapatric.

Bombus jonellus is a polytypic species (cf. Richards 1933, Rasmont 1983). Our

electrophoresis survey covers the subspecies B. j. martes Gerstaecker, 1869 (spec-

imens from Switzerland) and B. j. subborealis Richards, 1933 (specimens from

Scandinavia). These taxa were identical in their enzyme patterns (Fig. 1). However,

considering the very wide range of this species, one might expect electrophoret-

ically detectable genetic differentiation within the range of this species even if

levels of enzyme polymorphism are very low, as commonly observed in bumble

bees (e.g., Obrecht & Scholl 1981, Pamilo et al. 1984, Scholl et al. 1990). Un-

fortunately, B. jonellus specimens from Siberia have not been available for elec-

trophoretic studies, but we speculate that future studies might show that B. al-

boanalis is conspecific with B. jonellus. Specimens of B. jonellus and B. cingulatus

from a transect from Siberia to Scandinavia would be needed to determine whether

the electrophoretic pattern of B. alboanalis grades into those of European pop-

ulations.

ACKNOWLEDGMENT

The competent assistance of Mrs. V. Siegfried-and Mrs. L. Frauchiger in the

electrophoretic studies is gratefully acknowledged; S. Hunziker made the computer

1995 SCHOLL ET AL.: TAXONOMY OF BOMBUS 119

graphics; H. J. Geiger provided specimens from Alaska for initial studies; P.

Rasmont contributed stimulating discussions. Supported by travel grants (AS)

from the Dr. Karl Bretscher Foundation (Berne).

LITERATURE CITED

Ayala, F. J., J. R. Powell, M. L. Tracey, C. A. Mourao & S. Perez-Salas. 1972. Enzyme variability

in the Drosophila willistoni group. IV. Genic variation in natural populations of Drosophila

willistoni. Genetics, 70: 113-139.

Bulnheim, H. P. & A. Scholl. 1981. Genetic variation between geographic populations of the am-

phipods Gammarus zaddachi and G. salinus. Mar. Biol., 64: 105-115.

Burks, B. D. 1951. Tribe Bombini. pp. 1247-1255. In Muesebeck, Krombein and Townes (eds.).

Hymenoptera of America north of Mexico. Synoptic catalog. Agriculture Monogr. 2.

Dalla Torre, K. W. 1896. Catalogus Hymenopterorum. 10. Apidae (Anthophila). Verlag Wilhem

Englemann, Leipzig.

Franklin, H. J. 1913. The Bombidae of the New World. Trans. Am. Entomol. Soc., 38: 177-486;

39: 73-200.

Frison, T. H. 1929. Additional descriptions, synonymy and records of North American bumblebees

(Hymenoptera: Bremidae). Trans. Am. Entomol. Soc., 55: 103-118.

Geiger, H. J. & A. Scholl. 1985. Systematics and evolution of holarctic Pierinae (Lepidoptera). An

enzyme electrophoretic approach. Experientia, 41: 24-29.

Gerstaecker, A. 1869. Beitrige zur néheren Kenntnis einiger Bienengattungen. Stettin. ent. Ztg., 30:

315-367.

Harris, H. & D. A. Hopkinson. 1976. Handbook of enzyme electrophoresis in human genetics.

North-Holland, Amsterdam.

Hurd, P. D., Jr. 1979. Apoidea. pp. 1741-2209. In Krombein, K. V., P. D. Hurd Jr., D. R. Smith

& B. D. Burks (eds.). 1979. Catalog of Hymenoptera in America north of Mexico. Smithsonian

Inst. Press, Washington, D.C.

Nei, M. 1972. Genetic distance between populations. Am. Nat., 106: 283-292.

Nei, M. 1987. Molecular Evolutionary Genetics Columbia University Press, New York.

Obrecht, E. & A. Scholl. 1981. Enzymelektrophoretische Untersuchungen zur Analyse der Ver-

wandtschaftsgrade zwischen Hummel- und Schmarotzerhummelarten (Apidae, Bombus). Ap-

idologie 12: 257-268.

Packer, L. & R. E. Owen. 1992. Variable enzyme systems in the Hymenoptera. Biochem. Syst. Ecol.

20: 1-7.

Pamilo, P., S.-L. Varvio-Aho & A. Pekkarinen. 1984. Genetic variation in bumblebees (Bombus,

Psithyrus) and putative sibling species of Bombus lucorum. Hereditas, 101: 245-251.

Panfilov, D. V. 1951. Map 150. In K. B. Gorodkova, Arealy nasekomykh evropejkoskoj chasti SSR.

Karty 126-178. Akademiya Nauk SSR, Zoologicheskij Institut, Leningradskoe Otdelenie.

Prys-Jones, O. E., E. Olafsson & K. Kristjansson. 1981. The Icelandic bumble bee fauna (Bombus

Latr., Apidae) and its distributional ecology. J. apic. Res., 20: 189-197.

Rasmont, P. 1983. Catalogue commenté des Bourdons de la région ouest-paléarctique (Hymenoptera,

Apidae, Bombinae). Notes faunistiques de Gembloux, 7: 135-160.

Reinig, W. F. 1936. Bombus cingulatus Wahlb. (Hym. Apid.). Sber. Ges. natf. Freunde Berl. Jahrg.

1936: 130-138.

Reinig, W. F. 1939. Die Evolutionsmechanismen, erléutert an den Hummeln. Zool. Anz. suppl.,

12: 170-206.

Richards, O. W. 1933. Variation in Bombus jonellus Kirby (Hymenoptera, Bombidae). Ann. Mag.

nat. Hist., 12: 59-66.

Scholl, A., B. Corzillius & W. Villwock. 1978. Beitrag zur Verwandtschaftsanalyse altweltlicher

Zahnkarpfen der Tribus Aphaniini (Pisces, Cyprinodontidae) mit Hilfe elektrophoretischer

Untersuchungsmethoden. Z. zool. Syst. Evolut.-forsch. 16: 116-132.

Scholl, A., E. Obrecht & R. W. Thorp. 1988. Biochemical systematics in bumble bees: The subgenus

Pyrobombus (Hymenoptera: Apidae). Proceedings 18th International Congress of Entomology,

Vancouver, B.C., Canada, 62.

Scholl, A., E. Obrecht & R. E. Owen. 1990. The genetic relationship between Bombus moderatus

120 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Cresson and the Bombus lucorum auct. species complex (Hymenoptera: Apidae). Can. J. Zool.,

68: 2264-2268.

1992. Specific Distinctiveness of Bombus

Scholl, A., R. W. Thorp, R. E. Owen & E. Obrecht.

nevadensis Cresson and B. auricomus (Robertson) (Hymenoptera: Apidae)- Enzyme Electro-

phoretic Data. J. Kansas Entomol. Soc., 65: 134-140.

Stephen, W. P. 1957. Bumble bees of western America (Hymenoptera: Apoidea). Tech. Bull. Oreg

agric. Exp. Stn., 40.

