Vol. 54 JULY 1978 No. 3

THE

PAN-PACIFIC ENTOMOLOGIST

PAPP, R. — Ecology and Habitat Preferences of High Altitude Coleoptera from the

SiS thAING VAG arenas nee. Mn came y het, wuss ldob ans cee eats cel ete Wiad Stes a eee nmetieg aha rs 161

RENTZ — A New Ragirenogen etic Timema from California (Phasmatodea: Timemi-

CAG) Re ee ete ey ere eee a Mee Nc ae seo a Sew™ ahs cis May ci ares he gMa scott bat ox Afict/ eh OE te 173

ELLIS — Over-winter Occurrence and Maturation of Gonads in Adult Psychoglypha

subborealis (Banks) and Glyphopsyche irrorata (Fabricius) (Trichoptera:

imme DGS) Bae nok Sere ace ae Pet DYE Soca cts GR a ao 178

WILLIS — Eugene Graywood Smyth, Ph. D. (1886-1975): Obituary and Bibliography

OMANFEMLOMOIOGISK. «cscs tase ceteh See ue. s SeMay celles a sone ccc.) PRRs) Mates ale cece rye 181

HASSAN and KEIFER — The Mango Leaf-coating Mite, Cisaberoptus kenyae K.

(Eniophyidae, Aberoptinae),-. 2.8) oka. ba a Oh ee Set oe, ho ee 185

KRITSKY — The Neotropical Species of Oncylocotis (Hemiptera: Enicocephalidae).. 194

ELLIS — Seasonal Abundance and Distribution of Adult Caddisflies of Sashin

Creek, Baranof Island, Southeastern Alaska (Trichoptera)................. 199

JANSSON — A New Species of Graptocorixa from Mexico (Heteroptera, Corixidae).. 207

PAPP, C. — Corrections of Homonyms in Conotrachelus, and a Species List for

Pheloconus (Coleoptera: Curculionidae) ............ 2... eee ee eee eee eee 210

ALCOCK — Notes on Male Mate-locating Behavior in Some Bees and Wasps of

Arizona (Hymenoptera: Anthophoridae, Pompillidae, Sphecidae, Vespidae).. 216

PIPER — Life History of Zygogramma disrupta in Southeast Texas (Coleoptera:

ChinySOmeliGae)its;saten. hes cy kote cs atnemal s © ose b Beach dS REE ANN Ean ores OMe 226

MANLEY and TABER — A Mating Agoreoatlon of Dasymutilla foxi in Southern Ari-

zona (Hymenoptera: Mutillidae)......... 20.0... ce ee eee 231

WIRTH and BLANTON — Two New Species of Neotropical Culicoides (Diptera: Cera-

TOPO GOMIG AGC) hit eek ae cols nz age ces TT ctepe ee ans, 2 oe eee eee eco, Sereenenal sus 236

ZOOEOGICALNOMENGLATURE, «14% 02 A.cacm.o tkhapete ees. Sage oe ce ee 240

= GIIEINITIEUGIN ©] Bie ie Aare eee ce Mies ae ds tairatae ite.ce eee see Me ate dala vo hassy nia ge Mens 206

RECENTER ATURE Bete et mat Niet src ey. als api hap sete tans CUE oe ear Aedes 172; 214

BOOKIRIEVIEVVEs wat -ehrri ae te, ea akn ct sit etme. SAL NNEC gehepece's o abe. aueterste sacecte ebay tes 235

SAN FRANCISCO, CALIFORNIA ¢ 1978

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

THE PAN-PACIFIC ENTOMOLOGIST

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Title of Publication: The Pan-Pacific Entomologist.

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Publication #419440

The Pan-Pacific Entomologist

Vol. 54 JULY 1978 No. 3

Ecology and Habitat Preferences

of High Altitude Coleoptera from the Sierra Nevada’

Richard P. Papp

Dept. Entomological Sciences, University of California, Berkeley 94720?

Extensive pitfall trapping in the central Sierra Nevada in 1973-74

was conducted to obtain information on species composition, diver-

sity and associations for terricolous arthropods in five alpine and

subalpine plant communities. Over 1200 specimens comprising 37

species in 4 families of Coleoptera were collected. The information

presented here provides details of the location, habitat type and time

of capture for the 37 species of Coleoptera which were trapped.

Methods and Materials

Pitfall traps consisted of 4.5 x 4 cm glass jars filled to a depth of 1

cm with technical grade ethylene glycol. Traps were collected at 14-

day intervals and replaced by new traps. Ten stations of 3 traps each,

with traps set 1m apart in a line, were established in each of the 5

habitats sampled. Some supplemental collecting was done by hand

in areas adjacent to the pitfall trap sites. Voucher specimens of

insects collected are deposited in the collections of the author, and

at the University of Alberta (Edmonton), Biosystematics Research

Institute (Ottawa) and the California Academy of Sciences (San Fran-

CiSCO).

Study Sites

All of the trap sites were located in the Harvey Monroe Hall Area of

Inyo National Forest along the crest of the Sierra Nevada in Mono

County, California. The history, climate and plant affinities of the Hall

Area are described in Clausen (1969).

Details of the pitfall trap sites selection, vegetation analysis of the

plant communities sampled and arthropod species diversity analyses

may be found in Papp (1975). Brief descriptions of the 5 plant com-

munities follow. ;

‘Portion of Ph.D. thesis, Department of Entomological Sciences, University of California, Berkeley, 1975.

2Present Address: Department of Entomology, Bernice P. Bishop Museum, P.O. Box 6037, Honolulu, HI

96818.

The Pan-Pacific Entomologist 54:161-172. July 1978.

162 THE PAN-PACIFIC ENTOMOLOGIST

1. Lodgepole Pine Forest

Elevation 3048-3170 m. Located immediately NNW of the Carnegie

Institution Timberline Station (CITS) on a SE - facing slope. Contains

a considerable variety of understory subhabitats ranging from xeric,

almost pure stands of Pinus contorta Dougl. to hygric sites with under-

story vegetation dominated by Hordeum, Veratrum californicum

Durand and Senecio triangularis Hook. Vegetation in the drier sites is

diverse, with common species including Perideridia Parishii (Coult.

and Rose) Nels. and Macbr., Poa epilis Scribn. and Potentilla glandulosa

Lindl. Pitfall trap collections 11 July to 25 September, 1973 and 26

June to 18 September, 1974.

2. Sagebrush Community

Elevation 3078 m. Located on a gently sloping gravel area, facing

ESE and sloping to Slate Creek. Surrounded by scrubby lodgepole

pine forest. Dominant shrub is Artemesia cana Pursh, with other

common species including Carex, Horkelia fusca Lindl. ssp. capitata

(Lindl.) Keck and Sphenosciadium capitellatum Gray. Pitfall trap collec-

tions 17 July to 25 September, 1973 and 26 June to 18 September,

1974.

3. Subalpine Meadow

Elevation 3048 m. Located on the floor of Lee Vining Canyon, about

1500 m WNW of CITS. The meadow vegetation is dominated by Poly-

gonum bistortoides Pursh, and is bordered along Slate Creek by thick

clumps of Salix. Pitfall trap collections 17 July to 25 September, 1973

and 26 June to 18 September, 1974.

4. Alpine Meadow

Elevation 3353 m. Located on the E ridge of Mt. Conness, on site of

nival (Snow-surface) aeolian ecosystem investigations conducted by

Papp (1975, 1978). Dominant herbaceous vegetation includes Juncus

Parryi Engelm., Poa Hanseni Scribn., Antennaria alpina (L.) Gaertn. var.

media (Greene) Jepson and Lupinus. Pitfall trap collections 18 July to

26 September, 1973 and 11 July to 5 September, 1974.

5. Elfinwood Whitebark Pine Community

Elevation 3383 m. Located ESE of the alpine meadow on the SW

slope of the unnamed peak (3426 m) which forms the W border of

Saddlebag Lake. Pinus albicaulis Engelm. forms scattered thickets

over much of the slope. Various species of alpine grasses and

sedges, in addition to Ribes cereum, are found around the margins of

the pines. Frequent disturbance of pitfall traps by the alpine rabbit

Lepus townsendii in 1973 forced abandonment of trapping here. Pit-

fall trap collections 18 July to 26 September, 1973; no traps set in

1974.

VOL. 54, NO. 3, JULY 1978 163

Results

Distributional and ecological information for Coleoptera was ob-

tained either from the literature, or from available museum specimen

data. Distributions of some Carabidae, Elateridae and Staphylinidae

were based on collection data in the Department of Entomology, Cali-

fornia Academy of Sciences (CAS), and the California Insect Survey

(CIS), University of California, Berkeley. Ecological information from

the literature is given for each species, followed by additional (and

sometimes contradictory) information obtained in the course of this

study. The following species were previously unrecorded from the

conterminous United States: Dyschirius alticola Lindroth, D. truncatus

Lindroth (Carabidae); Limonius pappi Becker (Elateridae). The follow-

ing species were recorded for the first time from California: Amara

erratica Duftschmid, Bembidion complanulum Mannerheim, B. improvi-

dens Casey, B. quadrifoveolatum Mannerheim, Cymindis unicolor Kirby,

Dyschirius alticola Lindroth, D. truncatus Lindroth, Harpalus seclusus

Casey (Carabidae); Boletobius pygmaeus Fabricius, Mycetoporus

consors LeConte (Staphylinidae); Limonius pappi Becker (Elateridae).

Carabidae

Agonum fallianum Leng

Distribution: California (Lindroth, 1966)

Ecology: Previously unknown. All of the specimens (42) from the Hall Area were trapped

in meadows; 29 specimens (69%) in alpine meadow and 13 specimens (31%) in subalpine

meadow. In addition, 14 specimens were collected from the snow surface on Mt. Conness

in the alpine zone. This species forages on the snow in both alpine and subalpine areas

during the afternoon, but disappears at dusk. Of the specimens from alpine meadow

stations, 33 (79%) were not trapped until late September; all but one of the remaining

specimens trapped in this habitat were captured immediately following snow melt. It

appears that this species either diapauses as an adult during most of the snow-free

period of the summer, or spends much of the summer as an immature. A mating pair of

this species was observed on June 22, 1974 on wet soil near the sagebrush area.

Amara erratica Duftschmid

Distribution: Circumpolar; in North America transcontinental, south to New England in

the east and Mt. Rainier, Washington, and Colorado in the west (Lindroth, 1968); may also

occur in Oregon and Idaho (Hatch, 1953); boreoalpine in Europe and Asia, occurring in the

Caucasus, and in Siberia east to Kamchatka (Lindroth, 1954). Mani (1968) also records this

species from the Alps, the Pyrenees, Lapland and Iceland.

Ecology: lonescu (1969) trapped this species in five alpine stations in Romania at alti-

tudes of 1800-2035 m, with most specimens taken from May to mid-July; occurs in open

country with grass or meadow vegetation, usually on sandy moraine, on both sides of the

timber limit (Lindroth, 1968). Conspicuously xerophilic, on sandy moraines in the Betula-

zone up to 1000 min Lapland and 1200 m in Iceland (Mani, 1968).

Most of the 26 Hall Area specimens were trapped in meadow situations (alpine or sub-

alpine). One individual was taken in the lodgepole pine forest in late September and four

others were trapped in the sagebrush area in early August. Additionally, two specimens

were caught foraging on the snow on Mt. Conness on July 12, 1973.

164 THE PAN-PACIFIC ENTOMOLOGIST

Anisodactylus pitychrous LeConte

Distribution: A species of the dry interior, in Canada occurring only east (sic?) of the

Rockies, south to Colorado and California (Lindroth, 1968).

Ecology: Previously unknown. Sixty-eight specimens were taken in the Hall Area, of

which 88% occurred in the moist subalpine meadow. Five were recovered from forests, 3

from lodgepole and 2 from elfinwood whitebark pine; 3 specimens were trapped in sage-

brush. Within the subalpine meadow, 73% were from marginal, relatively well-drained

stations adjacent to young lodgepole pines.

Bembidion commotum Casey

Distribution: In western mountains, from Alberta and British Columbia south to Sierra

Nevada of California, east to Wyoming (Lindroth, 1963).

Ecology: Apparently restricted to high mountains; in Sonora Pass, California, recorded

from gravelly border of a small creek below timber limit, September 3, 1961 (Lindroth,

1963).

In the Hall Area, specimens were trapped in lodgepole pine forest, sagebrush and

alpine meadow, but 76% were taken in alpine traps, with the greatest number (41%) taken

immediately after snow-melt.

Bembidion complanulum Mannerheim

Distribution: In western mountains from Alberta, British Columbia and Alaska, south to

northern Oregon (Lindroth, 1963).

Ecology: Taken at margin of snow patches in Waterton Park, Alberta, Chilkot Pass,

British Columbia, and on Unalaska Island; apparently not riparian; normally imaginal

hibernation (Lindroth, 1963).

Collected in alpine meadow, subalpine meadow, and lodgepole forest in the Hall Area.

Two specimens were taken on snowfields in late June 1972 and 1973. Most of the speci-

mens (77%) were taken in the alpine meadow, with catches quite evenly distributed over

the season. This species is a facultative nival predator in alpine areas.

Bemibidion humboldtense (Blaisdell)

Distribution: Northern California, southwestern Oregon (Lindroth, 1963).

Ecology: Previously unknown. A single specimen was taken in the alpine meadow soon

after snow melt in late July, 1973.

Bembidion improvidens Casey

Distribution: Mt. Rainier (above 4000 ft. elevation, Washington) (Lindroth, 1963).

Ecology: Previously unknown. Occurs sparsely only in the alpine meadow, where it was

trapped at four stations; none of the alpine meadow stations differ substantially from

each other with respect to exposure, plant cover or soil type.

Bembidion obscuripenne Blaisdell

Distribution: In western mountains from Washington south to California (Lindroth,

1963).

Ecology: In the alpine and subalpine regions (Lindroth, 1963). In the Hall Area, a single

specimen was trapped in the alpine meadow soon after snow melt in late July 1973; at this

time this area was noticeably wetter than other trap stations in the alpine meadow.

Bembidion quadrifoveolatum Mannerheim

Distribution: In the western mountains and along the coast south to Oregon (Hatch,

1953).

Ecology: Strictly riparian, occurring among stones and gravel on barren banks of

usually small streams. Above timber limit in Chilkot Pass, British Columbia; hibernation

apparently in both larval and adult stages (Lindroth, 1963).

VOL. 54, NO. 3, JULY 1978 165

This is a widespread species in the Hall Area. Although most common in the alpine

meadow, where 70% were taken, it also occurs in elfinwood whitebark pine stands, sub-

alpine meadow and in Pinus contorta Doug]. forest. Of those specimens taken above the

timberline, 54% were captured within the first two weeks following snow melt. Those

specimens trapped in the lodgepole pine forest were taken much later, with 60% captured

during the last trapping period in late September. It seems probable that this species is

active throughout most of the snow-free period of the year, moving to sheltered locations

at lower altitudes during winter.

Specimens of B. quadrifoveolatum were taken on snowfields in late June and early July

in 1972 and 1973, where it is a facultative nival predator. Another specimen was taken near

the Hall Area in Glacier Canyon on the northwest side of Mt. Dana at an altitude of 3200 m

on May 18, 1968.

Bembidion timidum LeConte

Distribution: Interior areas of western Ontario, Manitoba, Saskatchewan, Alberta,

Northwest Territories and British Columbia south to Oregon, California and Colorado

(Lindroth, 1963).

Ecology: Occurs in a diversity of moist habitats often with Bembidion versicolor;

hibernates as an adult (Lindroth, 1963).

Five specimens were taken in the Hall Area, two in alpine meadow (one immediately

after snow melt and the other in late September, 1973), two in sagebrush during late

August and early September, 1973, and one in subalpine meadow during late August

1973. This is the only species of Bembidion which was trapped in the sagebrush com-

munity, which is drier than typical Bembidion habitats. B. timidum does not seem to occur

in forest habitats in the Hall Area, despite its broad habitat occurrence in other areas.

Cymindis unicolor Kirby

Distribution: Transamerican, from Canada and Alaska south along the mountains to

Colorado and Utah and isolated on New England mountains (Lindroth, 1969).

Ecology: In treeless country, above timberline and on tundra (Lindroth, 1969). Of the 70

Hall Area specimens 57% were trapped in alpine meadow stations; twenty were also

taken in sagebrush, and ten in subalpine meadow. None were trapped in forest habitats.

Dyschirius alticola Lindroth

Distribution: Previously known only from type locality: Canada, British Columbia; the

eastern of the two passpoints on Rossland Trail (10 miles west of trail) about 5000 ft. (1524

m), May 21, 1958 (Lindroth, 1961).

Ecology: On open, slightly moist, fine sandy soil, half shaded by A/nus bushes, at the

margin of mixed forest, with almost no vegetation, except in spots of the moss (Ceratodon

purpureus). Associated with Bembidion dyschirinum (common) and Notiophilus simulator

(Lindroth, 1961).

In the Hall Area this species was recorded only in the subalpine meadow, where the soil

is very finely textured and vegetative cover is almost complete. In 1973 most individuals

- (58%) were trapped during the last two weeks of August when the meadow was becoming

quite dry. In 1974 most (57%) were caught within a few days of snow melt in late June.