Tkalcu, B. 1969. Ergebnisse der Albanienexpedition 1961 des Deutschen Entomologischen Institutes

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PAN-PACIFIC ENTOMOLOGIST

71(2): 121-124, (1995)

ADULT OVERWINTER MORTALITY IN

OSMIA LIGNARIA PROPINQUA CRESSON

(HYMENOPTERA: MEGACHILIDAE)

RICHARD W. RUST

Department of Biology and Program in Ecology,

Evolution and Conservation Biology,

University of Nevada, Reno, Nevada 89557

Abstract.— Adult Osmia lignaria propinqua Cresson from Reno, Nevada that were overwintered

in Logan, Utah had increased winter mortality, reduced emergence, and lower reproductive

success compared to a similar population of bees from Logan, Utah. Both lower average monthly

temperatures and extreme minima were responsible for the winter loss.

Key Words.—Insecta, Osmia, overwinter, mortality

Unusual events, such as atypical precipitation or temperature patterns, were

invoked by researchers to explain unusual events in field data in 9% percent of

insect studies (n = 54) and in 9% of all studies (n = 354) analyzed by Weatherhead

(1986). When separated by ecosystem, unusual events were used to explain 5%

(38) of temperate, 12.5% (8) of tropical, and 25.0% (8) of desert insect studies

(Weatherhead 1986). These unusual events may be more common than expected,

greater than one in 20 chance, and may function to keep populations below their

carrying capacity (MacArthur 1972, Wiens 1977).

Here I present both observational and experimental data on the effects of un-

usual density independent temperature events on the overwinter losses in two

populations of the bee, Osmia lignaria propinqua Cresson. One population (Logan,

Utah) with frequent exposure to extreme low winter temperatures showed little

measurable winter mortality. Whereas, another population (Reno, Nevada) when

exposed to extreme low winter temperatures showed reduced survivorship.

Osmia I. propinqua overwinter as adults inside the larval cocoon (Rau 1937,

Krombein 1967, Rust 1974, Torchio 1989). Pupation occurs during June and

July, and by August all individuals that successfully complete immature devel-

opment are adults. In the laboratory, bees are successfully overwintered at 4° C

until their release in apple, prune, or almond orchards the following spring (5 to

6 months) (Torchio 1976, 1982, 1985).

MATERIALS AND METHODS

Bees used in this study were from trap-nests placed out in Reno and Logan

during the spring and summer of 1988. In September, trap-nests were recovered

and some Reno trap-nests were moved to Logan for overwintering. In Reno, bees

were provided with pine blocks drilled with 5, 6, 7, 8, and 9 mm diameter holes

to a depth of 135 to 140 mm (Rust 1990). In Logan, bees were provided with

drilled wooden blocks with 7 mm by 140 mm paper straw inserts (Torchio 1976,

1985). Nests were overwintered in an unheated, metal storage building on the

Utah State University campus, Logan, Utah. In March, the nests were brought

into the laboratory for examination and data recording. Nests from Reno con-

122 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Table 1. Overwintering and emergence parameters for two populations of Osmia lignaria propinqua

Cresson. One population from Reno, Nevada and the other Logan, Utah. Both populations were

overwintered in Logan, Utah. All paired data sets are significantly different at P < 0.001. Data presented

are means and standard deviations.

Males Females

Parameter Reno Logan Reno Logan

Weight—mg Ji.3 + 7.3 65.7 + 24.4 61.4 + 22.6 92.3 + 43.3

Hours emerge 28.9 + 9.8 59.7 + 15.8 50.0 + 18.5 106.4 + 22.1

Hours lived 18.0 & 215 140.4 + 28.8 88.7 + 19.7 170.4 + 42.9

% weight loss 17.6 + 8.0 40.1 + 6.1 18.2 + 6.4 33.3 + 6.8

tained both dead adults and live adults that showed limited movement when

exposed to laboratory room temperature (21 + 2° C). Individuals from 19 ran-

domly selected nests from each test site were left inside their cocoons, weighed

(0.1 mg) and placed in individual plastic vials (21 mm diameter x 40 mm). These

individuals were placed in a temperature cabinet held at 21° C + 1.0° C, 70 +

5% relative humidity, and 24 h dark. Individuals were checked at 12 h intervals

for emergence from their cocoon and length of life. Cocoons were removed,

weighed and adult weight was determined by subtraction. Adult dead weight was

also measured.

Climatic data for Reno and Logan were assembled for the winter months (Oc-

tober through March) (USDC-NOAA 1949-1991; USDC-NOAA 1988-1989).

Males and females from Reno and Logan were compared for emergence weight,

hours to emerge, hours lived and percent weight loss by t-test (Zar 1974). Mean

monthly minimum temperatures for October through March for both sites were

also compared by f-test.

RESULTS

The 19 Reno nests contained 77 cells (40 males and 37 females) with adult

bees of which 11 (six males and five females) were dead when opened in March.

The 19 Logan nests contained 79 live individuals (56 males and 23 females). Both

male and female O. /. propinqua from Reno were significantly lighter at emergence,

required less time to emerge form their cocoons, lived a shorter period of time,

and had less weight loss from emergence until death than Logan individuals (Ta-

ble 1).

Twice during the overwintering period (October to March) the populations

experienced six day periods (24 to 29 Jan and 5 to 10 Feb) of minimum tem-

peratures at or below —17.0° C with a minimum low of —27.2° C (7 Feb).

Mean minimum temperatures over a 30 year period were significantly lower

in Reno during October (—0.3 + 1.0° C and 3.6 + 2.2° C; t = 7.73, df = 58, P

< 0.001) and November (—3.7 + 2.1° C and —2.4 + 2.0°C; t = 2.45, df = 58,

0.02 > P > 0.01). Whereas, Logan temperatures were lower during January (—8.7

+ 3.2° Cand —6.6 + 2.5° C; t = 2.79, df = 58, 0.01 > P > 0.005) and February

(—6.6 + 3.0° Cand —4.2 + 1.8° C; t = 3.84, df = 58, P < 0.001). There was no

difference in December (Reno —6.8 + 2.8° C and Logan —7.2 + 2.1°C; t = 0.44,

df = 58, P > 0.50) or March (Reno —2.8 + 1.5° C and Logan —3.1 + 2.1°C;t

= 0.56, df = 58, P > 0.50) (Fig. 1). The 30 year monthly minimum temperatures

1995 RUST: OVERWINTER MORTALITY IN OSMIA 123

30 year mean monthly minimum

10 and standard deviation

re)

Bo a

= “10 on :

a. P nine, NV

= = . e cegan: UT

-30 s 8 &

30 year monthly minimum

Oct Nov Dec Jan Feb Mar

MONTHS

Figure 1. Thirty year mean and standard deviation of the monthly temperatures for October

through March for Logan, Utah and Reno, Nevada and the 30 year monthly minimum temperatures.

were from four to eight degrees lower in Logan from November through February

(Fig. 1).

DISCUSSION

The Reno population of O. /. propinqua experiencing lower winter temperatures

in Logan had an increased adult winter mortality, reduced emergence longevity,

and limited offspring production (Rust & Torchio 1991) when compared to the

Logan population exposed to the same overwintering conditions. Several ques-

tions emerge about these observations and measurements. First, do the temper-

ature differences between Logan and Reno allow for adaptation of Logan bees to

colder winter condition but not the Reno bees? Second, how often, if ever, have

the Reno bees been exposed to these cold temperatures? The mean monthly winter

minimum temperatures for Logan and Reno were somewhat similar, but the

recorded monthly minimums were colder by 4° to 8° C in Logan during four of

the six winter months. This suggests selection and possible adaptation to lower

cold temperatures in Logan bees.

If we assume an event is rare, as measured statistically at P < 0.05, then one

would expect —27.2° C in one of 20 years. The coldest recorded minimum tem-

perature in Reno, Nevada was —28.3° C in January 1890 (USDC-NOAA 1974)

124 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

and from 1949 to 1991 the coldest Reno, Nevada temperature was —26.6° C.

This low temperature was recorded in three of the 42 years (P = 0.09), not a rare

event. Raising the low temperature to — 17.2° C in Reno increased the cold periods

to 27 in 42 years (P = 0.64) or a common annual event. Also, the —17.2° C was

recorded in 43 of the 252 winter months (P = 0.17). Thus, Reno bees frequently

experience cold temperatures but less frequently the extreme cold temperatures

of the Logan, Utah area.

Little is known about the biology or physiology of overwintering in O. /. pro-

pinqua or for any solitary bee species (Krombein 1967, Michener 1972, Gauld &

Bolton 1988). Thus, the effect of one period of extreme cold versus the accu-

mulated effects of average colder winter temperatures are unknown. However,

this study shows that Reno bees exposed to both colder minimum and colder

average minimum temperatures did not survive to produce offspring and those

that did survive were short lived and produced no offspring. A Reno population

experiencing these winter minima should be severely reduced in size and should

show selection for potentially more cold hardy individuals.

ACKNOWLEDGMENT

I thank P. F. Torchio and G. Trostle (USDA, Logan, Utah) for their assistance

with this project, and D. Charlet and C. Cripps (Univ. Nevada, Reno) for reviewing

the manuscript.

LITERATURE CITED

Gauld, I. & B. Bolton. 1988. The Hymenoptera. Oxford University Press, Oxford, Great Britain.

Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests, and associates. Smithsonian

Institution, Washington, D.C.

MacArthur, R. H. 1972. Geographical ecology. Harper & Row, New York.

Michener, C. D. 1974. The social behavior of bees. Harvard University Press, Cambridge.

Rau, P. 1937. The life history of Osmia lignaria and Osmia cordata, with notes on Osmia conjuncta.

Ann. Entomol. Soc. Am., 39: 324-342.

Rust, R. W. 1974. The systematics and biology of the genus Osmia, subgenera Osmia, Chalcosmia,

and Cephalosmia (Hymenoptera: Megachilidae). Wassmann J. Biol., 32: 1-93.

Rust, R. W. 1990. Spatial and temporal heterogeneity of pollen foraging in Osmia lignaria propinqua

(Hymenoptera: Megachilidae). Environ. Entomol., 19: 332-338.

Rust, R. W. & P. F. Torchio. 1991. Induction of Ascosphaera (Ascomycetes: Ascosphaerales) infec-

tions in field populations of Osmia lignaria propinqua Cresson (Hymenoptera: Megachilidae).

Pan-Pacific Entomol., 67: 251-257.

Torchio, P. F. 1976. Use of Osmia lignaria Say (Hymenoptera: Megachilidae) as a pollinator in an

apple and prune orchard. J. Kansas Entomol. Soc., 49: 475-482.

Torchio, P. F. 1982. Field experiments with Osmia lignaria propinqua Cresson as a pollintor in

almond orchards: I, 1975 studies. J. Kansas Entomol. Soc., 54: 815-823.

Torchio, P. F. 1985. Field experiments with the pollinator species, Osmia lignaria propinqua Cresson,

in apple orchards: success, seed counts, fruit yields (Hymenoptera: Megachilidae). J. Kansas

Entomol. Soc., 58: 448-464.

Torchio, P. F. 1989. In-nest biologies and development of immature stages of three Osmia species.

Ann. Entomol. Soc. Am., 82: 599-615.

USDC-NOAA. 1949-1991. Climatological data annual summary. Nevada.

USDC-NOAA. 1988-1989. Climatological data annual summary. Utah.

Weatherhead, P. J. 1986. How unusual are unusual events? Am. Nat., 128: 150-154.

Wiens, J. A. 1977. On competitive and variable environments. Am. Sci., 65: 590-597.

Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Englewood Cliffs, New Jersey.

PAN-PACIFIC ENTOMOLOGIST

71(2): 125-129, (1995)

INCIPIENT SEXUAL ISOLATION IN

DROSOPHILA PSEUDOOBSCURA BOGOTANA

AYALA & DOBZHANSKY

(DIPTERA: DROSOPHILIDAE)*

MOHAMED A. Noor

Department of Ecology and Evolution, University of Chicago,

Chicago, Illinois 60637

Abstract. —Incipient sexual isolation was found between the subspecies Drosophila pseudoobscura

bogotana (Ayala and Dobzhansky) and D. p. pseudoobscura (Frolova). Males took longer to

initiate courtship with heterosubspecifics than consubspecifics. This divergence could represent

an early step in the speciation of these taxa. The subspecies’ pheromones do not appear to have

diverged, so the difference is attributed to female behavior.

Key Words.—Insecta, Drosophila pseudoobscura, Drosophila pseudoobscura bogotana, mating

behavior, sexual isolation, reproductive isolation, speciation

Behavioral characters in geographically isolated populations of a species can

diverge over time. If they are separated long enough, their sexual behavior might

differentiate to the extent that individuals of one population would not recognize

individuals of the other population as potential mates. The two populations will

then be different species.

Many Drosophila species have courtship patterns that prevent them from hy-

bridizing with other species in nature (Brown 1965, Spieth 1951, Carson et al.

1989). Conspecific populations which are geographically isolated and diverging

genetically provide ideal subjects for studying incipient sexual isolation. Does the

divergence evolve rapidly, or are these behavioral changes gradual? Studies of the

initial stages of ethological differentiation have been largely neglected; researchers

have focused instead on estimating sexual isolation between sibling species pairs

(Ryan & Wilczynski 1991). I present a test of incipient sexual isolation in two

subspecies of Drosophila.

Drosophila pseudoobscura bogotana occurs at high elevations in the vicinty of

Bogota, Columbia, nearly 2400 km from the main body of the species (Dob-

zhansky et al. 1963). It has differentiated genetically from the North American

mainland D. pseudoobscura pseudoobscura Frolova, as evidenced by allozyme

(Ayala & Dobzhansky 1974, Coyne & Felton 1977, Singh 1983) and DNA-se-

quence analyses (Schaeffer & Miller 1991); the latter estimated that they became

geographically isolated approximately 150,000 years ago. Although there appears

to be extensive gene flow among the North American populations, there is no

evidence for gene exchange between the subspecies (Schaeffer & Miller 1992).

Intersubspecific hybrid male offspring with bogotana mothers are sterile, while

all other hybrid offspring are fertile.