Dyschirius truncatus Lindroth

Distribution: Widely distributed but apparently everywhere rare, probably lacking in the

Atlantic region. There are records from Manitoba, Saskatchewan, Alberta, Northwest Ter-

ritories, British Columbia and Alaska (Lindroth, 1961).

Ecology: On barren clay-mixed sand at the margins of standing or slowly running

waters; probably constantly in company with Bledius (Lindroth, 1961).

A single specimen was taken in the Hall Area on August, 1973 in the lodgepole pine

forest. The microhabitat at this station was as follows: traps set in wet soil under shelter

of a large clump of Salix on a south-east-facing slope; also common at this station were

clumps of Veratrum californicum Durand, Allium validum Watson, Carex and mosses. The

166 THE PAN-PACIFIC ENTOMOLOGIST

station lies at the base of a large granite outcrop and received abundant snow melt runoff

throughout the summer.

Harpalus seclusus Casey

Distribution: From Alberta and British Columbia southward in the mountains to

Colorado (Lindroth, 1968).

Ecology: On meadows at high latitudes; specimens from Alberta were captured at the

margin of a snowfield about timber limit; in Manitoba it is associated with Harpalus

animosus (Lindroth, 1968).

A very common species in the Hall Area; 51% of the 285 specimens trapped were from

meadows, either alpine or subalpine. A few were taken in lodgepole pine forest and one

was trapped in the elfinwood whitebark pine community, but the vast majority came from

the sagebrush community, where 81% were trapped. This habitat is exceptionally well-

drained, with a gravel substrate which provides for warmer surface temperatures than in

the other communities sampled (Papp, 1975). Since many harpaline Carabidae are now

known to be phytophagous (e.g. Johnson and Cameron, 1969; Zhavoronkova, 1969;

Kabcik-Wasylik, 1971; Kirk, 1972) itis probable that this species is a seed-cacher.

Lebia viridis Say

Distribution: Transcontinental, entire United States south to Mexico; the most wide-

spread and abundant member of the genus (Lindroth, 1969).

Ecology: On open, sun-exposed substrates, often in goldenrod flowers or resting on

leaves of A/nus or U/mus in company with more abundant Altica chrysomelids of the same

size and color (Lindroth, 1969). Madge (1967) has shown that the larvae of Lebia viridis are

parasitoids on the Chrysomelidae, and suggests that Lebia mimics its host Altica.

Four specimens were collected, all in 1973: 3 in subalpine meadow and 1 in the lodge-

pole pine forest. Interestingly, many specimens of Al/tica torquata LeConte were also

trapped in the Hall Area (Papp, 1975).

Nebria ovipennis LeConte

Distribution: Sierra Nevada, from Sequoia National Park north to Yuba Pass

(Kavanaugh, pers. comm., 1978).

Ecology: Previously unknown. Four specimens were collected in 1973. A submature

male was found foraging on snow at 3353 m on Mt. Conness, on 9 July. A teneral male was

trapped in the sagebrush community and a mature male in the subalpine meadow, both in

late August. A mature female was also trapped in the subalpine meadow in mid-Septem-

ber.

Nebria spatulata Van Dyke

Distribution: Sierra Nevada, from Sequoia National Park north to Yosemite National

Park (Kavanaugh, pers. comm., 1978).

Ecology: Previously unknown. Four males and five females were trapped all in the

alpine meadow, in 1973. A teneral male was taken in mid-September, and a submature

female in early August. Other specimens, all of which were fully mature, were captured

from 15 July until 26 September.

Pterostichus inanis Horn

Distribution: The Cascade Range from British Columbia through Washington and

Oregon, into northern Nevada and the Sierra Nevada of California; (Hacker, 1968).

Ecology: Altitudinal range of this species is from 3000 ft. (914 m) in northern Oregon to

9400 ft. (2865 m) in Fresno County, California (Hacker, 1968).

All specimens in the Hall Area were trapped in lodgepole pine forest; most of these

were taken early in the season. Fourteen specimens (88%) were trapped from a single

station which was located adjacent to Cabin Creek, on a south-southeast facing slope.

Most abundant vegetation here was Phyllodoce breweri (Gray) Heller, Solidago and some

grasses, but all of these were present at some other stations where no P. inanis were

VOL. 54, NO. 3, JULY 1978 167

taken; its presence may therefore be restricted to forested situations near running water,

rather than to a specific plant association.

Pterostichus protractus LeConte

Distribution: In the Rocky Mountains of Canada, from Alberta and British Columbia,

south to the southern Cascades and into California (Lindroth, 1966).

Ecology: Apparently confined to montane forests; generally imaginal hibernation

(Lindroth, 1966). 66% of the 133 specimens were trapped in forested situations (71 in

lodgepole pine and 17 in elfinwood whitebark pine). This species also occurred in all of

the other habitats sampled: subalpine meadow (18), sagebrush (25) and alpine meadow

(2). Two hand collected specimens from the lodgepole pine forest were found near dead

trees, one under a rotten log and the other in the soil at the base of a standing dead tree.

The species shows a definite preference for dry niches with mature trees and abundant

pine needle litter: 4 of the lodgepole forest stations which were of this type accounted for

88% of the specimens taken.

Trachypachus gibbsi LeConte

Distribution: From southernmost British Columbia to the Southern Sierra Nevada of

California (Lindroth, 1961).

Ecology: Stenotopic, restricted to a special habitat on banks of big rivers at some

distance from the water under Populus balsamifera. The soil, consists of finest sand, is

dust dry on surface and is covered by an almost continuous layer of dead leaves.

Equisetum arvense was present in one place, otherwise no vegetation. This habitat is

doubtless flooded in the spring (Lindroth, 1961).

This species was trapped exclusively in the Pinus contorta forest in both 1973 and 1974.

The majority of the specimens were trapped in the first period following snow melt (1973:

65%, 1974: 76%), while the soil was still moist. Litter was present as pine needles and

cones, together with leaves of Salix, Senecio, Veratrum and Lupinus.

Cicindelidae

Cicindela longilabris Say

Distribution: Transamerican, from Alaska to Newfoundland, south into California, New

Mexico, Colorado, Wisconsin, Illinois and New York. (Leng, 1920).

Ecology: Of the 41 specimens taken in the Hall Area, 38 (93%) were captured in the

sagebrush community. The other specimens were taken in the subalpine meadow and in

the lodgepole pine forest. Temporal distribution was bimodal. In both years most speci-

mens were taken either soon after snow melt, or during the last trap period in September.

Cicindela oregona LeConte

Distribution: Alaska, south through British Columbia into California, Nevada, New

Mexico and east to Montana, Utah and Idaho (Leng, 1920).

Ecology: Guppy (1948) collected this species on bare gravel below the winter tide lines

on Vancouver Island. Twelve specimens were captured in the Hall Area: sagebrush (6),

subalpine meadow (5), and lodgepole pine forest (1). As with C. /ongilabris, this species

was trapped only in either the first or the last trap period of each year.

Staphylinidae

Aleochara affluens Casey

Distribution: British Columbia, northwestern Washington, northern Idaho and western

Oregon (Hatch, 1957).

Ecology: According to Hatch (1957), all members of this genus probably live as ecto-

168 THE PAN-PACIFIC ENTOMOLOGIST

parasites on pupae of cyclorraphous Diptera.

In the Hall Area, 6 specimens were taken crawling on snow on the east ridge of Mt.

Conness, 3353 m elevation, on June 25, 1973; in addition, 2 were taken in the alpine

meadow in early September 1973 and 3 were taken in the subalpine meadow in early

August and mid-September 1973.

Boletobius pygmaeus Fabricius

Distribution: British Columbia, northwest Washington, southeastern Idaho and

northern Oregon (Hatch, 1957).

Ecology: Previously unknown. Four specimens were taken in the Hall Area, all in lodge-

pole pine forest, from mid-July until late August, 1973.

Mycetoporus consors LeConte

Distribution: British Columbia, Washington and western Oregon (Hatch, 1957). New York,

Michigan, Indiana (Moore and Legner, 1975).

Ecology: Previously unknown. Three specimens were taken in the Hall Area: 1 from the

elfinwood whitebark pine habitat in late July, 1973; 2 from subalpine meadow, late July

and early September, 1973.

Platystethus americanus Erichson

Distribution: Common throughout the United States and in Mexico (Moore and Legner,

1971). Quebec and British Columbia (Moore and Legner, 1975).

Ecology: Common in manure and in isolated field droppings; occasionally in other de-

caying organic matter and at lights (Moore and Legner, 19771).

In the Hall Area, 1 specimen was taken in the subalpine meadow in late July, 1973; 6

others were found foraging on the snow on Mt. Conness, at 3353 m elevation.

Quedius breviceps (Casey)

Distribution: In western mountains from Alaska through British Columbia, Washington,

Oregon to southern Sierra Nevada (Mineral King) in California (Smetana, 1971).

Ecology: Specimens from Valdez, Alaska, were taken beneath an A/nus bush at a creek.

In the southern part of its range it occurs in wet biotopes in moss, under fallen leaves and

often near water; in high mountain elevations (alpine and subalpine zones) it occurs in

moss, lichens, grass patches, under stones, etc., and often in rather dry biotopes

(Smetana, 1971).

Of the 85 specimens collected in the Hall Area, all but one were taken in lodgepole pine

forest traps. Most of the specimens (58%) were taken soon after snow melt in late July.

No subhabitat preferences within the lodgepole forest were evident.

Quedius validus Smetana

Distribution: From type materia! only: holotype male, California: Stanford University,

May 13, 1950, PSB; allotype female, California: Grassy Lake, September 27, 1914, Lassen

National Forest, Lassen County, ex R. Hopping collection, California Academy of

Sciences (Smetana, 1971).

Ecology: No details of habitats known; a related species, Q. explanatus LeConte, occurs

under stones; development may be in mammal burrows (Smetana, 1971).

Eight specimens of this distinctive species were collected in the subalpine meadow

and in the lodgepole pine forest, in August and September of both years. As this is evi-

dently a boreal species, the data for the male holotype is probably erroneous, and the

designation of that specimen as the holotype is unfortunate.

Tachyporus californicus Horn

Distribution: From southern British Columbia, south into Wyoming, Idaho and Oregon

(Blackwelder, 1936); also in California (Fall, 1901).

Ecology: Members of this genus are commonly taken sweeping herbage (Blackwelder,

VOL. 54, NO. 3, JULY 1978 169

1936).

A single specimen was taken crawling on snow on the east ridge of Mt. Conness, 3353

m elevation, on July 5, 1973.

/

Elateridae

Ampedus phelpsi (Horn)

Distribution: From Alaska, across most of Canada, and south into New York (Mt.

Marcy), New Hampshire (Mt. Washington), Michigan, Wisconsin and Minnesota; in the

west it occurs in the Rockies south to the Parowan Mountains in Utah, and in the Sierra

Nevada south to Bubbs Creek Canyon on the Kings River.

Ecology: Previously unknown. A single specimen was collected beneath the bark of a

dead whitebark pine (Pinus albicaulis) near the Carnegie Institution Timberline Station on

June 17, 1973.

Ctenicera aeripennis (Kirby)

Distribution: From Alaska, south and east through the Yukon, Northwest Territories,

Alberta and British Columbia, along the mountains into Washington, Oregon and Cali-

fornia; also in the Rocky Mountains through Montana, South Dakota, Wyoming, Colorado,

Utah and New Mexico.

Ecology: Previously unknown. A single specimen was taken in the sagebrush

community in early August, 1973.

Ctenicera candezei (Leng)

Distribution: In the Sierra Nevada of California, from the Lake Tahoe area south into

Madera County (Lake Ediza).

Ecology: Previously unknown. Three specimens were collected, all crawling on the

snow on the east ridge of Mt. Conness, elevation 3353 m; one was taken on June 21 and

the others on July 12, 1973.

Ctenicera edwardsi (Horn)

Distribution: From localities around Vernon, British Columbia south through

Washington and Oregon, and into California as far south as Marin County along the coast

and Fresno County (Lower Hot Springs) in the Sierra Nevada; there are additional records

from what appears to be an isolated distribution on Mt. Timpanogos in north central Utah.

Ecology: One specimen was taken in the Hall Areain the subalpine meadow during late

July, 1973. Also recorded from 1981 min Nevada Co., California, on Juniperus.

Ctenicera tahoensis Van Dyke

Distribution: In the high Sierra Nevada of California from Placer County south to Lake

Alpine in Alpine County.

Ecology: Previously unknown. In the Hall Area, 2 specimens were captured while crawl-

ing on snow, one on the east ridge of Mt. Conness at 3353 m elevation on June 28, 1972,

and the other near Spuler Lake, elevation 3024-3194 m, 0815-0845 hours. Five others were

taken in pitfall traps, as follows: Pinus contorta Dougl. forest, late July, 1973 (2 specimens)

and late July-early August, 1974; subalpine meadow, late July, 1973; alpine meadow, late

July, 1973. BOF

Hypolithus bicolor Esch.

Distribution: A common species transcontinental in Canada below treeline and extend-

ing into the northern United States along mountain chains (Becker, 1978, pers. comm.).

Ecology: Previously unknown. Of the 176 specimens captured in the Hall Area, 97%

were taken in meadow situations: subalpine meadow, 99 specimens (56%), alpine

meadow, 71 specimens (40%). Becker (1978, pers. comm.) reported that all specimens

170 THE PAN-PACIFIC ENTOMOLOGIST

examined from the Hall area were wingless females, and that parthenogenesis is known

in this species.

Limonius nitidulus Horn

Distribution: From southern British Columbia and southwestern Alberta south to the

high mountains of California (Becker, 1978, pers. comm.).

Ecology: Apparently unknown. Three specimens were captured crawling on the snow

on the east ridge of Mt. Conness, elevation 3353 m, on June 21, June 25, and July 2, 1973.

One specimen was taken in the Artemesia community in late July, 1973, and two others

were trapped in the Pinus albicaulis habitat in late July, 1973.

Limonius pappi Becker

Distribution: Recently described from material collected in the Harvey Monroe Hall

Natural Area, Inyo National Forest, Mono County, California (Becker, 1976).

Ecology: The most commonly collected elaterid in the Hall Area, where 226 specimens

were trapped. All but 4 of these were taken in meadow habitats: 196 specimens (87%)

from the alpine meadow, and 26 specimens (12%) from the subalpine meadow. A few

additional specimens were collected in pitfall traps under elfinwood whitebark pine (Pinus

albicaulis), on foliage of P. albicaulis, and on snowfields on the E ridge of Mt. Conness,

3353.m elevation.

Discussion

Most of the 37 species treated herein represent extensions of

widespread boreoalpine species into southern latitudes. For

example, in the Carabidae 13 species including Amara erratica, Aniso-

dactylus pitychorus, Bembidion commotum, B. complanulum, B. obscur-

ripenne, B. quadrifoveolatum, B. timidum, Cymindis unicolor, Dyschirius

truncatus, Harpalus seclusus, Pterosticus inanis, P. protractus and

Trachypachus gibbsi are reported from other boreoalpine localities. Of

these only Amara erratica occurs outside of North America. The

cicindellid species Cicindela longilabris is also boreoalpine. Of the

Staphylinidae, 6 of the 7 species collected are broadly boreoalpine,

as are 7 of the 8 elaterid species collected.

Five boreoalpine species, including the carabids Agonum fallianum,

Bembidion humboltense, B. improvidens and Dyschirius alticola and the

elaterid Limonius pappi have rather limited distributions, in some

cases restricted to type localities, and probably reflect the formation

of disjunct islands during Pleistocene glaciations.

Acknowledgements

| wish to thank the following systematists for identifying material

from the Harvey Monroe Hall Natural Area: G.E. Ball, University of

Alberta, Edmonton (Carabidae); E.C. Becker, Biosystematics

Research Institute, Ottawa (Elateridae); JM. Campbell, Biosystem-

atics Research Institute, Ottawa (Staphylinidae); K.S. Hagen,

University of California Berkeley (Cicindelidae); J.T. Howell,

California Academy of Sciences, San Francisco (vascular plants); D.

Kavanaugh, California Academy of Sciences, San Francisco

VOL. 54, NO. 3, JULY 1978 171

(Carabidae: Nebria); A. Smetana, Biosystematics Research Institute,

Ottawa (Staphylinidae: Quedius); J. Strother, Jepson Herbarium,

University of California, Berkeley (vascular plants).

| also wish to thank: K.S. Hagen, F.A. Pitelka and E.I. Schlinger for

critical reading of my dissertation manuscript, of which this paper

formed a part; the Carnegie Institution of Washington, for use of the

Timberline facilities; the Mono Ranger District, Inyo National Forest,

for permission to conduct research in the Hall Area; and the Depart-

ments of Entomology, California Academy of Sciences and University

of California Berkeley, for permission to examine Coleoptera

collections.

Literature Cited

Becker, E.C. 1976. A new species of Limonius (Coleoptera: Elateridae) from California.

Can. Entomol. 108:689-692.

Blackwelder, R.E. 1936. Revision of the North American beetles of the subfamily Tachy-

porinae. Part |. Genus Tachyporus Grav. Proc. USNM 84:39-54.

Clausen, J. 1969. The Harvey Monroe Hall Natural Area. Carnegie Inst. Wash. Dept. Plant

Biol. Publ.no. 459,48 pp.

Fall, H.C. 1901. List of the Coleoptera of southern California. Occ. Papers Calif. Acad.

Sci. 8:282 pp.

Guppy, R. 1948. A list of Coleoptera collected near Lagoon Head, Wellington, V.I Coleopt.

Bull. 2(3):21-23.

Hacker, H.F. 1968. The species of the subgenus Leptoferonia Casey (Coleoptera: Cara-

bidae: Pterostichus). Proc. USNM 124(3649):1-16.

Hatch, M.H. 1953. The beetles of the Pacific Northwest. Part |. Introduction and Ade-

phaga. Univ. Wash. Publ. Biol. 16:1-340.

Hatch, M.H. 1957. The beetles of the Pacific Northwest. Part 2. Staphyliniformia. Univ.

Wash. Publ. Biol. 16:1-384.

lonescu, M. 1969. Observatii cu privire la entomofauna din zona alpina a Muntilor Cibin.

Stud. Ceret. Biol., Ser. Zool. 24:347-384.

Jeannel, R. 1940. Carabidae: Trechinae. British Museum (Nat. Hist.) Ruwenzori Expedi-

tion 1934-1935, 3:123-127.

Johnson, N.E. and R. Cameron. 1969. Phytophagous ground beetles. Ann. Entomol. Soc.

Amer. 62(4):909-914.

Kabacik-Wasylik, D. 1971. Studies on the diet of three field species of Carabidae. Ekol.

Polska 19(33):501-508.

Kirk, V.M. 1972. Seed-cacheing by larvae of two ground beetles, Harpa/us pennsylvanicus

and H. erraticus. Ann. Entomol. Soc. Amer. 65(6):1426-1428.

Leng, C.W. 1920. Catalogue of the Coleoptera of America, North of Mexico, Jonn D. Sher-

man, Jr. Mount Vernon, New York, 444 pp.

Lindroth, C.H. 1954. Carabidae common to Europe and North America. Coleopt. Bull.

8(2):35-52.

Lindroth, C.H. 1961-1969. The ground beetles of Canada and Alaska. Parts 1-6. Opuscula

Entomologica Supplementa XX, XXIV, XXIX, XXXII, XXXIV, XXXV.

Madge, R.B. 1967. A revision of the genus Lebia Latreille in America north of Mexico

(Coleoptera: Carabidae). Quaest. Entomol. 3:139-242.

Mani, M.S. 1968. Ecology and Biogeography of High Altitude Insects. W. Junk N.V. Pub-

lishers, The Hague, 527 pp.

Moore, |. and E.F. Legner, 1971. A review of the nearctic species of Platystethus (Coleop-

tera: Staphylinidae). Pan-Pacific Entomol. 47(4):260-264.

172 THE PAN-PACIFIC ENTOMOLOGIST

Moore, |. and E.F. Legner, 1975. A Catalogue of the Staphylinidae of America North of

Mexico (Coleoptera). Special Publ. 3015, Univ. Calif. Div. Agr. Sciences, 514

Pp.

Papp, R.P. 1975. Ecological interrelations among arthropods in some high altitude com-

munities in the central Sierra Nevada of California. Ph.D. Dissertation, Univ. of

California, Berkeley, 158 pp.

Papp, R.P. 1978. A nival aeolian ecosystem in California. Arctic and Alpine Res., 10(1):

117-131.

Smetana, A. 1971. Revision of the tribe Quediini of America north of Mexico (Coleoptera:

Staphylinidae). Mem. Entomol. Soc. Canada 79, 303 pp.

Yablakov-Khnzoryan, S.M. 1961. Opit vosstanovlieniya genezisa faunii zhestkokriilikh

Aremnii. Akad. Nauk Armyanskoi SSR. Zool. Inst., Erivan, 264 pp.

Zhavoronkova, T.N. 1969. Certain structural peculiarities of the Carabidae (Coleoptera)

in relation to their feeding habits. Entomol. Obozr. 48(4):729-739.

RECENT LITERATURE

The. Atlas of Insect and Plant Viruses including Mycoplasmaviruses

and Viroids. Karl! Maramorosch Ed. 1977. Academic Press,

N.Y. 512 pp. $49.50.

Perspectives in Forest Entomology. J. F. Anderson and H. K. Kaya

Eds. 1976. Academic Press, N.Y. 438 pp. $19.25.

Classification of the Higher Categories of the New World Tree-

hoppers (Homoptera: Membracidae). L. L. Deitz. 1975. Bull.

N. Carolina Agric. Exp. Sta., No. 225. 177 pp. $2.50.

Taxonomic Study of the Cicadellinae (Homoptera: Cicadellidae) Part

2. New World Cicadellini and the Genus Cicadella. D. A.

Young. 1977. Bull. N. Carolina Agric. Exp. Sta., No. 239. 1,135

pp. $15.00.

A New Parthenogenetic Timema from California

(Phasmatodea: Timemidae)

D.C.F. Rentz

Division of Entomology, CSIRO, Canberra, Australia

The timemas are an endemic western family. Species have been

described from Arizona, Nevada and California. California claims the

greatest number of described species with five now known. Timemas

are of interest because of their unique robust appearance and the

presence of distinctive asymmetrical genitalia in the males. Species

are uSually locally abundant and appear as adults from mid to late

spring. They are most often associated with chaparral plants and can

be found on the most pungent of shrubs. Most species emit an acrid

odor when handled. Such an odor is not associated with the species

herein described. The taxonomy of the Timemidae was reviewed

most recently by Tinkham (1942) and later Strohecker (1966) added

two species. The total number of known species now stands at 7.

Parthenogenetic phasmids are not unknown, (see White 1973). To

date only a single parthenogenetic phasmatid has been recorded

from the United States, but this is the first purported case in the

Timemidae. Such a conclusion was achieved by two collectors in-

dependently. Mr. H.B. Leech, California Academy of Sciences

(retired) wrote the author in 1971 suggesting the possibility after

studying a large collection of nymphs and adults which he had made.

Subsequent trips to the same location yielded additional specimens,

but there were no males. Concurrently, Ms. Genevieve Priain and Dr.

C.D. MacNeill of the Oakland Museum discovered the species at

Arroyo Mocho and began extensive studies to determine if any males

could be located. They reared individuals from eggs of captive

females and discovered no males in the progeny. Continuous collect-

ing each season from 1973 through 1977 has failed to uncover a

single male. In all other species of timemas known to this author,

adult females are encountered in the field with males atop, not

necessarily in copulation. Males are seldom found singly. It would

seem that if males were. present in the populations examined here,

they would have appeared by now. Adults have been reared by Ms.

Priain from adult females collected as immatures during the previous

season. Therefore, in any case, males are not necessary in these

‘populations to produce subsequent generations.

This species is being described at the request of the scientists at

the Oakland Museum and because of its potential interest to

geneticists due to its parthenogenicity.

The Pan-Pacific Entomologist 54:173-177. July 1978.

174 THE PAN-PACIFIC ENTOMOLOGIST

Figure 1. Timema genevievae Rentz, new species on chamise.

Timema genevievae, new species

HOLOTYPE FEMALE. California: Alameda County: Arroyo Mocho, 29 May 1974. C.D.

MacNeill, R.E. Love collectors. Holotype in California Academy of Sciences, number

13025. Type series taken mostly on chamise, Adenostoma fasciculatum H. & A. and

mountain mahagany Cercocarpus betuloides (Hook.) Nutt.

Description.—Form normal for genus. Head broad and flat, dorso-ventrally flattened;

eyes prominent, globular, much more produced than in any other known species.

Antenna with 22 articles, positioned well in front of eyes; scape very elongate, four times

longer than basal width, somewhat constricted basally; pedicel hardly distinguishable

from succeeding antennal articles; internal margins of antennal sockets produced

between eyes. Thorax; prothorax 1.65 times broader than long, sides straight, anterior

margin somewhat produced (Fig. 2), posterior margin concave; mesothorax slightly

smaller than prothorax, sides convex, posterior margin more concave, anterior margin

indistinct; mesothorax with lateral margin more strongly convex. Legs stout, undif-

ferentiated, unspined. Abdomen narrowing apically, not modified. Supra-anal plate rather

elongate, lying just anterior to middle of cercus, without central sinuation as seen in T.

boharti Tinkham; tenth tergite produced, emarginate. Subgenital plate (Fig. 3) constricted

basally, lateral margins obtusely produced mesad, converging to rather blunt apex.

Cercus elongate (Fig. 5), apices subacute, internal dorsal and ventral margins minutely

serrate. Ovipositor completely concealed by subgenital plate, valvulae strongly recurved

in apical portion, aciculate; dorsal valves longer, more sharply recurved than ventral

valves.

Coloration.—Surface of body glabrous; overall coloration intensely greyish brown,

dorsum with indistinct longitudinal stripes; margins of thoracic nota and abdominal terga

dotted with yellow; ventral surface light grey, each sternum bearing a pair of dark brown

spots; legs pale grey, speckled, except apex of hind femur and adjacent tibia solid dark

brown, antenna grey brown, first twelve articles somewhat darker.

Holotype measurements.—(in mm) Length body 23.7, length pronotum 2.2, width

pronotum 3.5, length mesonotum 2.0, width mesonotum 3.7, length metanotum 2.0, width

VOL. 54, NO. 3, JULY 1978 175

£ £

e E 3 3 e E

3 =} ° ° a) rs)

ae = <3 = 5 =5 Sc 2s =

GD De =a Do 5 2 ea) os 23

oS 358 = 52 2 5 2 se oo

- ® Sa Sa SE SE SE Se Sx

Paratop- X24.14 X2.38 X3.44 X1.83 X3.96 X1.45 X4.11 X2.56

type n=21 (+1.15) (+£0.08) (40.24) (+£0.14) (40.13) (+0.10) (40.26) (+0.26)

Mines rd X19.97 X2.11 X1.75 X1.75 X3.48 X1.45 3.67 X2.0

n=11 (+£0.97) (+£0.12) (40.15) (+0.11) (+0.18) (+0.10) (+0.16) (+0.14)

metanotum 3.8, length posterior femur 3.1, length posterior tibia 3.3, length subgenital

plate 2.5

Measurements from the type series were made from liquid-preserved specimens.

Specimens from localities other than those listed were pinned and too distorted for

accurate measurement. All measurements made with ocular micrometer and are in

millimeters. The number in parentheses is the standard deviation.

Records.—CALIFORNIA: ALAMEDA COUNTY: Arroyo Mocho, San Antonio Valley,

Taravel Creek, 21-V-1974, 29-V-1974, I-VI-1973 (C.D. MacNeill, G. Priain, R.E. Love, 629,

including type, CAS). SANTA CLARA COUNTY: South end of Mines road, 30 mi. by road

S.E. of Livermore, 720 m alt., 14-V-1972 (H.B. Leech, 159, 1 3rd instar in alcohol, 359

pinned, CAS). 28 mi. by Mines road S. of Livermore, 720 m alt., 12-V1I-1971 (H.B. Leech, 439,

pinned, CAS). STANISLAUS COUNTY: Del Puerto Canyon, 2-V-1970, 2-V-1972 (R.M.

Brown, 22, CAS); Frank Raines Park, Del Puerto Canyon, 2-V-1970 (R.M. Brown, 4¢, CAS).

All specimens designated as paratypes. Paratopotypes will be deposited in the

Academy of Natural Sciences of Philadelphia and Smithsonian Institution at the request

of the Oakland Museum.

Derivation of name.—This species is named in honor of Ms. Genevieve Prlain in recog-

nition of her interest in the bionomics of the species.

E, genevievae Rentz, new species differs from its closest relative 7.

boharti Tinkham (from mountainous desert regions of western

southern California) in its greyish, never green, coloration, smaller,

less robust size and sculpture of the internal surface of the cercus

(Fig. 5). In T. boharti the apex of the cercus is truncate to emarginate

and always with a prominent dorsal tooth often with several smaller

teeth in females. In 7. genevievae the apex is more or less evenly

serrate. The dorsal internal margin of the cercus of 7. genevievae is

serrate along the entire margin but smooth in 7. boharti. The sub-

genital plate of the two is also quite different. In 7. genevievae this

structure is more constricted basally with the proximal margins

nearly parallel. In 7. boharti the base of the subgenital plate is not

parallel but the margins obtuse. 7. genevievae can be immediately

distinguished from 7. chumash Hebard (coastal and inland central

California) by the much broader and obtuse subgenital plate and the

absence of prominent serrations on the cercus.

Ms. Prlain reports that 7. genevievae shows a preference for

chamise both in the field and in the laboratory. Chamise samples

from Sierran localities were reluctantly accepted, but the captive

timemas always fed on chamise from the type locality and they

176 THE PAN-PACIFIC ENTOMOLOGIST

Cty =

SS ay

cee

Bs

\ Seas aye

2mm LB

ao .

Ss Ca

| ‘Cahiers Ly “f ra eee

Ware Cae 5) h - wes

Mu llinex r 5 2

Figures 2-5.

Fig. 2, dorsal view anterior portion of paratopotype;

Fig. 3, subgenital plate; Fig. 4, apex of abdomen, dorsal view; Fig. 5,

individual cercus, dorsal view. Figs. 3-5, holotype female, Timema

genevievae Rentz.

preferred the budding tips. Even though, a few individuals were

collected on buckbrush (Ceanothus cuneatus (Hook.) Nutt.) and

mountain mahogany (Ceroocarpus betuloides.Nutt.), these shrubs were

rejected as food in the laboratory.

VOL. 54, NO. 3, JULY 1978 177

Eggs from captive females were placed outdoors and moistened

occasionally over winter. In general, they hatched in early January.

Usually six weeks and 4-5 instars later, adults were obtained. At the

type locality, adults were present from mid to late May and mostly

gone by the end of June. Eggs are freely broadcast by females but are

sticky and adhere to the first surface touched.

Timemas should be preserved in a fixative, then stored in alcohol.

Pinned specimens are usually hopelessly shriveled and distorted. |

prefer the fixative described by Williams (1968) for arachnids which is

a modification of the widely used Pampel’s fixative. Pierce (1942)

described a method for drying specimens and then pinning them.

Color and form was reportedly preserved. His method was to place

the living specimens in 70% alcohol for 1 hour, then transfer to 95%

alcohol for 1 hour, then move into xylol for 24 hours. The dried

specimen could then be pinned. He also stated that the same results

could be obtained by placing the timema into xylol and foregoing the

dehydration in alcohol.

Literature Cited

Pierce, W.D., 1942. The preservation of color in soft Orthoptera. Bull. S. Calif. Acad. Sci.,

41:79.

Strohecker, H.F., 1966. New Timema from Nevada and Arizona Pan-Pacific Entomol.,

42:25-26.

Tinkham, E.R., 1942. A new California species of Timema with Zoogeographical notes.

Bull. S. Calif. Acad. Sci., 41:72-79.

White, M.J.D., 1973. Animal cytology and evolution., 3rd. ed. Cambridge Univ. Press,

London, 961 pages.

Williams, S.C., 1968. Scorpion preservation for taxonomic and morphological studies.

Wasmn. J. Biol., 26:133-136.

Over-winter Occurrence and Maturation of Gonads in Adult

Psychoglypha subborealis (Banks) and

Glyphopsyche irrorata (Fabricius)

(Trichoptera: Limnephilidae)

Robert J. Ellis

Northwest and Alaska Fisheries Center Auke Bay Laboratory

National Marine Fisheries Service, NOAA

P.O. Box 155, Auke Bay, AK 99821

Adults of several species of limnephilid caddisflies are collected

only from fall to spring and are variously referred to (for example,

Leonard and Leonard, 1949; Nimmo, 1971; Anderson, 1976) but

essentially they are members of a winter fauna. The actual time of

emergence of adults of these species is usually unknown and the

functional significance of over-wintering adults (in contrast to winter-

emerging adults such as Dolophilodes distinctus (Walker)) has not

been established.

In this paper | describe observations on the occurrence and gonad

maturation of adult Psychoglypha subborealis (Banks) and

Glyphopsyche irrorata (Fabricius) from fall to spring in the vicinity of

Juneau, Alaska, and suggest a functional significance for over-

wintering of these adult caddisflies.

From 1962 to 1974 | casually collected adult caddisflies in the

vicinity of Juneau, Alaska (58°18’N, 134°20’W) (most collections were

made around my home). Because of their presence from fall to spring,

P. subborealis and G. irrorata were especially noticed. | have 30 collec-

tions of P. subborealis (17 males, 22 females) and 20 collections of G.

irrorata (9 males, 21 females; Table 1). Psychoglypha subborealis

occurred every month from October through May, and G. /rrorata

occurred every month from October through May except January.