There has been little research on the behavioral divergence between these sub-

species. Prakash (1972) found that matings between consubspecifics were as fre-

* Author page charges partially offset by a grant from the C. P. Alexander Fund, PCES.

126 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

quent as matings between heterosubspecifics in chambers containing both sub-

species. However, Singh (1983) noted that bogotana females preferred pseudoob-

scura males to their own males in such mixed cages.

Here, I report a new test of behavioral divergence of the subspecies D. p.

bogotana and D. p. pseudoobscura. Since these subspecies are truly allopatric, any

behavioral differences must have arisen by drift or correlated selection pressures

imposed by their different environments rather than selection in sympatry to

reduce hybridization (see Dobzhansky 1940).

MATERIALS AND METHODS

I observed the courtship of bogotana and pseudoobscura in several intrasub-

specific and intersubspecific matings. Differences in length of time until courtship

was initiated (courtship latency) and length of time between courtship initiation

and copulation (courtship duration) were noted. The former shows the length of

time the male took to identify the female as a potentially receptive mate. The

latter shows how quickly the female allowed the male to copulate. These two

characters represent differences in the specific behavioral components that can

confer sexual isolation, such as differing acoustical, visual, olfactory, or tactile

cues.

The bogotana strains used were an isofemale line from el Recreo (hereafter

BOG1), originally captured in 1978, and line 0121.35 from the National Species

Resource Center (hereafter BOG2), originally captured in 1960. They were chosen

because they were hardy and showed no obvious symptoms of inbreeding de-

pression, such as a consistent lack of intrastrain matings in 20 min observations.

The USA1 stock was a mixture of four isofemale lines captured in the Gundlach-

Bundschu Winery in the Sonoma Valley of California in 1988. The USA2 stock

was an isofemale line collected in Flagstaff, Arizona in 1993.

Stocks were kept at 20° C on cornmeal/Karo syrup/yeast/agar food. Carbon

dioxide was used for anesthetization. Virgin males and females were collected

daily and kept individually in vials for 7 d to reach sexual maturity. Single male

and female pairs were then placed without anesthesia in an 8 dram food-containing

vial and observed for up to 20 min. Each combination was repeated 20 to 30

times. The time of first male courtship after introduction and time of copulation

were recorded. First courtships were usually wing-vibrations (Brown 1963), but

occasionally a male attempted to mount a female directly. The courtship duration

was estimated as the time of first courting subtracted from the time of copulation.

Any pairings in which the male or female was clearly weak or injured were

excluded. The rare pairings (approximately 5%) in which copulation did not occur

in the first 20 min were also excluded, since they likely resulted from nonvirginity

or abnormalities in the specific flies. These unsuccessful pairings were randomly

distributed among the crosses.

Because some strains had been in culture for several years, their behavior may

have diverged from that of flies in nature. To correct for any selection within the

strains, I compared interstrain crosses within subspecies to crosses with the other

subspecies (see Tables 1 and 2). I performed Kruskal-Wallis and Mann-Whitney

U-tests on courtship latency and courtship duration to assess differences in sexual

behavior.

1995 NOOR: SEXUAL ISOLATION IN DROSOPHILA 127

Table 1. Median courtship latencies (sec) and P value as estimated by two-tailed Mann-Whitney

U-test.

Intrasubsp. n Median T Intersubsp. n Median T P

BOGI Male x BOG2 Female 29 #615 USAI1 Female 28. 32 0.0043

BOG2 Male x BOGI1 Female 29 14 USA1 Female 22 19.5 0.5875

BOGI1 Female x BOG2 Male 29 «14 USA1 Male 29 39 0.0180

BOG2 Female x BOGI1 Male 29 «#15 USAI1 Male 22 15.5 0.5873

USAI1 Male x USA2 Female 27 ~=16 BOGI Female 29 39 0.0136

USAI1 Male x USA2 Female 27 ~=16 BOG? Female 22 15.5 0.8721

USAI1 Female x USA2 Male 24 12.5 BOGI Male 28 32 0.0001

USAI1 Female x USA2 Male 24 12.5 BOG2 Male 22 19.5 0.0942

RESULTS

A Kruskal-Wallis test on courtship latencies showed significant heterogeneity

among all crosses (two-tailed H = 25.34, 7 df, P = 0.0007), with the mean ranks

of the intrasubspecific crosses all smaller than the mean ranks of the intersub-

specific crosses (Table 1). The probability values from the Mann-Whitney U-tests

were combined for two broad tests of significance, as described by Fisher (see

Sokal & Rohlf 1969); the composite statistics showed significant tendencies for

the male flies to court consubspecific females more quickly than heterosubspecific

females (two-tailed x? = 20.83, 8 df, P < 0.025) and for the females to be courted

more quickly by consubspecific males than heterosubspecific males (two-tailed x?

= 32.24, 8 df, P < 0.001). These courtship latency data clearly indicate some

subspecies discrimination.

The data for courtship duration were less conclusive. The Kruskal-Wallis test

showed heterogeneity among all crosses (H = 19.38, 7 df, P = 0.0062), but

intrasubspecific matings were not all faster than the intersubspecific matings (Table

2). However, no intersubspecific crosses were significantly faster than the corre-

sponding intrasubspecific crosses.

DISCUSSION

These experiments demonstrate that D. p. bogotana has differentiated ethol-

ogically from the mainland D. p. pseudoobscura. Although the differences are

minor compared to the level of sexual isolation between D. pseudoobscura and

Table 2. Median courtship durations (sec) and P value as estimated by two-tailed Mann-Whitney

U-test.

Intrasubsp. n Median T Intersubsp. n Median T P

BOGI Male x BOG2 Female 27 15 USA1 Female je. “13S 0.9919

BOG2 Male x BOGI1 Female 29 9 USAI Female 17 45 0.0060

BOGI1 Female x BOG2 Male 29 9 USAI1 Male 28 36 0.0549

BOG2 Female x BOGI1 Male at | Bs USA1 Male 20 7 0.3231

USAI Male x USA2 Female 27 8 BOGI1 Female 28 36 0.0046

USAI1 Male x USA2 Female 27 8 BOG2 Female 20 7 0.4872

USAI Female x USA2 Male 25 10 BOGI1 Male a2 “33 0.7133

USAI1 Female x USA2 Male 25 10 BOG2 Male 17 45 0.0067

128 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

its sibling species, D. persimilis Dobzhansky and Epling (Tan 1946), they could

represent an early stage in the development of stronger, and perhaps complete,

isolation. The minor behavioral differences associated with the one-way hybrid

male sterility in these taxa agree with previous observations that premating and

postmating isolation may evolve in a correlated fashion among allopatric sibling

species pairs (Coyne & Orr 1989).

Singh (1983) also noted that intersubspecific matings took longer than intra-

subspecific matings in nonchoice experiments in these taxa, but he attributed the

delays to female discrimination. The results of this study demonstrate that female

subspecies discrimination may play little or no part in the observed mating delays.