The adaptation to cold indicated by Denning (1970) for

Psychoglypha in general is confirmed for P. subborealis and G. irrorata

by the minimum temperatures for each month | made a collection

(Table 1) and by my subjective observation of no correlation between

occurrence and warming periods during the winter season. Adults

were collected during very cold weather when all known fresh water

was covered by ice and snow. The implication is not that these adult

caddisflies had recently emerged but that they had emerged during

Open-water periods and survived into winter. The adults could have

Originated from two small permanent streams and several inter-

mittent ponds and streams within 2 km of the collection sites. Nimmo

(1971) refers adults of P. subborealis and G. irrorata to habitats ranging

from ponds to rivers but does not mention temporary waters.

The Pan-Pacific Entomologist 54:178-180. July 1978.

VOL. 54, NO. 3, JULY 1978 179

Table 1. Summary of minimum air temperatures for each month a

collection of Psychoglypha subborealis or Glyphopsyche irrorata was

made, number of collections, and numbers of males and females

collected near Juneau, Alaska, 1962-1974.

Number of:

Minimum air

temperature’ Psychoglypha subborealis Glyphopsyche irrorata

Month (range, °C) collections males females collections males females

October -4 to 1.5 3 2 3 5 6 0

November -16 to -7 4 3 1 1 0 1

December -21.5 to -6 3 1 2 1 0 1

January -30 to -19 2 2 0 0 — _

February -19 to -15.5 3 4 0 1 1 0

March -20 to -9 10 5 11 3 2 2

April -14.5 to -4 4 0 4 6 0 11

May -4 to -2 1 0 1 3 0 6

Totals 30 17 22 20 9 21

‘Local Climatological Data, Juneau, Alaska, National Weather Service Forecast Office, Municipal

Airport, National Oceanic and Atmospheric Administration Environmental Data Service.

Emergence dates of adults (not collection dates), seasonal deposi-

tion of eggs, and life span of P. subborealis and G. irrorata are not

recorded in the literature, but the sequence of maturation of the

ovaries implies certain aspects of their life history. Comparison of

ovaries from my fall to spring collections indicates that Ova mature

during the winter and oviposition occurs in early spring in both

species. During the fall, ovaries are undeveloped and minute and the

abdomen is distended with what appears to be fatty tissue. By mid-

winter the ova are enlarged but remain within the ovariole; much of

the fat has disappeared. In early spring the eggs are fully developed

for oviposition and most of the body fat has disappeared.

Deposition of eggs of P. subborealis appears to commence in early

March since abdomens of two females collected then were only

partially filled with eggs (147 and 150), which is about half the average

of 302 (range 226 to 353) for six subjects ‘‘apparently full” of eggs. No

partially spent females of G. irrorata were collected. The average

number of eggs found in six G. irrorata collected from 24 March to 5

May was 360 (range 243 to 472). The female G. irrorata with the fewest

eggs (243) was collected on 5 May and her abdomen appeared to be

full.

Apparently both species emerge as sexually immature adults in the

fall; the males and females remain active during the winter months,

gradually become sexually mature, and mate and oviposit in the

spring. Males have abundant fatty tissue in the fall and it is almost

totally absent in the spring; however, progressive development of

testes was not apparent from examination of my preserved speci-

180 THE PAN-PACIFIC ENTOMOLOGIST

mens. | did not find pairs in copula but the absence of males in April

and May indicates that mating occurred by March followed soon after

by death of males. Although the evidence is circumstantial, it appears

that the life span of adults is 5 to 7 months.

In a study of adaptations of caddisflies to life in temporary ponds,

Wiggens (1973) reviewed the literature on long-lived adults that

emerge in early summer and survive to oviposit in the fall. These

species have egg matrices which are adapted to permit survival of

drought and freezing. Wiggens’ studies involved species adapted to

live in ponds which were dry from summer to fall (autumnal ponds) or

from summer to following spring (vernal ponds). Over-wintering ina

gelatinous matrix resistant to drought and cold as described by

Wiggens (1973) would be hazardous in climates where freezing

droughts are often interspersed with periods of warming and heavy

rains. Warming and flooding of the gelatinous matrix could result in

dissolution of the matrix and release of larvae susceptible to freezing

and desiccation. Functionally, over-wintering adults with gradually

maturing gonads and spring oviposition permits survival of winter

drought without requiring specially resistant egg matrices, the dry

period in this instance being due to freezing of surface drainage

waters rather than absence of precipitation.

Literature Cited

Anderson, N..H. 1976. The distribution and biology of the Oregon Trichoptera. Oreg.

Agric. Exp. Stn. Tech. Bull. 134. 152 pp.

Denning, D. G. 1970. The genus Psychoglypha (Trichoptera: Limnephilidae). Can.

Entomol. 102:15-30.

Leonard, J. W., and F. A. Leonard. 1949. An annotated list of Michigan Trichoptera.

Occas. Pap. Mus. Zool. Univ. Mich. No. 522. 35 pp.

Nimmo, A: P. 1971. The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta

and eastern British Columbia and their post-glacial origin. Quaest. Entomol.

7:3-234.

Wiggens, G. B. 1973. A contribution to the biology of caddisflies (Trichoptera) in

temporary pools. Life Sci. Contr., R. Ont. Mus. No. 88. 28 pp.

Eugene Graywood Smyth, Ph. D. (1886 - 1975):

Obituary and Bibliography of an Entomologist

Ronald C. Willis

377 Conejo Road

Santa Barbara, CA 93103

The death of Eugene Graywood Smyth, one of the old-time ento-

mologists, is a great loss to the scientific world. He was born in

Topeka, Kansas on July 28, 1886 and began studying insect life at an

early age. He achieved local fame as a teenager for his butterfly

collection.

Dr. Smyth would talk for many hours about the days he was under

the tutorage of Dr. F.H. Snow of the University of Kansas. This was

the era of coal oil lamp use for night collecting and traveling by horse

and wagon. He was a student under Dr. Snow during the years from

1903 to 1907. Between July 22 and August 1 in 1903, he was collecting

insect specimens around Congress Junction in Yavapai County,

Arizona; between August 3 and September 3 near the Tappan’s Ranch

on the Bill Williams fork of the Colorado River; in 1904, at Thomas’

* Ranch in Oak Creek Canyon below Flagstaff, Arizona in Coconino

County; and in 1905, he collected on the San Bernardino Ranch in the

valley of Sycamore Creek east of Douglas, Arizona in Cochise County.

The San Bernardino Ranch belonged to J.H. Slaughter, presumably

the John Slaughter who was sheriff of Tombstone after the Earp-

Clanton shoot-out in the early 1880’s. On an expedition in 1906, Mr.

S.E. Crumb, one of Dr. Smyth’s fellow students, was collecting near

the craggy peaks of the Baboquivari Mountains south-west of

Tucson, Arizona, and while stopped for a lunch break, Crumb dis-

covered and collected some Amblycheila baroni, a very rare tiger

beetle at that time, which was climbing over rocks. The other

members of the expedition then searched and collected as many

other specimens as they were able to locate. They discovered that

these particular tiger beetles would emerge in late afternoon at dusk.

But, the collectors were supposedly endangered by bear and cougar

and could not collect after dark. Each day, Dr. Snow would awaken

his students with the cheery call: ‘‘Blyc, blyc, blyc it again boys, let’s

be off.’’ Referring to the 1905 expedition, Dr. Snow wrote ‘My

assistants on this trip were three students: Eugene Smyth, Ebb

Crumb, and Rollin Perkins. And | have never had a more indefatigable

corps of collectors.’ On these expeditions, the students were

allowed to keep one specimen out of ten for each species collected.

Dr. Smyth would recall the many times he became so absorbed in

collecting in one place too long and would have to run to catch the

disappearing wagon.

The Pan-Pacific Entomologist 54:181-184. July 1978.

182 THE PAN-PACIFIC ENTOMOLOGIST

Economic Entomology was the field of science Dr. Smyth pursued

for most of his life. He won a scholarship to Chicago University. Upon

graduation he joined the United States Department of Agriculture in

Washington, D.C. Working with L.O. Howard, he helped pioneer the

research which proved that house flies contribute to epidemics of

typhoid. Eyebrows were lifted by local residents as Dr. Smyth would

pedal by on his bicycle collecting sticky paper containing fly

specimens for this research.

In 1910, he was sent to Smyra, Turkey to investigate the fig

damaging moth (Esphestia cautella Walk.). After solving this control

problem, he was sent to Arizona in 1911 to work on alfalfa pests. At

this time, blister beetles were defoliating acres of alfalfa. Between

the years 1913 and 1920, he worked with the Department of

Agriculture in Puerto Rico, where he devised new control measures

to eliminate the spread of sugar cane pests. His methods are still

used today for fumigating the ships importing and exporting produce

to and from Puerto Rico. While in Puerto Rico, he worked out the life

history of ten species of white grubs (Lachnosterna), of which five

were species new to science. In 1922 and 1923, L.O. Howard, Chief of

the Bureau of Agriculture, sent Dr. Smyth to southern Mexico and

Guatemala in search of parasites for use in the control of the bean

beetle. Working for the United States Department of Agriculture, he ©

searched for agricultural pests, mainly Epilachna, and encountered

many discomforts such as diarrhea, strange foods, and bed bugs, to

name a few. Traveling into the steep baranncas around Coatepeque

by horse or mule was quite dangerous during the rainy season. Ina

letter to F.H. Chittenden, Chief of Entomology in Washington, D.C.,

Dr. Smyth commented how slippery it became and preferred to trust

his own feet to riding a mule. While in Guatemala and Mexico, he also

collected many specimens of the local flora, which were later

donated to the University of California in Santa Barbara.

During World War |, Dr. Smyth was a U.S. Army officer in Puerto

Rico. While stationed in Puerto Rico, he met and married Laya

Machat, a prima donna with a touring opera company. She later gave

up her singing career to take care of Gene and his insects for the rest

of her life. In 1926, Mrs. Smyth, accompanied by Dr. Smyth, traveled

to Italy for a singing engagement. While there, Dr. Smyth collected

many beetles and visited different research stations in France and

Italy. In 1925, Dr. Smyth was elected a Fellow of the American Asso-

ciation for the Advancement of Science.

From 1929 to 1938, he worked for the Grace and Company sugar

estates in Trujillo, Peru. Here he reared sixty million parasitic wasps

to be turned loose in the sugar cane fields to work where insecticides

could not reach in the heavy sugar cane stalks. In February of 1933,

while on one collecting trip to the remote parts of the Chancamayo

Valley in Peru, Dr. Smyth recalled how he was almost done in by some

VOL. 54, NO. 3, JULY 1978 183

local Indians. In one particular steep canyon, Dr. Smyth is quoted as

saying “The Indians started rolling large boulders down upon me.

With my Colt .45, | sent a volley of shots into the air; and the small,

brown, naked men departed with great haste.” While in South

America, he received his Ph.D. at the University of Peru in Lima.

Semiretired in 1943, Dr. and Mrs. Smyth moved to their home in

Glen Ellen, California, aptly named ‘‘The Perch.” After three years in

The Valley of the Moon, Dr. and Mrs. Smyth moved to Los Angeles in

1946, where he became Associate Curator of Entomology at the Los

Angeles County Museum of Natural History (on a temporary budget

item, July 1, 1946 to June 30, 1951). In 1954 Dr. Smyth retired and

moved to Scarab Hill, the name of his home in Santa Barbara,

California. His love of Entomology led Dr. Smyth to the Chiricahua

Mountains in southern Arizona, with the aid of a grant from the David

Rockefeller Foundation, American Museum of Natural History, New

York, to collect the eggs of Plusiotis gloriosa and to work out its life

history back at his home in Santa Barbara in 1958. He resided in this

coastal city until he passed away on July 30, 1975, three months after

the death of Mrs. Smyth. There were no children surviving them.

Even after seventy years of collecting insects around the world, Dr.

Smyth, when picnicing with his wife, would be sure to have a vial of

formaldehyde along in order to perhaps add a few more specimens to

his collection of over 100,000 insects. Many of his specimens were

donated to the University of Kansas, Los Angeles County Museum of

Natural History, and National Museum of Natural History. He also

donated many unmounted specimens to my own personal collection,

along with his correspondence, papers, books and journals. | am

currently undertaking the task of fulfilling his wish to have his insects

cared for and mounted. As a scientist, Dr. Eugene Graywood Smyth

left behind much completed work and much yet to be completed. His

special love was tiger beetles (Cicindelidae), of which he was able to

on-sight identify over a hundred species. Surely a man who loved

Entomology as much as he, must, even now, have with him a vial of

formaldehyde.

Partial List of Publications by E.G. Smyth

1905. Notes on Collecting Cicindelidae, Trans. Kansas Acad. Sci., 19:425-432.

1907. Notes on Collecting Cicindelidae !/, Trans. Kansas Acad. Sci., 21(1):180-188.

1911. Report on the Fig Moth in Smyrna, U.S.D.A. Bureau Entomol. Bull. 104, pp. 41-65.

1914. Report of the So. Coast Lab., Board of Comm. of Agri. of Puerto Rico, Third Report,

pp. 40-52.

1915. Report of the So. Coast Lab., Board of Comm. of Agri. of Puerto Rico, Fourth Report,

pp. 45-50.

1917. Report of the Entom. Dept., Annual Report of the Insular Exper. Stn. of Puerto Rico,

Dept. of Agri. and Labor of Puerto Rico, pp. 99-106.

184 THE PAN-PACIFIC ENTOMOLOGIST

1919. A Resume of Plant Quarantine Work in Puerto Rico from July 1910 to July 1919, Dept. of

Agri. of Puerto Rico, Plant inspection and Quarantine Report, Bull. 32, pp. 5-56.

1919. Report of the Division of Entomology, Annual Report of the Insular Experiment

Station of the Dept. of Agri. and Labor of Puerto Rico, pp. 27-31 and pp. 109-129.

1919. Dominio de la Plaga de la Mosca Cornupeta de! Ganado en Puerto Rico, Revista de Agri.

de Puerto Rico, Dept. of Agri. and Labor, 3(6):17-28.

1920. Annual Report of the Division of Entomology, Annual Report of the Insular Experiment

Station of the Dept. of Agri. and Labor or Puerto Rico, pp. 83-89.

1920. Cotton Insects in Puerto Rico. Entomol. News, 31:121-125.

1920. White Grubs Injuring Sugar Cane in Puerto Rico, Journal of the Dept. of Agri. of

Puerto Rico, 4:No. 11 Rhinoceros Beetles, pp. 1-31.

1921. La Mosca del Canado (The Horn Fly), Circular 39, Dept. of Agri. in Rio pleura Puerto

Rico, pp. 1-17.

1933. Techniques in the Mass Production of Trichogramma, Journal of Economic Entomol-

. ogy, 25(8).

1933. On the Nicolay & Weiss ‘Synopsis of the Cicindelidae; Entomol. News, 44(8):197-204.

1934. The Gregarious Habit of Beetles, J. Kansas Entom. Soc., 7(334):102-119.

1935. An analysis of the Cicindela Purpurea Group (Coleo: Cicindel), Entomol. News, Vol.

XIVI, #35, pp. 14-19 and pp. 44-49.

1940 Trichogramma Proves Itself in Sugarcane Borer Control, Proc. Inter. Soc. Sugar Cane

Tech., Baton Rouge, La., pp. 367-377.

1960. A New Tephritid Fly Injurious to Tomatoes in Peru, The Bulletin, Dept. of Agri. for the

State of Calif., 49(1):16-22.

1960. Life History of the Silver-Striped Scarabaeid, Plusiotis Gloriosa Le Conte, Bull. So. Calif.

Acad. Sci., 59(2):89-94. ,

The Mango Leaf-Coating Mite, Cisaberoptus kenyae K.

(Eriophyidae, Aberoptinae)

E| Fatih Osman Hassan

University of Khartoum, Shambat, Sudan

and

H. H. Keifer

Sacramento, Cal. ; Collaborator, Agricultural Research Service, USDA

Cisaberoptus kenyae Keifer 1966 is a tiny leaf-coating mite wide-

spread in the tropics, having been transported on its host, mango,

Mangifera indica L. It occurs in southern Asia, east Indian islands,

where mango is grown in Africa, and in South America. The white leaf

coatings it develops on mango leaves are notable in Kenya and the

Sudan. Leaf coatings are conspicuous in Bali, but probably less so in

Thailand. These notable white coatings develop primarily along

the midrib on the upper surface, but as the colony grows the coating

follows lateral veins and goes to leaf margins (Fig. 1). :

Leaf specimens submitted from Thailand by Dr. L. C. Knorr, of the

Plant Protection Service of the United Nations, reveal that the activi-

ties of this mite first can begin inconspicuously on the underside of

the leaf petiole. Petioles thus attacked then become covered with the

typical coating, and show surface roughening under the coating. But

as the colonies enlarge the mites move up the petiole and transfer for

the most part to upper leaf surface areas where they start the typical

development. ;

When first examined this mite seemed to be a leaf miner and as

such was thought to damage the leaf mechanically. Investigations in

the Sudan have shown that there is no leaf mining. Leaf sections from

coated leaves do not disclose any direct damage to internal tissue

(Fig. 4). What eventually does happen is that as coatings become

more and more extensive the leaves tend to yellow and drop pre-

maturely.

A recently published paper by Knorr, Phatak, and Keifer (1976),

entitled Web-spinning eriophyid mites, may have a suggestion as to the

relationship of the white coating to the kenyae mites. In the case of

the web-spinning species, Aculops knorri Keifer (1976), the webs con-

sist of fine discrete strands over the colonies on the upper leaf sur-

faces. Aculops knorri infests a sapindaceous tree, Lepisanthes rubigi-

nosa (Roxb.) Leenh., in Thailand. Serological tests proved that the pro-

teinaceous nature of the web these mites cover themselves with, and

the mite body proteins, are antigenically closely related.

The Pan-Pacific Entomologist 54:185-193. July 1978.

186 THE PAN-PACIFIC ENTOMOLOGIST

Fig. 1 — Mango leaves showing upper surface leaf coatings caused

by Cisaberoptus kenyae. Left leaf with early infestation; right leaf with

more advanced infestation. Ca. 0.57 X.

While the white coating on mango leaves does not readily disclose

separate strands, like the webbing on the Lepisanthes leaves, micro-

scopic examination of the coating (Fig. 3) reveals that it consists of

what might be called crude strands that are irregular. These strands

assume various forms and bunches. The view adopted here is that the

white leaf coating is some sort of regurgitation.

VOL. 54, NO. 3, JULY 1978 187

Fig. 2 — Heteromorphic females as they appear under coating. Ca.

228 X.

Cisaberoptus kenyae is not only notable for the coatings it makes,

but it is also of interest because of different structural forms in

colonies under the coatings. There are three adult forms or types as

follows:

One form is the male (Plate 1) which is a rather ordinary, somewhat

flattened eriophyid, with generalized structures, and which fits easily

into the genus Eriophyes. It has an ordinary-type gnathosome. The

propodosomal shield has dorsal setiferous tubercles on the rear

margin directing the setae to the rear. This male shield is almost en-

tirely devoid of any markings or pattern, but it does show a curved

line just on the inner side of the dorsal tubercle that is similar to the

curved line present in the same position on the shields of females in

the colonies. The abdomen is completely microtuberculate and it

tapers regularly to the terminal lobes. Males vary somewhat in size

and the feather-claws (empodia) have 5 to 8 or 10 rays. The genital

188 THE PAN-PACIFIC ENTOMOLOGIST

Fig. 3 — a. Bunched strands in coating |

b. Separate strands forming compartments. Ca. 1 140 X.

setae are quite small. Average male length from the anterior shield

edge to the termen is 140u.

The second form in these colonies is the female anieh more orless

resembles the male, except for genitalia, and is therefore the proto-

gyne or primary female. These protogynes vary somewhat in size and

in featherclaw rays, and it is possible to divide them into subgroups.

Featherclaw rays on these protogynes run from 7 to 12. Protogyne .

gnathosomes are normal in shape. Protogyne lengths range from

145, to 165y.

The third adult form under the coatings is the most numerous, the

most standardized, and usually the only type easily recovered for ex-

amination (Fig. 2). It is the female that differs notably from the more

generalized types described above (Plate II). This third form is the

deutogyne. It is flattened, but its most outstanding features are the

VOL. 54, NO. 3, JULY 1978 189

Fig. 4 — Cross section of infested leaf showing interior tissue is

not entered by the mites. a. upper epidermis; b. palisade cells; c.

parenchyma Cells; d. lower epidermis.

stocky legs, large complicated featherclaws, and most notably the

stiffened gnathosome. This gnathosome projects ahead and termi-

nates in a pair of expanded spatulate structures. (Fig. 2 and Plate Il).

These spatulate or ‘shovel-nosed’ females are therefore the secon-

dary females and differ mainly by the modified gnathosomal struc-

tures and enlarged legs. These deutogynes are similar to the primary

females, or protogynes, as regards genital structures and the curved

lines on the inner side of the propodosomal shield tubercles. The

large complicated featherclaws have what appears to be 16 to 18 rays.

Deutogynes average 170y in length.

No nymphs of kenyae on slides have shown spatulate gnatho-

somes, but a few second stage nymphs have disclosed developing

adults inside that have spatulate gnathosomes.

190 THE PAN-PACIFIC ENTOMOLOGIST

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Plate | — Cisaberoptus kenyae male.

Abbreviations on Plate | and II

API! — internal female genital structures; CS — caudal section of mite; DA — dorsal view

of anterior section of mite; ES — detail of side ring structures and microtubercles; F —

featherclaw or empodium; GF1 — female genitalia and coxae; GM1 — male genitalia and

coxae; L1 — left foreleg; L2 — left second leg; S — lateral diagram of mite; SA — lateral

view of anterior section of mite

The first stage nymphs of kenyae have about the same basic

arrangement of the dorsal shield tubercles and setae, and the same

interruption of anterior abdominal dorsal rings as other first nymphs

VOL. 54, NO. 3, JULY 1978 191

DA \ x

ia, h fi A

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Cee LE Comcecget yy yh eel Pes

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Plate Il — Cisaberoptus kenyae deutogyne.

in various genera of the Eriophyidae have (Jeppson, et. al. 1975:432,

442). The dorsal tubercles are set ahead of the rear shield edge and

the setae point up and ahead. Several rings behind the shield are dis-

continuous dorsally. The first three subventral abdominal setae on

these first stage nymphs are reduced to small or minute size. The last

subventral abdominal setae before the terminal lobes are of usual

size. These first stage nymphs lack the genital setae which is a feature

not previously observed on other first stage nymphs. There are no ab-

dominal microtubercles present on the first stage nymph.

192 THE PAN-PACIFIC ENTOMOLOGIST

Second stage nymphs have setae about as on adults, perhaps

shorter, but the second stage nymphs share with first stage nymphs

the lack of genital setae. Abdominal microtubercles on the second

nymphal abdomen are present behind the shield, behind the coxae,

and just before the terminal lobes.

This mango leaf coating eriophyid was the first tropical member of

this family that has proved to be deuterogynous. (For a second tropi-

cal deuterogynous species see Keifer 1977:1,2). Deuterogyny is

common among temperate and cold region eriophyid species, mainly

among those that live on deciduous hosts. The female present in tem-

perate regions that displays structures not on the male is the deuto-

gyne or secondary female. These temperate region deutogynes serve

to carry the species over unfavorable periods, chiefly through winter.

In the spring these overwintering deutogynes start new colonies and

then disappear from the colonies. New deutogynes develop later in

the summer in response to changing host and weather conditions.

While these new deutogynes carry sperm over winter they are not

known to be able to lay eggs until they are subjected to winter chill.

On the other hand kenyae deutogynes, with — spatulate

gnathosomes, are always present in colonies on mango leaves, and

they are active egg layers, as shown by slide mounted females of this

type that contain eggs. Thus the kenyae deutogyne performs a func-

tion different from the temperate region deutogyne.

Leaf coating deutogynes in colonies, as well as being active egg

layers, are evidently the form principally instrumental in tending the

coatings. These females congregate around the edges of the covered

areas where they presumably expand the coating, and by means of

their stiff, spatulate gnathosomes keep the coating raised enough to

provide mite space for the colony members. Inactive living kenyae

deutogynes can aestivate for a period of a month or more. These

deutogynes also travel as shown by their presence on grease coated

plates hung in infested groves.

There is at present only one other eriophyid known to be a close

relative of kenyae. This is Aberoptus samoae Keifer (1951). It occurs on

mango in the Samoan Islands. It differs from kenyae principally by

having the spatulate organ on each of the foretibiae, and the forelegs

are stiffened. While this suggests that the female of samoae as de-

scribed is the deutogyne, and has functions similar to kenyae, the

bionomy of samoae is as yet unknown. -

Together, Aberoptus samoae and Cisaberoptus kenyae constitute the

eriophyid subfamily Aberoptinae. |

_ The mango genus, Mangifera, has several species in it. The host of

kenyae is Mangifera indica L., the well known mango. The precise

identification of the Samoan host of samoae is unknown, and the

differences between the position of the spatulate organ on the two

species of eriophyids suggests that possibly the Samoan tree is not

indica. Further exploration may well disclose additional members of

VOL. 54, NO. 3, JULY 1978 193

the Aberoptinae.

As far as presently known all deutogynes in the large subfamily

Eriophyinae of the Eriophyidae are more generalized in their struc-

tures than the males and protogynes, and therefore do not furnish

characters useful in defining taxa. So the practice has been to ignore

them for such definitions.

But this rule now proves inapplicable to members of the subfamily

Aberoptinae of the Eriophyidae. The stiff spatulate gnathosome of

the kenyae deutogyne, and the stiff spatulate forelegs of samoae,

compel their use in defining the genera and subfamily to which they

belong.

But the Aberoptinae are not the only group in the Eriophyoidea that

have deutogynes with useful defining features. In the Diptilomiopi-

dae there are two species possessing flat-backed deutogynes that

constitute the genus Abacoptes (Keifer, 1939b, 1944). The two dip-

tilomiopids are now quoted as Ahyncaphtoptus ulmivagrans Keifer

(1939a), and Rhyncaphytoptus fagifoliae Keifer (1940). These two

species should now be referred to as Abacoptes ulmivagrans (Keifer),

and Abacoptes fagifoliae (Keifer). For an account of the rearing that

proved that the genotype of Abacoptes, which is platynus Keifer, is the

deutogyne of u/mivagrans see Keifer (1944).

Acknowledgement

We are indebted to Dr. L.C. Knorr of the United Nations Plant Pro-

tection Service for his contributions to the bionomy and distribu-

tion of this curious mango leaf coating mite.

References

Jeppson, L.R., Keifer, H.H., Baker, E.W., 1975 Mites Injurious to Economic Plants, 614 p.

photos, University of California Press, Berkeley.

Keifer, H.H., 1939a. Eriophyid Studies VI, Bull. Cal. Dept. Agr. XXVIII (6):420, Aug.

Keifer, H.H., 1939b. Eriophyid Studies III, Bull. Cal. Dept. Agr. XXVIII (7-8-9): 491, Nov. 1939.

Keifer, H.H., 1940. Eriophyid Studies X, Bull. Cal. Dept. Agr. XXIX(3):167, Oct. 1940.

Keifer, H.H. 1944. Eriophyid Studies XIV, Bull. Cal. Dept. Agr. XXXIII(1):30, January 1944.

Keifer, H.H. 1951. Eriophyid Studies XVII, Bull. Cal. Dept. Agr. XL(3):97, Sept. 1951.

Keifer, H.H. 1966. Eriophyid Studies B-18: 1,2, Cal. Dept. Agr., Bur. Ent. April 1966.

Keifer, H.H. 1976. Eriophyid Studies C-12:1 and 2, Agricultural Research Service, USDA,

April 1976

Keiter, H.H. 1977. Eriophyid Studies C-14: 1 and 2, Agricultural Research Service, USDA,

August 1977.

Knorr, L.C. Phatak, H.C., Keifer, H.H., 1976. Web-spinning eriophyid mites, Wash. Acad.

Sci., 66(4):228-234.

The Neotropical Species of Oncylocotis

(Hemiptera: Enicocephalidae)

Gene Kritsky

Dept. Biology, Tri-State University, Angola, IN. 46703

Oncylocotis was erected by Stal in 1855 for the African species

nasutus. Since then the number of Oncylocotis species has greatly

increased with 48 species in the Ethiopian region (Villiers 1969), 2

species in the Palearctic region (Jeannel 1942, Stys 1970), 12 species

in the Oriental region (Usinger 1945, Miyamoto 1965), 2 species in the

Australian region (Usinger 1945), 5 species from the Oceanic region

. (Usinger and Wygodzinsky 1960, Usinger 1945), and 6 species from

the Neotropical region (Stal 1860, Champion 1898). Oncylocotis has

not been collected in the Nearctic region.

Presented herein are descriptions of the six Neotropical species of

which three are new. A key to the Neotropical species is also pro-

vided.

Oncylocotis Stal 1855

Type-species by original designation.—Oncylocotis nasutus Stal (1855).

Moderate to large enicocephalids, 3- 11mm.

Head with deep postocular impression. Posterior lobe usually with transverse impres-

sion. Rostrum short, eyes and ocelli of moderate size.

Pronotum distinctly divided into three lobes. Anterior lobe sometimes with slight

tubercles; intermediate lobe with deep inverted T shaped sulcus and impressions on each

side.

Scutellum triangular shaped.

Foreleg with two claws and four spines on tarsus, one hook-shaped spine and other

three spines curved (Fig. 1). Apical end of the Neotropical species tibia with seven spines,

innermost two spines pear shaped, remainder spines conical and erect (Fig.1).

Forewings with basal cell present and discal cell closed.

Male genitalia without distinct parameres, posterior apophysis of pygophore opening

below the anus.

Female stouter than male. Eyes smaller, posterior lobe of head wider, middle lobe of

pronotum larger, and forelegs more robust. Genitalia reduced to opening below anus.

Key to the Neotropical Species of Oncylocotis

1. Posterior lobe of pronotum greatly wider than intermediate lobe

CRG se) casted pearson has oy cele me oe Beene wee bean ee rhyparus

Posterior lobe of pronotum not greatly wider than intermediate

POD rhage Ores, RIL ee Me nen Eval seen Se Sede I Weert a roe Ree PR all tp ee 2

The Pan-Pacific Entomologist 54:194-198. July 1978.

VOL. 54, NO. 3, JULY 1978 / 195

2. Posterior lobe of pronotum with deep median diverging suture

Ld (9 (GS le Peo ee ae on ne, DIN a By ee UC re PUY, Sha mexicanus

Posterior lobe of pronotum without deep median diverging suture

SB Sn Ee arcana ot Sale 2, SO ocean -2 8.1 aso se rales Op gan 3

3. Posterior lobe of pronotum deeply separated from intermediate

es] sd ocho We: | aN ae dA a mA na ie A emt for Pah be stannardi

Posterior lobe of pronotum not deeply separated. from inter-

CTR NE ahah Ashe Mee dang ibe in re WO he ona ete eets 4

4. Posterior margin of intermediate lobe of pronotum with deep

indentations, and posterior margin of posterior lobe of head

shallowly emarginate (Fig.5) ...............2200 eee braziliensis

Posterior margin of intermediate lobe without deep indentations,

posterior margin of posterior lobe of head not emarginate...... 5

5. Sparsely covered with short setae (Fig.6).............. concolor

Sparsely covered with long setae .................... annulipes

Oncylocotis rhyparus (Stal 1860)

(Fig. 2)

Type-locality.— Rio de Janeiro, Brazil.

Male length 5.00 mm. Body sparsely covered with setae. Head, pronotum, and wings

dull black; rostrum, antennae and legs dark brown.

Head 1.06 mm long. Posterior lobe convex with a slight transverse impression. Eyes

and ocelli of moderate size. Length of antennal segments I, 0.10 mm; II, 0.27 mm; III, 0.30

mm; IV, 0.34 mm.

Pronotum with trapazoidal intermediate lobe with median sulcus and Y-shaped sutures

on both sides. Posterior lobe much wider than intermediate lobe, shallowly emarginate.

Forelegs slender; femur length to width ratio 3.27; tibia length to width ratio 3.70.

Forewings venation complete, small basal cell equal in length to discal cell length.

Material examined collected in Utcuyacu, Peru, Prov. Tarma, 1600-2800 meters eleva-

tion, March 15, 1948 (collected by F. Woytkowski).

Oncylocotis mexicanus, new species

(Fig. 3)

Female length 3.50 mm. Moderately covered with short setae. Body deep brown in

color, legs lighter brown than rest of body.

Head 0.87 mm long. Posterior lobe strongly convex with a deep transverse impression.

Ocelli of moderate size, directed laterally. Length of antennal segments I, 0.08 mm; II, 0.25

mm; Il, 0.27 mm; IV, 0.27 mm.

Anterior lobe of pronotum with two small median tubercles. Intermediate lobe with

deep median sulcus and deep lateral Y-shaped sutures. Posterior lobe shorter than inter-

mediate lobe with a transverse impression dividing posterior lobe into three parts.

Forelegs stout; femur length to width ratio 2.72, tibia length to width ratio 2.81.

Forewing venation complete with long narrow basal cell, discal cell shorter.

Holotype; female; Tapachula, Mexico. May 1902 (collected by Cook and Collins). The

type is deposited in the United States National Museum.

196 THE PAN-PACIFIC ENTOMOLOGIST

Oncylocotis stannardi, new species

(Fig. 4)

Female length 3.50 mm. Sparsely covered with short setae. Legs, rostrum, and wing

bases yellow; remainder of body dark brown.

Head 0.87 mm long. Posterior lobe of head very convex with deep transverse furrow.

Ocelli of moderate size, laterally directed. Length of antennal segments I, 0.08 mm; II, 0.27

mm; III, 0.25 mm; IV, 0.25 mm.

Anterior lobe of pronotum with two small tubercles. Intermediate lobe with a deep

median sulcus and deep lateral impressions. Posterior lobe as long as intermediate lobe,

distinctly separated from intermediate lobe, with emarginate posterior margin.

Forelegs slender; femur length to width ratio 3.11, tibia length to width ratio 3.50.

Forewing with basal cell equal in length to discal cell.