The specific components of courtship or mate preference that have diverged

are not clear. The male flies appeared to not recognize the heterosubspecific

females as quickly as they recognized the consubspecific females. This could have

resulted from a difference in the females’ behavior or pheromones. Mayr (1946)

presented evidence that D. pseudoobscura males use smell in species discrim1-

nation, so a slight change in the female bogotana pheromone could have produced

the observed results. However, we could detect no difference in the cuticular

hydrocarbons of the subspecies using gas chromatography (Noor unpublished

data). Thus, a female behavioral difference seems more likely, although D. pseu-

doobscura females appear to play a passive role in courtship (Brown 1963).

Finally, the results of this study further support the idea that populations can

undergo substantial genetic divergence without developing strong behavioral iso-

lation (Coyne & Orr 1989).

ACKNOWLEDGMENT

I thank Jerry Coyne, Steve Schaeffer, Phil Service, and the National Drosophila

Species Resource Center for help in obtaining fly stocks; Jerry Coyne, Norman

Johnson, Shyril O’Steen, and two anonymous reviewers for helpful comments

and criticisms of the manuscript; and Anne Crittenden for technical assistance

and moral support. Financial support was provided by a Genetics Training Grant

from the Department of Health and Human Services, a Grant-in-Aid of research

from Sigma Xi, a Hinds Fund grant from the University of Chicago, and grant

GM 38462 from the National Institutes of Health to Jerry Coyne.

LITERATURE CITED

Ayala, F. J. & T. Dobzhansky. 1974. A new subspecies of Drosophila pseudoobscura. Pan-Pacif.

Entomol., 50: 211-219.

Brown, R. G. B. 1963. Courtship behaviour in the Drosophila obscura group. I.: D. pseudoobscura.

Behaviour, 23: 61-106.

Brown, R. G. B. 1965. Courtship behaviour in the Drosophila obscura group. Part II. Comparative

studies. Behaviour, 25: 281-323.

Carson, H. L., K. Y. Kaneshiro & F. C. Val. 1989. Natural hybridization between the sympatric

Hawaiian species Drosophila silvestris and Drosophila heteroneura. Evolution, 43: 190-203.

Coyne, J. A. & A. A. Felton. 1977. Genic heterogeneity at two alcohol dehydrogenase loci in

Drosophila pseudoobscura and D. persimilis. Genetics, 87: 285-304.

Coyne, J. A. & H. A. Orr. 1989. Patterns of speciation in Drosophila. Evolution, 43: 362-381.

Dobzhansky, T. 1940. Speciation as a stage in evolutionary divergence. Am. Nat., 74: 312-321.

Dobzhansky, T., A. S. Hunter, O. Pavlovsky, B. Spassky & B. Wallace. 1963. Genetics of natural

populations. XX XI. Genetics of an isolated marginal population of Drosophila pseudoobscura.

Genetics, 48: 91-103.

1995 NOOR: SEXUAL ISOLATION IN DROSOPHILA 129

Mayr, E. 1946. Experiments on sexual isolation in Drosophila. VII. The nature of the isolating

mechanisms between Drosophila pseudoobscura and Drosophila persimilis. Proc. Natl. Acad.

Sci. USA, 32: 128-137.

Prakash, S. 1972. Origin of reproductive isolation in the absence of apparent genic differentiation

in a geographic isolate of Drosophila pseudoobscura. Genetics, 72: 143-155.

Ryan, M. J. & W. Wilczynski. 1991. Evolution of intraspecific variation in the advertisement call

of a cricket frog (Acris crepitans, Hylidae). Biol. J. Linn. Soc., 44: 249-271.

Schaeffer, S. W. & E. L. Miller. 1991. Nucleotide sequence analysis of Adh genes estimates the time

of geographic isolation of the Bogota population of Drosophila pseudoobscura. Proc. Natl. Acad.

Sci. USA, 88: 6097-6101.

Schaeffer, S. W. & E. L. Miller. 1992. Estimates of gene flow in Drosophila pseudooscura determined

from nucleotide sequence analysis of the alcohol dehydrogenase region. Genetics, 132: 471-

480.

Singh, R. S. 1983. Genetic differentiation for allozymes and fitness characters between mainland

and Bogota populations of Drosophila pseudoobscura. Can. J. Genet. Cytol., 25: 590-604.

Sokal, R. R. & F. J. Rohlf. 1969. Biometry. W. H. Freeman and Company, San Francisco.

Spieth, H. T. 1951. Mating behavior and sexual isolation in the Drosophila virilis species group.

Behaviour, 3: 105-145.

Tan, C. C. 1946. Genetics of sexual isolation between Drosophila pseudoobscura and Drosophila

persimilis. Genetics, 31: 558-573.

PAN-PACIFIC ENTOMOLOGIST

71(2): 130-132, (1995)

DESCRIPTION OF THE FEMALE OF

NOMADA DREISBACHORUM MOALIF

(HYMENOPTERA: APOIDEA: NOMADINAE)

Byron A. ALEXANDER

Snow Entomological Museum, Snow Hall, University of Kansas,

Lawrence, Kansas 66045

Abstract. —The female of the cleptoparasitic bee Nomada (Pachynomada) dreisbachorum Moalif

is described for the first time, on the basis of specimens collected in the Distrito Federal of

Mexico in October and November, 1991.

Key Words. —Insecta, Hymenoptera, Apoidea, Nomadinae, vincta group, Nomada dreisbacho-

rum

In 1988, a revision (Broemeling & Moalif 1988) of the species of the subgenus

Pachynomada Rodeck, a monophyletic group within the genus Nomada Scopoli

(equivalent to the vincta species group sensu Alexander 1994), was published that

recognized fourteen species and subspecies, five of them new. For four of these

fourteen species, only one sex is currently known: the female only for N. saltillo

Broemeling and N. vitticollis Cresson, the male only for N. tepoztlan Moalif and

N. dreisbachorum Moalif.

Broemeling and Moalif’s revision shows that the vincta group has its greatest

diversity in Mexico. Over the past two years, a survey of the wild bee fauna of

Mexico has been underway, supported by the U.S. National Science Foundation

and the Instituto de Biologia, Universidad Nacional Autonoma de Mexico

(UNAM). A collecting trip through parts of the Altiplano and coastal Veracruz

in the late autumn of 1991 (described in Griswold 1992) turned up a modest

series of Nomada dreisbachorum that included nine female specimens. Because

no further representatives of any species of the vincta group have been collected

in any of the three other expeditions funded by this grant, and the recent revision

by Broemeling and Moalif was based upon a comprehensive survey of the material

that was already available in collections, a description of the hitherto undescribed

female of Nomada dreisbachorum is presented below. Format and terminology

follow that of Broemeling & Moalif (1988); abbreviations S and T refer to meta-

somal sternum and tergum, respectively. All measurements are presented as arith-

metic mean, with range in parentheses. Specimens are deposited in the collections

of the Instituto de Biologia at UNAM, the Bee Biology and Systematics Laboratory

at Utah State University, and the Snow Entomological Museum at the University

of Kansas.

NOMADA DREISBACHORUM MOALIF

Nomada (Pachynomada) dreisbachorum Moalif, in Broemeling & Moalif, 1988.

Pan-Pacif. Entomol., 64: 222. male.