Holotype: female; Bocaiuva, Brazil, 35° 11’ N, 49° 04’W; May 1964 (collected by Fritz

Plaumann). This species is named after Dr. Lewis J. Stannard, Jr., who has made many

valuable contributions to this study. The type is deposited in the United States National

Museum.

Oncylocotis braziliensis, new species

(Fig. 5)

Female length 3.5-4.0 mm. Moderately clothed with long setae. Legs, rostrum, and wing

bases yellow; remainder of body deep brown color.

Head 0.95 mm long. Posterior lobe very convex, with deep transverse impression. Ocelli

of moderate size, laterally directed.

Pronotum with two small tubercles on anterior lobe. Intermediate lobe with deep

inverted T-shaped sulcus and lateral impressions ending in deep indentations in

. posterior margin. Posterior lobe with rounded sides. Shallowly emarginate posterior

margin.

Forelegs stout; femur length to width ratio 2.80, tibia length to width ratio 2.81.

Forewings with narrow and equal length discal cell and basal cell.

Holotype: female; Nova Teutonia, Sta. Caterina, Brazil; June 30, 1955 (collected by F.

Plaumann). Six paratypes; females; same locality as holotype; 2 specimens collected on

April 8, 1953; 1 specimen collected on July 2, 1953; 2 specimens collected on June 30,

1955; and 1 specimen collected on August 15, 1955. The types are deposited in the United

States National Museum.

Oncylocotis concolor (Champion 1989)

(Fig. 6)

Type-locality.— Guatemala, Guatemala.

Female length 4.25 mm. Densely covered with short setae. Legs and rostrum pale

brown, remainder of body reddish brown color.

Head 1.00 mm long. Posterior lobe very convex with a transverse suture. Ocelli of

moderate size and laterally directed. Length of antennal segments I, 0.10 mm; I!, 0.30 mm;

I, 0.32 mm; IV, 0.27 mm.

Pronotum with two small tubercles on anterior lobe. Intermediate lobe with deep

Se

Figs. 1-6, Oncylocotis sp. Fig. 1, O.braziliensis, foreleg; Fig. 2, O.

rhyparus; Fig. 3, O. mexicanus; Fig. 4, O. stannardi; Fig. 5, O. brazilien-

sis; Fig. 6, O. concolor.

198 THE PAN-PACIFIC ENTOMOLOGIST

sulcus and lateral impressions. Posterior lobe with rounded sides posterior margin

emarginate.

Forelegs stout; femur length to width ratio 3.07, tibia length to width ratio 2.84.

Forewings with basal cell equal in length to discal cell, discal cell narrow.

Material examined was collected in Union Juarca, Chiapas, Aug. 11, 1950 (collected by

Goodnight & Goodnight); and from Panama collected by R. Hyssey.

Oncylocotis annulipes (Champion 1898)

Type-locality.—Volcan de Chiriqui, Panama.

Length 4 mm. Sparsely covered with long setae. Body dull black with antennae,

abdomen, and legs lighter color.

Posterior lobe of head convex with a weak transverse impression. Ocelli larger than

concolor.

Intermediate lobe of pronotum with a deep median impression and lateral impressions

on each side. Posterior lobe with rounded sides and shallowly emarginate posterior

margin.

Forewing venation with long narrow basal cell and discal cell shorter.

Acknowledgments

The author would like to thank Dr. Lewis J. Stannard for his help

while completing the study. The author also acknowledges the

following individuals and institutions for the loan of specimens: Dr.

Richard C. Froeschner,; United States National Museum; Dr. P. A.

Arnaud, California Academy of Science; Dr. Jerry Powell, University

of California, Berkeley; and Dr. Thomas Moore, University of

Michigan.

Most of this work was completed while the author was associated

with the Department of Entomology, University of Illinois and the

Illinois Natural History Survey.

Literature Cited

Champion, G. C., 1898. In ‘Biologia Centrali-Americana, Heteroptera’. 2 Vols. 1881-1901,

Il, pp. 158-162.

Jeannel, R., 1942. Les Hénicocephalides, Monographie d’un groupe d’Hémipteres

hematophages. Ann. Soc. entomol. France 110:273-368.

Miyamoto, S., 1965. Enicocephalidae in Taiwan (Hemiptera). Sieboldia 3: 295-304.

Stal, Carl., 1855. Hemiptera fran Kafferlandet. Ofv. Kong. Vet.-Akad. Forh. 12: 27-46.

Stal, Carl, 1860. Bidrag til Rio Janeiro-trakens Hemipter-Fauna. Kong]. Svenska Vet.-

Akad, Hand. 66: 352-365.

Stys, P. 1970. A review of the Palaearctic Enicocephalidae (Heteroptera). Acta entomol.

bohemoslov. 67:223-240.

Usinger, R. L., 1945. Classification of the Enicocephalidae (Hemiptera, Reduvioidea).

Ann. Entomol. Soc. Amer. 38:321-342.

Usinger, R.L., and Wygodzinsky, P., 1960. Heteroptera: Enicocephalidae. Insects of

Micronesia. 7: 219-230.

Villiers, A., 1969. R&évision de Hémipteres Hénicocephalidae Africains et Malgaches.

Ann. Mus. r Afr. cent. 176: 1-232, 335 figs.

Seasonal Abundance and Distribution of Adult Caddisflies

of Sashin Creek, Baranof Island, Southeastern Alaska

(Trichoptera)

Robert J. Ellis

Northwest and Alaska Fisheries Center Auke Bay Laboratory

National Marine Fisheries Service, NOAA

P.O. Box 155, Auke Bay, AK 99821

In this paper | record 21 species of adult caddisflies collected

along Sashin Creek and vicinity and describe their relative abundance

and spatial distribution. Also, incidental observations on biology are

recorded for several species. The collections were made intermit-

tently from 1965 through 1972.

Sashin Creek is located at approximately lat 56°23’N, long

134°44’°W, on the southeast shore of Baranof Island, southeastern

Alaska (Fig. 1). The annual range in volume of flow is about 0.2 to 20

m/s. This study was conducted in and near the stream between its

mouth at tidewater and a 30-m falls about 1,200 m upstream. The

stream water is generally cool, clear, and fast-moving, and the sub-

strate is mostly gravel and rubble. Further details of the physical and

biological setting are available in Ellis (1975).

Materials and Methods

Most of the collecting was done along the stream below the falls

but some was done above the falls and at nearby lighted windows. |

collected with an air net and by hand-picking with forceps at lights. A

few species were collected only at lights. A gasoline lantern and

white sheet were used at three sites below the falls — one close to

tidewater, one about 300 m, and another about 500 m above tide-

water. Lighted windows were visited at residences at the mouth of

the stream (residence l) and about 300 m away along the shore of the

estuary (residence Il). None of the sampling was done on a regular

time or area basis, but on most sampling days, collections were made

from tidewater to falls. The following discussion and list of adult

caddisflies from Sashin Creek are based on specimens collected

along the stream below the falls with air net unless otherwise noted.

Seasonal Distribution and Abundance

On 48 days between 6 June and 4 October | made 206 collections

which yielded 735 adult caddisflies. To compare seasonal occur-

The Pan-Pacific Entomologist 54:199-206. July 1978.

200 THE PAN-PACIFIC ENTOMOLOGIST

rence, it is assumed that timing and sequence of emergence of cad-

disflies are essentially the same from year to year in Sashin Creek

and therefore all collections are combined from 1965 through 1972.

The distribution of collecting by period (approximately a week) is

summarized in Figure 2 together with periods of occurrence for each

species. No collections were made between the first weeks of

October and March which may explain the absence of G/yphopsyche

irrorata and presence of only one specimen of Psychoglypha

subborealis, both winter caddisflies common in the Juneau, Alaska

area (Ellis, 1978).

Several species of caddisflies differed in abundance and length of

occurrence and | have combined the collection data under six general

categories (Table 1 and Fig. 2). The categories and species are:

1. Very abundant for several weeks:

Ecclisocosmoecus scylla (830 specimens in 52 collections made

on 31 days)

2. Abundant for several weeks:

Onocosmoecus unicolor (83 specimens in 41 collections made on

23 days)

Dicosmoecus atripes (88 specimens in 29 collections made on 18

days)

3. Very common for a few weeks:

Ecclisomyia conspersa (56 specimens in 20 collections made on 9

days)

Rhyacophila alberta (88 specimens in 24 collections made on 11

days)

Lenarchus vastus (82 specimens in 23 collections made on 15

days)

4. Common for a few weeks:

Lepidostoma roafi (19 specimens in 14 collections made on 10

days)

Rhyacophila narvae (16 specimens in 12 collections made on 9

days)

Polycentropus halidus (16 specimens in 13 collections made on 9

days)

5. Occasional over several weeks:

Rhyacophila vaccua (5 specimens in 3 collections made on 3

days)

Rhyacophila vao (10 specimens in 3 collections made on 3 days)

Limnephilus harrimani (7 specimens in 6 collections made on 6

days)

Limnephilus nogus (6 specimens in 6 collections made on 6 days)

Limnephilus sitchensis (6 specimens in 5 collections made on 5

days)

VOL. 54, NO. 3, JULY 1978 201

Fig. 1. Map showing location of Sashin Creek in southeastern Alaska.

6. Rare for up to a few weeks:

Glossosoma penitum (2 specimens collected on 2 days)

Rhyacophila rickeri (1 specimen)

Rhyacophila grandis (1 specimen)

Rhyacophila verrula (2 specimens collected on 2 days)

Ptilostomis ocellifera (2 specimens collected on 2 days)

Micrasema sp. (2 specimens in one collection)

Parapsyche elsis (1 specimen)

Chyranda centralis (1 specimen)

202 THE PAN-PACIFIC ENTOMOLOGIST

AND SPECIES 1.2 FUT 2 3a 12 34/123 4/12 34

COLLECTION PERIOD

PSYCHOGLYPHA SUBBOREALIS

ECCLISOMYIA CONSPERSA

ECCLISOCOSMOECUS SCYLLA

LENARCHUS VASTUS

RHYACOPHILA NARVAE

RHYACOPHILA GRANDIS

DICOSMOECUS ATRIPES

ONOCOSMOECUS UNICOLOR

GLOSSOSOMA PENITUM

HOMOPHYLAX ANDAX

LIMNEPHILUS HARRIMANI

POLYCENTROPUS HALINUS

LEPIDOSTOMA ROAFI

CHYRANDA CENTRALIS

LIMNEPHILUS NOGUS

RHYACOPHILA VACCUA

RHYACOPHILA VAQ_

PARAPSYCHE ELSIS

PTILOSTOMIS OCELLIFERA

LIMNEPHILUS SITCHENSIS

RHYACOPHILA ALBERTA

RHYACOPHILA VERRULA

RHYACOPHILA RICKERI

Fig. 2. Periods when collections of caddisflies were made in Sashin Creek 1965-72 (top

line), and periods of occurrence of each species by week.

Homophylax andax (2 specimens in 1 collection)

Psychoglypha subborealis (1 specimen).

The main period of appearance of adult caddisflies in Sashin Creek

is from mid to late summer (Fig. 2). The seasonal progression of

species emerging and the long period of emergence (or long adult

life) for some species make it likely that 8 to 12 species can be col-

lected on any day from about 15 July to 7 September.

Identification of Material

The caddisflies of Alaska have not been treated as a unit and the

descriptions are scattered in the literature. The basic reference work

for the material reported here is Denning (1956). Dr. D.G. Denning

either identified specimens or verified my identifications of all

species reported here. Responsibility for correct identifications of all

remaining material not seen by him is my own.

VOL. 54, NO. 3, JULY 1978 203

Table 1. Number of specimens, collections, and collection days

for each species of adult caddisfly collected in Sashin Creek, 1965-

12.

Number of Number of

specimens Number of collection

Family and species Males Females collections days

Rhyacophilidae

Rhyacophila alberta 36 2 24 11

Rhyacophila grandis 1 0 1 1

Rhyacophila narvae 13 3 12 9

Rhyacophila rickeri 1 0 1 1

Rhyacophila vaccua 5 0 3 3

Rhyacophila vao 8 2 6 3

Rhyacophila verrula 2 0 2 2

Rhyacophila spp. 0 14 9 9

Glossosomatidae

Glossosoma penitum 2 0 2 2

Phryganeidae

Ptilostomis ocellifera 0 2 2 2

Lepidostomatidae

Lepidostoma roafi 19 0 14 10

Lepidostoma sp. 0 8 8 6

Brachycentridae

Micrasema sp. 1 1 1 1

Polycentropodidae

Polycentropus halidus 1 15 13 ]

Hydropsychidae

Parapsyche elsis 0 1 1 1

Limnephilidae

Chyranda centralis 1 0 1 1

Dicosmoecus atripes 71 17 29 18

Ecclisomyia conspersa 47 9 20 9

Ecclisocosmoecus scylla 295 35 52 31

Homophylax andax 2 0 1

Lenarchus vastus 10 22 23 15

Limnephilus harrimani 7 0 6 6

Limnephilus nogus 1 5 5 5

Limnephilus sitchensis 1 5 5 5

Onocosmoecus unicolor 47 36 41 23

Psychoglypha subborealis 0 1 1 1

204 THE PAN-PACIFIC ENTOMOLOGIST

Collection Data

RHYACOPHILIDAE.—Seven species of Rhyacophila were found —

one was very common (Rf. narvae), one was common (R. alberta), and

the rest were occasional or rare.

Rhyacophila alberta Banks.—Most abundant rhyacophilid. Thirty-six

males, two females, August to September, in 24 collections on 11

days. Above and below falls, air net and light.

Rhyacophila grandis Banks.—One male. Captured at light at resi-

dence ll on 12 July.

Rhyacophila narvae Navas.—Thirteen males, seven females, in 12

collections on nine days, 11 July to 30 August, above and below the

falls.

Rhyacophila rickeri Ross.—One male. Captured above the falls on 1

September.

Rhyacophila vaccua Milne.—Five males, no females, in three col-

lections on three days, 1 August to 1 September, above and below the

falls.

Rhyacophila vao Milne.—Eight males, two females, in six collec-

tions on three days, 1 August to 1 September, above and below the

falls.

Rhyacophila_ verrula Milne.—Two males, no females in two

collections on two days, 29 August and 4 October.

Rhyacophila spp.—Fourteen females, in nine collections on nine

days, 19 July to 2 September.

GLOSSOSOMATIDAE.—A single species, G/ossosoma penitum

Banks. Two males in two collections, 19 July and 27 August, above

and below the falls.

PHRYGANEIDAE.—A single species, Ptilostomis ocellifera (Walker).

Two females in two collections on 9 and 13 August. Both collections

at light. One female contained 50 eggs.

LEPIDOSTOMATIDAE.—A_ single species was __ identified,

Lepidostoma roafi (Milne). Nineteen males, no females in 14 collec-

tions made on 10 days, 24 July to 1 September. Eight females in eight

collections are referred to as Lepidostoma sp.

BRACHYCENTRIDAE.—A single species, Micrasema sp. (bactro

group). One male and one female in one collection, 15 July.

POLYCENTROPODIDAE.—A single species, Polycentropus halidus

Milne. One male, 15 females in 13 collections from lights and air net

on nine days, 15 July to 2 September.

HYDROPSYCHIDAE.—A single species, Parapsyche elsis Milne.

One female above falls, 1 August.

VOL. 54, NO. 3, JULY 1978 205

LIMNEPHILIDAE.—The limnephilids were the dominant caddis-

flies in the study area both in total numbers (over 75% of all speci-

mens) and in number of species (11, or about 45% of the number of

species collected). The four most abundant species were

Ecclisocosmoecus scylla, Dicosmoecus atripes, Ecclisomyia conspersa,

and Onocosmoecus unicolor.

Chyranda centralis (Banks).—One male, 29 July.

Dicosmoecus atripes (Hagen).—Seventy-one males, 17 females, 11

July to 2 September, air net and lights (including residence l). The

second most abundant species in this study. The larvae of this large

limnephilid were conspicuous in the stream; they are often eaten by

the water ouzel as evidenced by the abundant empty cases on the

near-water sites frequented by this bird.

Ecclisomyia conspersa (Banks). — Forty-seven males, nine females

in 20 collections on nine days from 6 June to 30 August, above and

below the falls, air net and light. The fourth most abundant species in

this study.

Ecclisocosmoecus scylla (Milne).—Two hundred ninety-five males,

35 females in 52 collections on 31 days from 2 July to 2 September,

above and below the falls. This was the most abundant species in the

study area and was abundant in both air net and light collections.

Homophylax andax (Ross). — Two males in one collection from a

light on 15 July.

Lenarchus vastus (Hagen).—Ten males, 22 females in 23 collections

on 15 days from 10 July to 5 October. Although most collections were

from lighted windows at residence II, three collections were made

with air net along the Creek — one above the falls and two below.

Limnephilus harrimani (Banks). — Seven males in six collections, 18

July to 27 August. The type locality of this species is in Alaska.

Limnephilus nogus (Ross). — One male, five females in five collec-

tions, 30 July to 1 September. Collections made only at mouth of

Sashin Creek with air net (one collection) and at lights at residence Il.

Perhaps larvae do not live in Sashin Creek.