Types. — Holotype, male; data: ““Amecameca, Mex. 9-25-57, R. & K. Dreis-

bach”; deposited: Michigan State University, East Lansing, Michigan. Paratypes:

1995 ALEXANDER: FEMALE NOMIA DREISBACHORUM 131

same data as holotype, 4 males; deposited: 3 at Michigan State University, 1 at

Bee Biology & Systematics Laboratory, Utah State University, Logan, Utah (ex-

amined October, 1993). Type depositories are as reported in Broemeling & Moalif

(1988), but holotype and 3 paratypes could not be found at Michigan State Uni-

versity in October, 1993 (Virginia Scott, personal communication).

Male. —Described in detail in Broemeling & Moalif (1988).

Female. —Body length 11.2 mm (10.4—12.4); forewing length, measured from base of costal vein to

apex of marginal cell, 8.8 mm (8.5—9.2); hindwing length, measured from base of vein Sc+R to base

of vein Rs, 4.7 mm (4.3-5.0); antennal scape cylindrical, not swollen, anterior face covered with

shallow, flattened punctures, integument between punctures dull, coarsely shagreened; head densely

covered with round, shallow punctures separated by less than a puncture diameter except on apical

one-half of clypeus and malar area, where spacing is more irregular and sparser; integument between

punctures dull and coarsely roughened except on apical one-half of clypeus and malar area, where it

is smooth and polished; ocellocular distance 0.43 mm (0.42-0.50); mid-lateral ocellar distance 0.14

mm (0.12-0.14); mid-ocellar occipital margin distance 0.48 mm (0.44—0.54); mid-ocellar diameter

0.23 mm (0.22-0.26); interocellar distance 0.38 mm (0.36-0.38); paraocular “carina” weakly ex-

pressed, but surface between anterolateral corners of clypeus and inner margin of compound eye

angulate rather than evenly convex; labrum with irregular coarse, shallow punctures, integument

between punctures smooth and polished, apex produced into thick, prominent protuberance; acetabular

carina of mandible less prominent than in male, mandible otherwise as in male; pronotal collar narrow

mesally, lateral portion of collar rounded when viewed from front, but angulate along crest of curve;

scutum very densely punctate, punctures deep and contiguous, integument dull and rough, vestiture

a mixture of sparsely distributed setae that are erect, flexible, weakly plumose, pale yellow, about as

long as second flagellomere; and somewhat denser layer of very short, simple, recumbent setae (vestiture

of scutum most apparent in oblique lateral view, scarcely discernible in dorsal view); scutellum with

discernible dorsal and posterior faces, but evenly rounded in profile, median longitudinal depression

very weak or absent, punctation and vestiture as on scutum; sides of propodeum with distinct, shallow

punctures irregularly spaced, ranging from contiguous to separated by 1 or 2 puncture diameters,

integument between punctures rugulose or shagreened; vestiture resembles erect setae of scutum, with

no understory of short, recumbent setae; propodeal triangle bare, with prominent longitudinal rugae

on basal third (adjacent to metanotum), grading into apical one-half that is impunctate and weakly

shagreened; metapleuron dorsally with sculpturing and vestiture as on sides of propodeum, but ven-

trally impunctate, shiny, only slightly roughened, setae very short and fine, weakly plumose; meso-

pleuron with vestiture and sculpturing similar to sides of propodeum, but punctures slightly larger

and more evenly spaced; procoxal spine rudiment strongly developed; apex of hind tibia with 5-8

long, thin, somewhat dorsoventrally compressed bristles, each tapering to sharp point, these bristles

about as long as surrounding plumose setae, but much broader at base and ferruginous rather than

pale yellow; hind basitarsus not distinctly widened medially, anterior and posterior margins essentially

parallel; forewing with 2 submarginal cells of roughly equal size (1 specimen has short remnant of

second transverse cubital vein in 1 wing); wings weakly infumate throughout, more strongly clouded

along principal longitudinal veins, within apical one-half of marginal cell, and along apex of forewing.

S5, T5, S6 of typical form for female Nomada (see Alexander 1994: 179). COLOR: integument

primarily black except mandibles dark red-brown near apex; ocelli, tibial spurs, and apex of distalmost

antennal segment brown-yellow; and the following maculations yellow: irregular spot of variable size

near apical margin of clypeus; narrow lines along inner margin of compound eye and in malar space;

small spot on mandibular acetabulum (3 specimens) or on outer face of mandible about midway

between base and apex (1 specimen); lateral angles of pronotal collar; pronotal lobes; tegulae in part;

an irregular yellow strip on ventral one-half of mesopleuron (absent on 2 specimens, reduced to small

spots on 1 specimen); 2 spots on scutellum; mesal part of metanotum (extending onto scutellum on

7 specimens); narrow transverse median bands on T1-5, first 2 or 3 terga with band extending anteriorly

on each side to create a U-shaped maculation; large median patches on S1-3, small lateral spots on

S4 and SS.

Discussion. —The male of this species, characterized by Broemeling and Moalif

as ‘“‘the furthest [sic] from the norm of Pachynomada,” is easily recognized by a

132 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

suite of diagnostic characters. The nine female specimens collected at the same

time and place as twelve males can be readily assigned to the same species by the

following combination of characters that Broemeling and Moalif identified as

diagnostic: dull, shagreened interpunctural surface of the head, 2 submarginal cells

(due to loss of second transverse cubital vein), body black with greatly reduced

yellow markings, pro-coxal spine rudiment strong, hind basitarsus only slightly

(if at all) expanded medially. The last character listed is of some note because

one of the characters that has been considered diagnostic for the vincta group is

a hind basitarsus that is “expanded medially’’ (Broemeling & Moalif 1988) or

““somewhat swollen and widest near the middle” (Snelling 1986). This is a subtle

character at best, and scarcely discernible in Nomada dreisbachorum.

As usual in Nomada, the bristles at the apex of the hind tibia are somewhat

more robust and deeply pigmented in females than in males of Nomada dreis-

bachorum. Broemeling & Moalif’s (1988) description refers to “[yellow] patches

on sterna 1-3.”’ The new specimens (of both sexes) examined for this paper have

large median yellow maculations on S1-3 and small lateral spots on S4 and S5,

with the latter absent, or possibly hidden, in 3 specimens.

Material Examined. —Paratype, male, from Utah State University collection (see Types). MEXICO.

DISTRITO FEDERAL: 6 km SW of Milpa Alta, 28 Oct 1991, 2600 m, “‘wooded area just after cactus

gardens”, F. Nogueira, 1 female; 13 km SW of Milpa Alta, 28 Oct 1991, 2700 m, “‘steep field above

Salvia patches, bee hive in quarry,” R. Ayala, A. Rodriguez, 5 males; 13 km SW of Milpa Alta, 28

Oct 1991, 2700 m, T. Griswold, 5 males, 6 females; 6 km SW of Milpa Alta, 12 Nov 1991, 2600 m,

“‘wooded area after cactus gardens,” R. Ayala, A. Rodriguez, F. Nogueira, 2 females, 2 males.