Limnephilus sitchensis (Kolenati).—One male, five females in five

collections on five days from 9 August to 24 August, air net and light.

The females have not been illustrated but the wing patterns of males

and females are similar. Type locality is in Alaska.

Onocosmoecus unicolor (Banks).— Forty-seven males, 36 females in

41 collections on 23 days from 11 July to 4 October. Below falls only,

air net and lights (including residence l). The third most abundant

species in the study area.

Psychoglypha subborealis (Banks).—One female on 6 June, below

falls. This is a winter species usually collected only from fall to spring

(October to May in southeastern Alaska). Type locality is in Alaska.

206 THE PAN-PACIFIC ENTOMOLOGIST

Literature Cited

Denning, D.G. 1956. Trichoptera. /n Aquatic insects of California, pp. 237-270, edited by

Robert L. Usinger, Univ. Calif. Press, Berkeley.

Ellis, R.J. 1975. Seasonal abundance and distribution of adult stoneflies of Sashin Creek,

Baranof Island, southeastern Alaska. Pan-Pacific Entomol., 51:23-30.

Ellis, R.J. 1978. Over-winter occurrence and maturation of gonads in Psychoglypha

subborealis (Banks) and Glyphopsyche irrorata (Fabricius) (Trichoptera:

Limnephilidae). Pan-Pacific Entomol., 54:178-180.

SCIENTIFIC NOTE

Goniopsita oophaga (Diptera: Chloropidae) A Predator of Grasshopper Eggs. — At the

time of its description Goniopsita oophaga Sabrosky was associated as an egg predator of

Melanoplus sanguinipes (Fabricius) and its possible economic significance was pointed

out (Sabrosky, 1967. J. Kansas Entomol. Soc., 40:151-156). Information presented herein

extends the host range of G. oophaga to include a slant-faced grasshopper, Chloealtis

aspasma Rehn and Hebard and covers the frequency and extent of predation on that host.

During 1972 and 1973 numerous ovipositional sites of C. aspasma were noted in dead,

pithy stems of elderberry, Sambucus sp. Oviposition was also induced using stick traps

cut from elderberry (Foster, 1974. Pan-Pacific Entomol., 50:207-208). During both years

numerous fly larvae were observed infesting egg pods of C. aspasma. Egg pods returned

to the insectary during May, 1973 yielded larvae from which G. oophaga adults were

reared.

Predation data from the total of 204 C. aspasma egg pods examined during the study are

tabulated below. Of these, 138 (68 percent) were infested with G. oophaga larvae and

pupae. Substantial egg mortality due to the chloropid is further indicated by comparing

the average number of eggs destroyed per infested pod (7.1) with the observed average

oviposition rate of C. aspasma (7.9). Egg counts of the 204 pods revealed 1,621 grass-

hopper eggs, of which 981 (60.5 percent) were destroyed by G. oophaga larvae.

Present knowledge of the life history of G. oophaga is fragmentary. The presence of two

dead females entrapped on the surface of the recessed plugs of two egg pods indicates

that females may oviposit in the immediate vicinity of the egg pod cap. Although the

grasshopper’s ovipositional period extends from mid-July through August, there is no

evidence that more than one generation of the chloropid develops per year on this host.

All specimens taken during this study were collected 16 mi. northeast of Ashland,

Jackson County, Oregon.

| wish to thank Drs. Ashley B. Gurney and Curtis W. Sabrosky, both of the Systematic

Entomology Laboratory, USDA, Washington, D.C., for their respective identifications of

C. aspasma and G. oophaga, and Dr. W.F. Barr, University of Idaho, who provided labora-

tory space and helped with certain aspects of the field work. — David E. Foster, Depart-

ment of Entomology and The Museum, Texas Tech University, Lubbock, Texas 79409.

Total Average

Egg Pod Data Number Percent Number/Pod Range

Egg pods examined 204 oo + oo

C. aspasma eggs 1621 o--- 7.9 4-8

Egg pods infested 138 68 o-- ---

G. oophaga larvae and pupae 575 o-- 2.3 1-10

Number eggs destroyed 981 60.5 ns ~

Number eggs destroyed/fly ---- a--- 1.7 1-4

Number eggs destroyed/infested pod ao a--- 7.1 0-8

The Pan-Pacific Entomologist 54:206. July 1978.

A New Species of Graptocorixa from Mexico

(Heteroptera, Corixidae)

Antti Jansson

Department of Zoology, University of Helsinki,

P. Rautatiekatu 13, SF-00100 Helsinki 10, Finland

When identifying miscellaneous Mexican Corixidae | found a

species of Graptocorixa which did not agree with any of the descrip-

tions in Hungerford (1948). This new species is described herein and

compared to other species of the genus.

Graptocorixa breweri, new species

Size: Length 7.5-8.5 mm, width across head 2.3-2.7 mm.

Color: General facies medium to dark brown, head and legs yellow, mesosternum and

metasternum smoky to black. Abdominal venter smoky to black in males, yellowish in

females. Pronotum crossed by 8-10 dark bands which are anteriorly broader but

posteriorly narrower than the yellow bands. Transverse yellow bands of hemelytra

generally regular, but somewhat broken in the middle of corium.

Structural characteristics: Face not reduced, frontal depression of male shallow and

broad, slightly hairy; face of female also slightly hairy. Infraocular portions of genae broad

as seen in cephalic view. Rear margins of eyes slightly concave. Prothoracic lobe

elongate. The lower basal angle of fore femur conspicuously produced and anteriorly

covered by a patch of long hairs in male, somewhat produced and with a few hairs in

female. The pala of male as in Fig..1, possessing about 20 pegs. Length of middle leg

femur: tibia: tarsus : claw = 100: 32: 29: 27. Length of hind leg femur: tibia: tarsus 1:

tarsus 2 = 100: 98: 120: 41. Metaxyphus slightly longer than broad. The male genital

capsule as in Fig. 2, with Figs. 3-6 showing variation in shape of the right paramere (=

clasper). Male abdominal dorsum with a posteriorly pointing fingerlike projection near to

the right margin of the fifth tergite, and a small dorsolaterally pointing projection at outer

base of the right penultimate lobe (= seventh tergite) (Fig. 7). Strigil minute, with three

combs (Fig. 8).

In Hungerford’s (1948) key for identification of Graptocorixa species

G. breweri ends up near G. serrulata (Uhler) and G. gerhardi (Hunger-

ford). Clear differences between the three species appear in both

sexes in arrangement of caudo-ventral pegs of middle femora: G.

breweri has a multiple row of small pegs (Fig. 9), while G. gerhardi has

a single row of rather slender, basally almost hair-like pegs (Fig. 10)

and G. serrulata a closely set row of small but sturdy pegs (Fig. 11). In

males clear differences also appear in shape of the right paramere,

and only males of G. breweri have a patch of long hairs on fore

femora. Further, while males of G. breweri have both the fingerlike

projection on the fifth abdominal tergite and the small projection at

the base of the right penultimate lobe, males of G. gerhard/ have only

the former and males of G. serrul/ata only the latter.

Etymology: The species is named simply according to the collector

of the specimens that first caught my attention.

Holotype (male): Mexico, Zacatecas, 25 mi. W. Fresnillo, Laguna

Balderama, 7900 ft., 23.V1.1954, R.H. Brewer. Deposited in the col-

The Pan-Pacific Entomologist 54:207-209. July 1978.

208 THE PAN-PACIFIC ENTOMOLOGIST

/

/

f

i

/

f

t

t

!

i

!

!

f

!

i

!

i

{

{

Be A SS Ce

Figs. 1-9: Graptocorixa breweri. Fig. 1. Front leg of male. Fig. 2. Genital capsule of male.

Figs. 3-6. Variation in shape of the right paramere. Figs. 3-4. Zacatecas. Fig. 5. Michoacan.

Fig. 6. Hidalgo. Fig. 7. Dorsal view of male abdomen. Fig. 8. Strigil. Fig. 9. Middle femur of

female. Fig. 10. G. gerhardi, middle femur of female. Fig. 11. G. serru/ata, middle femur of

female. — Scale lines (1 mm) apply as follows: a = Figs. 7 and 9-11, b = Fig. 8,c = Figs.

1-6.

lections of the California Academy of Sciences.

Paratypes: 6dd and 5992, same data as the holotype (deposited in

the collections of the California Academy of Sciences and Zoologi-

cal Museum, University of Helsinki); 1d and 399, otherwise same data

as the holotype, but collected on 21.V1.1954 (Calif. Acad. Sci. collec-

tions).

Data on distribution: Besides the type series, | have seen the fol-

lowing: 12, Mexico, Zacatecas, 45 mi W Fresnillo, 7700 ft., 25.V1.1954,

VOL. 54, NO. 3, JULY 1978 209

R.H. Brewer (Calif. Acad. Sci.); 3dd and 229, Mexico, Zacatecas, 10 mi

NW of Sombarete, 7700 ft., 1.VII.1954, R.H. Brewer (Calif. Acad. Sci.

and Univ. Helsinki); 292, Mexico, Durango, 35 mi SW El Salto, 8400 ft.,

23.VI1.1953, Univ. Kansas Mexican Expedition, Slesnick Field No. 73

(Snow Entomological Museum, Univ. Kansas); 20d and 12 Mexico,

Hidalgo, Tasquillo, 23.V1.1955, R.E. Beer & party (Univ. Kansas); 19,

Mexico, Michoacan, Morelia, 4.1X.1938, H.D. Thomas (Univ. Kansas),

and 1d, Mexico, Michoacan, 15 mi S of Carapan, 7.XII.1948, H.B.

Leech collector (Calif. Acad. Sci.).

In general, G. breweri seems to be restricted to the central high-

lands of Mexico, and is probably rather common in this area. In the

various collections it might have been overlooked as G. gerhardi,

because the two have rather similar general appearance.

Acknowledgements

| am indebted to the following for loan of material: Dr. D.C. Rentz

(California Academy of Sciences) and Dr. P.D. Ashlock (University of

Kansas).

Literature Cited

Hungerford, H.B. 1948. The Corixidae of the Western Hemisphere (Hemiptera). Univ.

Kansas Sci. Bull., 32: 1-827.

Corrections of Homonyms in Conotrachelus,

and a Species List for Pheloconus

(Coleoptera: Curculionidae)

Charles S. Papp

Plant Industry, California Department of Food and Agriculture, Sacramento, 95814

In preparation for renewed studies in New World Coleoptera, the

following necessary changes have come to my attention.

Conotrachelus Schonherr

aculeatus Papp, new name for hirsutus Fiedler 1940:108 (not Montrouzier 1860:893) from

Bolivia. Type in the Kraatz collection, Deutsches Entomologisches Institut,

Berlin.

carli Papp, new name for humeridens Fiedler 1943:14, and 15-16 (not Hustache 1940:124)

from Brazil. Type in the Chevrolat Collection, Museum Stockholm. In honor of

the late Dr. Carl Fiedler.

exoptorostris Papp, new name for /ongirostris Bondar 1945:327 (not Champion 1904:354)

from Brazil. Type in the Bondar collection, American Museum of Natural

History, New York.

inquinoflavus Papp, new name for griseoflavus Fiedler 1943:26, and 28-29 (not Fiedler 1940:

329) from Colombia. Type in the Chevrolat collection, Museum Stockholm.

nigrivallus Papp, new name for quadraticollis Fiedler 1954:131, and 136-137 (not Fiedler

1940:348-349, and 358) from Brazil. Type in the Sahlberg collection, Museum

Stockholm.

pallidus Papp, new name for a/bosignatus Bondar 1945:321 (not Champion 1904:359) from

Brazil. Type in the American Museum of Natural History, New York.

paranus Papp, new name for tuberosus Fiedler 1940:244, and 266 (not LeConte 1876:233)

from Brazil and Colombia. Type in the British Museum (Natural History),

London.

quaestosus Papp, new name for pallidirostris Fiedler 1943:45, and 56-57 (not Fiedler 1940:

154,161-162) from Cayenne. Type in the Chevrolat collection, Museum Stock-

holm.

schoofi Papp, new name for tibialis Schoof 1942:91-94 (not Hustache 1939:101; not Fiedler

1940:35, and 48) from Mississippi, lowa, Illinois, Kansas, Louisiana, Massa-

chusetts, Missouri, West Virginia. Holotype (male, No. 54310) and Allotype

(female) in the United States National Museum, Washington, D.C.

secernosus Papp, new name for divisus Fiedler 1954:131, and 136 (not Champion 1904:401)

from Brazil. Type in the Kraatz collection, Deutsches Entomologisches

Institut, Berlin.

variabilis Papp, new name for vario/osus Fiedler 1943:15, and 25 (not Hustache 1939:99)

from Brazil. Type in the Chevrolat collection, Museum Stockholm.

vorruptus Papp, new name for interruptus Fiedler 1943:28, and 42-43 (not Pascoe 1889:330)

from Cayenne. Type and paratypes in the British Museum (Natural History),

London, additional paratypes in Museum Dresden.

In 1948, when | completed my papers on Cryptorhynchini (Papp:

1950 and 1951) | was unaware of the publication of Blackwelder’s

Checklist, Part 5 (1947), where adjustments were made for, among

The Pan-Pacific Entomologist 54:210-214. July 1978.

VOL. 54, NO. 3, JULY 1978 211

others, one of Fiedler’s names, which | had also corrected. The

correct citation should now be:

repetitio Blackwelder 1947:853 for /utulentus Fiedler 1940:358 (not Fiedler 1940:

86, and 92) from Brazil (type in the Faust collection, Museum Dresden).

syn.: fiedleri Papp 1951:484 (not Bondar 1944:195).

>

Pheloconus Roelofs

Hustache (1936:38) regarded Phelocomus as an independent genus

very closely related to Conotrachelus. Species shuffling between the

two genera is recorded in the bibliographical data by Hustache (1936).

Working with Fiedler’s types (Papp 1950, and 1951) and evaluating

Fiedler’s argument (1940:6, and 15; 1943: 4-5, this paper sometimes

erroneously cited as 1944) on the validity of the generic concept, we

(Papp and Varga, 1951) came to the conclusion that despite the mor-

phologically slight differences in generic characters, the genus

Pheloconus Roelofs should be recognized. Fiedler’s final position

was later clearly reflected in his writing (Fiedler 1954) and attempts to

break up the genus Conotrachelus.

There is considerable confusion as far as the type of the genus is

concerned. Roelofs selected pilosellus Boheman (1845) as genotype, a

species known in Conotrachelus. Then Champion made a statement

(1904:419) “‘l have seen the types of C. rubicundulus and C. pilosellus,

Boh., the former being abraded and the latter densely squamose.” On

this basis then pilosellus became a synonym of rubicundulus, as

recorded by Champion (1904:418), and Hustache (1936:38).

The following list of species includes three species from North

America, north of Mexico. Their inclusion in the genus is based solely

on description, as mentioned by Fiedler (1940:15; hispidus LeConte

mispelled ‘“‘hispidulus’”. Not identical with hispidulus Hustache

1924:184). To assist further study of the individual species of the

Neotropical region the location of the type specimens are mentioned.

Roelofs, Ann. Soc. Entomol. Belg., 18, 1875:193. — Fiedler, Monograph. London 1940:6.

— Papp and Varga, Portug. Acta. Biol., (B), 3(3), 1951:8. — Fiedler, Neue Sudamer.

Russelk., Jena 1954:138.

aequalis Fiedler, Monograph 1940:8, and 12 (Conotrachelus). Type inthe — Brazil

British Museum (Nat. Hist.). New combination.

albifrons Fiedler, Neue Stidamer. Ruisselk., Jena 1954:91, and 93. Type Paraguay

in the Kraatz collection, Deut. Entom. Inst., Berlin.

albomaculatus Fiedler, Monograph 1940:7, and 8-9 (Conotrachelus). Type _ Brazil

in the Kraatz collection, Deut. Entom. Inst., Berlin. New

combination.

ambiguus Faust, Stett. Ent. Zeit., 54,1893:361. Type in the Faust collec- Venezuela

tion, Museum Dresden. New combination.

212 THE PAN-PACIFIC ENTOMOLOGIST

atrofasciatus Fiedler, Neue Sudamer. Russelk., Jena 1954:90, and 91.

Type in Museum Hamburg.

cretatus Fiedler, Monograph 1940:7-8, and 11-12 (Conotrachelus). Type

and paratype in British Museum (Nat. Hist.). New combina-

tion.

cribricollis Say, Descr. Curc. North Amer., 1831:28 (Cryptorhynchus). Pro-

posed by Fiedler 1940: 15.

fasciolatus Kirsch, Abhandlg. Zool. Mus. Dresden (4), 1889:33

(Conotrachelus). Type in Museum Dresden. New combination.

flavicans Fiedler, 1940:8, and 12-13 (Conotrachelus). Type in Museum

Stettin. New combination.

flavosparsus Fiedler, Neue Sudamer. Riisselk., Jena 1954:138, and 140.

Type in Museum Stockholm.

glabriventris Champion, Biol. Centr.-Amer., Col. 4, 1904:356. Type in

the British Museum (Nat. Hist.). Proposed by Fiedler 1940:

15.

hispidus LeConte, Proc. Amer. Philos. Soc., 15,1876:235 (Conotrachelus).