ACKNOWLEDGMENT

I thank Virginia Scott of Michigan State University for attempting to find the

holotype of Nomada dreisbachorum, T. Griswold for loaning a paratype of N.

dreisbachorum from the Utah State University collection as well as the specimens

he collected in Mexico, W. E. LaBerge of the Illinois Natural History Survey for

loaning the specimens collected by Ayala, Nogueira, and Rodriguez, and Charles

D. Michener of the University of Kansas and one anonymous referee for reviewing

the manuscript. This project was supported by funds from NSF grant BSR 90-

24723 to W. E. LaBerge and R. J. McGinley. This paper is contribution No. 3123

from the Department of Entomology, University of Kansas.

LITERATURE CITED

Alexander, B. A. 1994. Species-groups and cladistic analysis of the cleptoparasitic bee genus Nomada

(Hymenoptera: Apoidea). Univ. Kansas Sci. Bull., 55: 175-238.

Broemeling, D. K. & A. S. Moalif. 1988. A revision of the Nomada subgenus Pachynomada (Hy-

menoptera: Anthophoridae). Pan-Pacif. Entomol., 64: 201-227.

Griswold, T. 1992. Report on the second NSF-funded PCAM expedition. Melissa, 5: 2-3.

Snelling, R.R. 1986. Contributions toward a revision of the New World nomadine bees: a partitioning

of the genus Nomada (Hymenoptera: Anthophoridae). Nat. Hist. Mus. Los Angeles County

Contrib. Sci., 376.

PAN-PACIFIC ENTOMOLOGIST

71(2): 133-134, (1995)

Scientific Note

THE BANDED ALDER BEETLE IN NATURAL AND URBAN

ENVIRONMENTS (COLEOPTERA: CERAMBYCIDAE)

Rosalia funebris Motschulsky, the banded alder beetle, is a striking black and

white insect [illustrations in situ, see: Chemsak J. & E. G. Linsley. 1971; Pan-

Pacif. Entomol., 47: 149-154; Linsley, E. G. 1964. Univ. Calif. Publ. Entomol.,

22; Linsley, E. G., J. N. Knull & M. Statham. 1961. Amer. Mus. Novitatas, 2050],

in a genus noted for large size and conspicuous coloration. It was among the first

insects discovered and named from western North America. It was captured at

Sitka, Alaska, during the early Russian survey expeditions and named by the

Russian entomologist Motschulsky in 1845. It was not reported by American

entomologists until 12 years later [Le Conte, J. L. 1857. Vol. 12, Book 2, Part 3.

Zoological Rpt. 1, pp. 1-72. House of Representatives, 36th Congress, Ex. Doc

56.], when Le Conte recorded it from Oregon in an account of the insects collected

during the Missouri to Pacific railroad survey. The next American record [Le

Conte, J. L. 1869. Ann. Mag. Nat. Hist. (4)4: 369-385] was from Vancouver

Island. Subsequently, its known geographical range has been extended southward

to California [Casey, T. L. 1912. Memoirs on the Coleoptera, 3: 313] and south-

eastward in the southern Rocky Mountain region into Arizona (Linsley et al.,

1961) and New Mexico (Casey 1912). For a map of the presently known distri-

bution, see Linsley (1964).

Beyond the fact that adults gather in small aggregations, on freshly cut logs and

recently dead trees, for mating and oviposition during warm, mid-summer nights,

little is known of their life history. Their adult environment is the streamside

habitats where their host trees, usually alder (A/nus) but sometimes apparently

California laurel (Umbellularia), grow. Of special interest, then, is their observed

attraction to artificial environments provided by man. One such attractant is

paint. First reported [Essig, E. O. 1948. Pan-Pacif. Entomol., 19: 91-92] and based

on the collection of large numbers of both sexes attracted to a paint shop in Ukiah,

California, on a hot summer day, this unique behavior was confirmed by Chemsak

& Linsley (1971) at a prefabrication building plant 7.3 km (4.5 mi) north of Santa

Rosa, California. Again, both sexes were attracted to the site, where they were

drowned in empty paint cans or resting immobilized in the paint shed. When

captured and placed in plastic bags the sluggish ones regained their mobility and

became very active.

The second unique behavior pattern involves the aggregating of the sexes on

the walls of buildings in a dense urban environment. In response to reports

received in early July, 1970, Chemsak and Linsley visited a bank in Santa Rosa,

where the beetles were gathering and attracting local attention. Upon their arrival

at 11:05 h the beetles were resting and sluggish, but by noon the sun shone brightly

on the front of the building and the beetles moved into the shade. Bank officials

stated that the beetles had been coming over a period of 10 days to 2 weeks,

reaching a peak during a hot spell when the daytime temperatures rose to 38.8°

C (102°F). The building had been newly painted recently.

134 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

These observations may now be supplemented with others from Oregon, by

Hariana Chilstron, an exhibitor of live insects:

“First noted in the summer of 1969, was a single adult male resting con-

spicuously on the newly painted wall of a stucco or concrete building in a

city park in West Portland in a predominantly residential area a few blocks

southwest of the Willamette River. At the time the significance of the sighting

was not recognized. However, between 1980 and 1993, usually in late July

and early August, both sexes were repeatedly brought to the Washington Park

Insect Zoo by visitors and zoo personnel who were puzzled by the fact that

all were found on walls of buildings and other structures. Most were people

from southwest Portland or adjacent communities west of the Willamette

River, although the area has numerous creeks running through it with abun-

dant Alnus and Salix, which may have provided a source for the beetles.”

In all of these situations, the common denominators are: (1) buildings or other

man-made structures, and (2) in at least some cases the presence of fresh paint

(status of the others not known). This suggests that a volatile ingredient, capable

in the California examples of being carried over a very significant distance, was

acting as an attractant, because a careful search of both areas failed to reveal a

streamside site or a growth of trees of either of the known hosts. The volatile

material must mimic a specific sex pheromone because a host attractant would

bring other insects. Further, it must be a sex pheromone that attracts both sexes—

not unknown but not the usually observed case in which the female attracts the

male.

Finally, it has been suggested [Ross, E. S. 1993. Pacific Discovery, Winter 1993:

40-41] that these beetles may have served as a model (otherwise unknown) for

the Hopi Indian Katchinas known as hamo clowns. Ross (1993) illustrates both

the indian ceremonial dancers and the beetle and coins the name “banded alder

beetle,’’ which we have adopted. The resemblance of Rosalia to the Hopi Katch-

inas, although not previously reported, was noted independently by Celeste Green

(artist and entomological illustrator), Joan Quay (artist and scientist), and by E.

G. and J. M. Linsley, who over a period of 15 years assembled a collection of

these dolls (some eating watermelon after the dance, which gave them another

common name, “the glutton’’).

Acknowledgment. —I thank Hariana Chilstron for discussions and information.

E. Gorton Linsley Dept. of Entomological Sciences, University of California,

Berkeley, California 94720.

PAN-PACIFIC ENTOMOLOGIST

71(2): 135-136, (1995)

Scientific Note

HARMONIA AXYRIDIS (PALLAS)

(COLEOPTERA: COCCINELLIDAE), FIRST

WESTERN UNITED STATES RECORD FOR THIS

ASIATIC LADY BEETLE

Harmonia axyridis (Pallas) is a predator of aphids and scale insects and is

distributed from Siberia through China and Japan. It was introduced into Cali-

fornia in 1916, 1964, and 1965, and into Nova Scotia, ten eastern states, from

Maine to Mississippi and Washington, from 1978 to 1985 (Gordon, R. D. 1985.

J. New York Entomol. Soc. 93: 1-912; Chapin, J. B. & V. A. Brou. 1991. Proc.

Entomol. Soc. Wash. 93: 630-635). No recoveries were reported until those of

Chapin & Brou (1991) during 1988-1990 in Louisiana and Mississippi, and in

Georgia in 1991 (Gordon, R. D. & N. Vandenberg. 1991. Proc. Entomol. Soc.

Wash. 93: 845-846).

In June, 1993, one of us (SHD) collected lady beetles that KSH later identified

as Harmonia axyridis. At least four adults and two pupae were collected by hand

from an aphid-infested elm tree, probably Ul/mus americana L., in Vancouver,

Washington. Other uncollected specimens of apparently the same species were

observed on an ornamental Pyrus species about 1 km away in Vancouver. Two

H. axyridis were also collected on a Quercus sp., about 13 km from the first site,

in Portland, Oregon. Jeffrey C. Miller and Michael LaMana (personal commu-

nication), Oregon State University, Corvallis, first discovered H. axyridis at sites

in northwestern Oregon during the spring of 1993.

Since April, 1993, Kent M. Daane and KSH have been introducing into Cal-

ifornia, Harmonia axyridis received from W. L. Tedders at the University of

Georgia. Releases were on pecan, Carya illinoensis Koch, infested with yellow

pecan aphid, Monelliopsis pecanis (Bissell). Release dates (and number of adults

released) during 1993 were 9 Apr (1400), 26 Apr (700), and 15 May (1400) in

Visalia (Tulare Co.) and 26 Apr (20) in Clovis (Fresno Co.), California. In May,

1994, LGB collected two adults of H. axyridis from fennel, Foeniculum vulgare

Miller, outside of a greenhouse near downtown Sacramento (Sacramento Co.),

California. These beetles, along with Hippodamia convergens Guerin, were feeding

on aphids. In October, 1994, larvae, pupae, and adults were abundant on cotton,

Gossypium hirsutum L., at the same location. In September, 1994, SHD collected

one adult H. axyridis from a potted chrysanthemum, Dendranthema grandiflora

(Tzvelev), infested with melon aphids, Aphis gossypii Glover, in Davis (Yolo Co.),

California. During April and May, 1995, H. axyridis appeared to be common in

the Davis and Sacramento area; it was observed on aphid-infested Liriodendron,

Lycopersicon, Prunus, Quercus, Rosa and Ulmus sp.

In addition to its apparent economic importance as a biological control agent,

Harmonia axyridis is of evolutionary interest because of its color variability

(Ayala, F. J. 1978. Scient. Amer., 239: 56-69); more than 100 forms of the beetle

have reportedly been described. The lady beetles we collected in Oregon and

136 THE PAN-PACIFIC ENTOMOLOGIST Vol. 71(2)

Washington have light orange elytra with 19 black spots. They have been des-

ignated H. axyridis var. 19-signata and are most prevalent in southern Siberia

and southwestern China (Dobzhansky, Th. 1933. Amer. Nat., 67: 97-126). Spec-

imens collected in California include individuals with pale yellow to light orange

elytra and little or no dark markings, in addition to the 19-signata pattern. All

specimens exhibited the transverse subapical elytral ridge characteristic of this

species.

Material Examined.-CALIFORNIA. SACRAMENTO Co.: Sacramento, North B Street, 17 May

1994, Foeniculum vulgare, L. G. Bezark. Sacramento, North B Street, 3 Oct 1994, Gossypium hirsutum,

L. G. Bezark & J. Brown. YOLO Co.: Davis, Orchard Park Drive, 29 Sep 1994, Dendranthema

grandiflora, S. H. Dreistadt. OREGON. MULTNOMAH Co.: Portland, Hoyt Arboretum, Quercus

sp., S. H. Dreistadt. WASHINGTON. CLARK Co.: Fort Vancouver Historical Site, Vancouver, 28

Jun 1993, (prob.) Ulmus americana L.

Steve H. Dreistadt,! Kenneth S. Hagen,” and Larry G. Bezark,? ‘JPM Education

and Publications, Statewide IPM Project, University of California, Davis, Califor-

nia 95616-8620; *Laboratory of Biological Control, University of California,

Berkeley, California 94720; *Biological Control Program, California Department

of Food and Agriculture, Sacramento, California 95814.

PAN-PACIFIC ENTOMOLOGIST

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Literature Cited. — Format examples are:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

Blackman, R. L., P. A. Brown & V. 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 Essige/la (Homoptera: Aphididae). Pan-Pacif. Entomol.

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THE PAN-PACIFIC ENTOMOLOGIST

Volume 71 April 1995 Number 2

Contents

EDMUNDS, G. F., JR. & C. M. MURVOSH— Obituary: Richard K. Allen (1925-1992)

POLHEMUS, J. T.—A new genus of Hebridae from Chiapas amber (Heteroptera)

GULMAHAMAD, H.—The genus Liometopum Mayr (Hymenoptera: Formicidae) in Califor-

nia, with notes on nest architecture and structural importance

SAVARY, W. E.—Dacne picta Crotch: a recently introduced pest of stored, dried shiitake

mushrooms (Coleoptera: Erotylidae)

MONTLLOR, C. B., E. A. BERNAYS, J. HAMAI & M. GRAHAM~—Regional differences in

the distribution of the pyralid moth Uresiphita reversalis (Guenée) on French broom,

and introduced weed

GULMAHAMAD, H.—The desert dampwood termite (Isoptera: Kalotermitidae) as a structural

pest in the Colorado desert of southern California

ANDREWS, F. G.—An atypical new species of Corticarina from the Clarion Islands, Mexico

(Coleoptera: Lathridiidae: Corticariini)

SCHOLL, A., R.W. THORP, J. A. BISHOP & E. OBRECHT — The taxonomic status of Bombus

alboanalis Franklin and its relationship with other taxa of the subgenus Pyrobombus

from North America and Europe (Hymenoptera: Apidae)

RUST, R. W.—Adult overwinter mortality in Osmia lignaria propinqua Cresson (Hymenoptera:

Megachilidae)

NOOR, M. A.—Incipient sexual isolation in Drosophila pseudoobscura bogotana Ayala &

Dobzhansky (Diptera: Drosophilidae)

ALEXANDER, B. A.—Description of the female of Nomada dreisbachorum Moalif (Hyme-

noptera: Apoidea: Nomadinae)

SCIENTIFIC NOTES

LINSLEY, E. G.—The banded alder beetle in natural and urban environments (Coleoptera:

Cerambycidae)

DREISTADT, S. H., K.S. HAGEN & L. G. BEZARK — Harmonia axyridis (Pallas) (Coleoptera:

Coccinellidae), first western United States record for this Asiatic lady beetle

105

125

130