Champion (1904:419) and Fiedler (1940:15) indicates that

this species probably synonym to rubicundulus Boheman

(1837:447) with distribution from Mexico to Argentina.

hystricosus Champion, Biol. Centr.-Amer., Col. 4, 1904:419 (Conotrache-

lus). Type in the British Museum (Nat. Hist.). Proposed by

Fiedler, 1904:15.

infector Boheman, in Schénherr, Gen. Spec. Curc. 8(2), (Conotrachelus).

Type in Museum Stockholm. Proposed by Fiedler, 1940:15.

lividipes Fiedler, Neue Sidamer. Rlisselk., Jena 1954:138, and 141-142.

Type in Museum Stockholm.

longirostris Fiedler, op. cit., 1954:91, and 93. Type in Museum Hamburg.

luticollis Fiedler, op. cit., 1954:138, and 142. Type in Museum Stockholm.

moestus Fiedler, op. cit., 1954:138, and 142. Type in Museum Stockholm.

nebulosus Fiedler, Monograph 1940:8, and 13-14 (Conotrachelus). Type

in the Kraatz collection, Deutsch. Ent. Inst., Berlin. New

combination.

nudoscutellatus Fiedler, Neue Sidamer. Russelk., Jena, 1954:138, and

139. Type in the Paris Museum.

nubiferus Fiedler, op. cit., 1954:90, and 92. Type in the Kraatz collection,

Deutsch. Ent. Inst., Berlin.

olivaceus Fiedler, op. cit., 1954:91, and 93-94. Type in the Kraatz collec-

tion, Deutsch. Ent. Inst., Berlin.

orchestoides Fiedler, Monograph 1940:7, and 11 (Conotrachelus). Type

and paratypes in the British Museum (Nat. Hist.). New com-

bination.

pallidus Faust, Stett. Ent. Zeit. 54, 1893:361 (Conotrachelus). Type in

Museum Dresden. New combination.

pallisteri Kissinger, Amer. Mus. Novit., 1962:15-18. Types in the Amer.

Mus. Nat. Hist., New York. New combination.

plagiatus Fiedler, Neue Stidamer. Riisselk., Jena 1954: 138-139. Type in

Museum Stockholm.

propinquus Fiedler, Monograph 1940:7, and 9 (Conotrachelus). Type in the

Kraatz collection, Duetsch. Ent. Inst., Berlin. New combina-

tion.

rubicundulus Boheman, in Schtnherr, Gen. Spec. Curc. 4 (1), 1837:447

* (Conotrachelus). Type in Museum Stockholm.

subcarinatus Fiedler, Neue Sudamer, Russelk., Jena 1954:90, 91 and 92.

Type in the Kraatz collection, Deutsch. Ent. Inst., Berlin.

Brazil

Brazil

Eastern half

of USA.

Ecuador

Brazil

Brazil

Panama

Georgia

Panama

New York,

Pennsylvania,

Indiana

Brazil

Brazil

Brazil

Brazil

Bolivia, Peru

Venezuela

Brazil

Bolivia

Brazil

Panama,

Colombia,

Brazil

Mexico

Brazil

Brazil

Mexico to

Argentina

Brazil

VOL. 54, NO. 3, JULY 1978 213

subtriangularis Fiedler, op. cit., 1954:138, and 141. Type in Museum _ Brazil

Stockholm.

sylvius Boheman, in Schonherr, Gen. Spec. Curc. 8(2), 1845:51. Type in Guiana

Museum Stockholm. New combination.

tricinctus Fiedler, Neue Sudamer. Rtisselk., Jena 1954: 90, and 92. Type _— Bolivia

in the Kraatz collection, Deutsch. Ent. Inst., Berlin.

tricolor Fiedler, op. cit., 1954:138, and 139-140. Type in Museum Stock- Chile

holm.

variegatus Boheman, Eugen. Resa 1859:143 (Conotrachelus). Type in Brazil

Museum Stockholm. New combination.

versicolor Fiedler, Monograph 1940:8, and 14-15 (Conotrachelus). Type Peru

in the Kraatz collection, Deutsch. Ent. Inst., Berlin. New

combination.

Literature Cited

Blackwelder, R.E. 1947. Checklist of the Coleopterous insects of Mexico, Central Ameri-

ca, the West Indies, and South America. Part 5. Bull. 185, Smithsonian Inst.,

U.S. Nat. Mus., pp. 765-925.

Boheman, C.H. 1837. /n Schonherr, Genera et species curculionidum. 4 (1):1-600.

Boheman, C.H. 1845. /n Schdnherr, Genera at species curculionidum. 8 (2):1-504.

Boheman, C.H. 1859. Coleoptera. Species nova descripsit. In Kongliga Svenska Fregatten

Eugenies resa, Zoologi, 1 (Insecta):113-118.

Bondar, G. 1944. Notas entomoldégicas. Revista Entomol. Rio de J., 15:191-204.

Bondar, G. 1945. Notas entomoldégicas. Revista Entomol. Rio de J., 16:89-112.

Champion, G.C. 1904. Biologia Centrali-Americana. Coleoptera 4(4), Curculionidae. Pp.

313-440.

Champion, G.C. 1906. Op. cit., pp. 727-728.

Faust, J. 1893. Reise von E. Simon in Venezuela. Curculionidae. Stett. Entomol. Zeit.,

54:313-368.

Fiedler, K. 1940. Monograph of the Southamerican Weevils of the Genus Conotrachelus.

Brit. Mus. (Nat. Hist.), London. 365 pp., frontisp. and 65 figs.

Fiedler, C. 1943. Neue sudamerikanische Conotrachelus aus der Sammlung Chevrolat’s

im Reichsmuseum in Stockholm. Ark. f. Zool., 35A, (6):1-63.

Fiedler, C. 1954. Neue suidamerikanische Arten der Gattung Conotrachelus Schonh. In

‘“‘Neue sudamerikanische Risselkafer’’. Fischer Verlag, Jena, pp. 90-137.

Fiedler, C. 1954. Neue sudamerikanische Arten aus der Subfamilie der Cryptorhynchiden.

In “Neue stdamerikanische Risselkafer’’. Fischer Verlag, Jena, pp. 138-149.

Hustache, A. 1924. Curculionides de l’exploration Lizer-Delétang au Chaco Bolivien. An.

Soc. Cient. Argent., Buenos Aires, 97:156-191.

Hustache, A. 1936. Curculionidae: Cryptorrhynchinae. In Junk and Schenkling Coleop-

terorum Catalogus, part 151, 317 pp.

Hustache, A. 1939. Descripcidn de une especie nueva del genero Conotrachelus Sch. Not.

Mus. La Plata, 14, Zool. (23): 323-325.

Hustache, A. 1939. Curculionides noveaux de |l’Argentine at autres régions Sud-

américaines. Ann. Soc. Cient. Argent.. Buenos Aires, 128:99-124.

Hustache, A. 1940. Curculionides nouveaux de |’Argentine et autres régions Sud-

américaines (deuxiéme note). Ann. Soc. Cient. Argent., 129:112-144.

Kirsch, T.F. 1875. Beitrage zur Kenntnis der Perusanischen Kaferfauna auf Dr. Aben-

droth’s Sammlugen basiert. Deut. Entomol. Zeit., 19:161-208.

Kirsch, T.F. 1889. Coleopteren gesammelt in den Jahren 1868-1877 auf einer Reise durch

Siid Amerika von Alphons Stubel. Abh. Zool. Mus., Dresden, No. 4:1-58.

Kissinger, D.G. 1962. The Curculionid Beetles Collected on the Explorers Club — Ameri-

can Museum of Natural History Entomological Expedition to Yucatan,

Mexico, in 1952. Amer. Mus. Novit., No. 2086:1-28, figs.

214 THE PAN-PACIFIC ENTOMOLOGIST

LeConte, J.L. 1876. In LeConte and Horn: The Rhynchophora of America, North of Mexico.

Proc. Amer. Philos. Soc. 15 (No. 96):VII-XVI, 1-12, and 112-455, (225-235).

Montrouzier, X. 1860. Essai sur la fauna entomologique de la Nouvelle-Caledonie (Balade)

et des Iles des Pins, Lifu, etc. Ann. Soc. Entomol. France, ser. 3, 8:867-916.

Papp, C.S. 1950. Verzeichnis der von Dr. Carl Fiedler beschriebenen sudamerikanischen

Cryptorhynchinen. Part 1. Ark. f. Zool., Ser. 2, 1(11):147-172.

Papp, C.S. 1951. Op. cit., Part 2. Ark. f. Zool., Ser. 2, 2(3):457-518.

Papp, C.S. and G.A. Varga, 1951. Uber die Cryptorhynchinen-Gattung Conotrachelus.

Portug. Acta Biol. (B), 3(3):7-29.

Pascoe, F.P. 1889. On the weevil genus Centrinus and its allies. Ann. Nat. Hist., 6(4):321-

330.

Roelofs, W. 1875. Diagnoses d’esp&ces nouvelles de curculionides. Ann. Soc. Entomol.

Belg., 18:193.

Say, T. 1831. Descriptions of North American curculionides and an arrangement of some

of our known species agreeably to the method of Schonherr. New Harmony,

Indiana, 30 pp.

Schoof, H.F. 1942. The Genus Conotrachelus Dejean (Col.:Curculionidae) in the North

Central United States. II. Biol. Monogr., 19(3):1-170, figs.

RECENT LITERATURE

The following Occasional Papers in Entomology has recently

appeared, and is available free, upon request.

No. 27. The Clearwing Moths of California (Lepidoptera: Sesiidae),

W. Donald Duckworth and Thomas D. Eichlin, 90 pp.

Write to: Library Chairman, Insect Taxonomy Laboratory, 1220 N

St., Sacramento, CA 95814.

Notes on Male Mate-Locating Behavior in Some Bees and Wasps

of Arizona

(Hymenoptera: Anthophoridae, Pompilidae, Sphecidae, Vespidae)

John Alcock

Department of Zoology, Arizona State University, Tempe 85281

Relatively little is known about the reproductive behavior of male

bees and wasps. Yet even from the limited data available it is clear

that the diversity of mate-locating strategies exhibited by males of

these groups is impressive (Alcock et al., in press). Through

comparative studies it may be possible to determine why so many

different kinds of mating adaptations have evolved in the

Hymenoptera. This paper provides brief descriptions of male

behavior in five species of bees and wasps for use in future analyses

of the relation between the reproductive behavior and ecology of

these insects.

The research was done at three study sites in southeastern

Arizona. Study site 1: A flat plain about 1 km north on the road to San

Simon near Portal, Arizona (Cochise County) at an elevation of about

1600 m. The area was covered with a relatively dense stand of

creosote bush. For a full description of the flora in this region see

Linsley and Cazier (1972). Study site 2: A dry stock tank 1 km to the

north of Study site 1. The floor of the tank was partly covered with a

flowering shrub, Baccharis sp., which was attracting many

Hymenoptera. The earthen bank of the stock tank had a sparse cover

of shrubs and gourds. Study site 3: A ridge in the Chiricahua

Mountains 2 km to the south of Portal, Arizona at an elevation of

about 2100 m. Peaks along this ridge supported scattered shrubs and

small pines.

Some individuals of all the species studied were captured and

given distinctive color marks of acrylic or enamel paints on the

dorsum of the thorax. Records were then made of the activities of

known individuals. A stop watch was used to time the duration of

flights from perches or along patrol routes.

Voucher specimens of the undescribed species have been placed

in the entomology collection at Arizona State University.

Results

Triepeolus n. sp. (in manuscript) (det. by Paul D. Hurd, Jr.)

Males of this parasitic anthophorid bee were observed at study site

1 from 3-15 August 1977. Individuals patrolled regular routes through

creosote bush stands from 0730-1330. They flew rapidly, briefly visit-

ing bushes that were an average of 8.1 m apart (N = 19; range = 4.8 - 24

The Pan-Pacific Entomologist 54:215-225. July 1978.

216 THE PAN-PACIFIC ENTOMOLOGIST

m). Upon reaching a creosote station on its route a male would

usually slow somewhat and spend 1-3 sec circling over a portion of

the bush, almost always 12-1 m above the ground, before flying on to

the next inspection point. On three occasions, all between 0815-0845,

a male alighted on creosote leaves on what proved to be a regularly

visited station and walked quickly over the foliage in a small area

(probably applying a pheromone in the process). If this was

pheromone-marking behavior, one brief (less than 5 sec) application

lasted for at least one morning.

The routes of males must have been roughly circular because

recognizable individuals always appeared at a station moving in the

same direction. The interval between visits to the same station

averaged 8.3 min (n= 20, Male “‘White’’), 8.3 min (n=7, Male ‘“‘Pink’’),

9.3 min (n=7, unmarked male), 12.0 min (n=9, Male ‘‘Red-White’’),

and 9.8 min (n= 16, unmarked male). | estimated conservatively that a

male required 5 sec to travel 10 m and to inspect one station. Assum-

ing this rate of travel and taking the minimum time between two visits

to a bush as the time needed to fly around the entire route, these five

males covered patrol routes with a circumference of at least 450-600

m. Because of the difficulty of following the rapid, low-flying males

through the creosote stand, | was never able to trace the path of a

male for more than 80 m.

Some males followed all or a portion of the same route over a

period of days with White, Pink and Red-White seen at one of their

stations on two consecutive days. Pink visited the same inspection

point on 4-6 and 8 August.

There was substantial overlap in routes travelled by different

males. On numerous occasions two males were seen at the same

station, sometimes almost simultaneously. Three marked males were

seen at one bush on 5 August. Often males arrived at an inspection

site from different directions but some individuals followed the same

routes for at least four inspection points.

The behavior of this species of Triepeolus resembles the behavior

of those male bumblebees that mark vegetation at intervals along a

“‘trapline’’ which they patrol (Free and Butler, 1959; Kullenberg et al.,

1973). Bumblebee females wait at an inspection point and copulate

with a male that finds them. This is probably true for this Triepeo/us as

well. | collected conspecific females foraging at creosote bushes and

crushed their thoraxes. The immobilized individuals were placed on

the foliage at stations visited by males. Some specimens elicited no

response but on six occasions, a male located the female (usually

after circling slowly in and around the portion of the creosote bush on

which she had been placed) and attempted copulation. When a male

landed on a female’s back, he quickly oriented himself so that his

head was above the female’s head. He then stroked the female with

his legs and antennae, while his abdomen rhythmically tapped the

VOL. 54. NO. 3, JULY 1978 217

female’s abdomen and his wings whirred in synchrony with the rapid

stroking and tapping movements. This pattern is similar to that of the

anthophorid Centris pallida Fox (Alcock et al., 1977) and the oxaeid

Protoxaea gloriosa (Fox) (Alcock, pers. observations).

The slow circling flight of males as they zeroed in on a female

indicated that they may initially use an odor cue to detect a female

that has landed at a station. Additional evidence for this possibility

comes from four observations of males circling and hovering for over

5 sec in the vicinity of creosote leaves from which a female specimen

had recently been removed. In one case, a male actually alighted on

leaves from which a female specimen had been removed some

minutes previously.

Nomada n. sp. (det. by R. R. Snelling)

This is an undescribed species near N. gutierreziae which was

observed at study site 2 between 3-7 August 1975. Males of this

anthophorid bee also appear to have a patrol route many meters in

length with stations that they visit and mark. Tengo and Bergstrom

(1977) report that males of some European species of Nomada follow

patrol routes used by males of the host species parasitized by their

females. | located a single station at the top of an earthen bank of a

stock tank in an area with cucurbit gourd ground cover and a few stalks

of grass about 15-25 cm high. A male that visited the site would circle

slowly in the vicinity of two grass stalks about 1 m apart before flying

to and alighting upon one or the other of these plants. It then walked

quickly up the stem and along the curving blade of the grass with its

abdomen and head held close to the substrate (Fig. 1). After walkinga

distance of about 10 cm it would fly off so quickly that | was unable to

follow the male to any other station. Tengo and Bergstrom (1977) do

not mention marking of vegetation by European Nomada although

they did find that males applied chemical substances to the backs of

females during copulation. These odors mimic those of the species

of Andrena parasitized by the female Nomada and may permit the

female to gain access to a host nest more easily. The behavior of

Fig. 1. Two views of amale of an undescribed species of Nomada walking along grass

blades that were visited repeatedly by this and other males.

218 THE PAN-PACIFIC ENTOMOLOGIST

males of European Nomada may be derived from marking substrate

with an odor similar to an Andrena host in order to lure a female to a

site where she might be found and induced to copulate.

The two grasses were visited primarily by one male “Orange” that

returned to the station at an average interval of 181 sec on 5 August

(N = 10; 0840-0925; range, 90-445 sec) and 242 sec on 6 August (N = 11;

1015-1100; range, 70-355 sec). On its first visit of the day on 5 August

it marked both grass stems four or five times and remained in the area

for several minutes. Thereafter it marked one grass stem once or

twice and left after less than 30 sec. Although Orange was the

primary visitor, other males were seen at the site on three of the five

mornings of the study. When Orange was collected a new male

became a regular visitor on the following day. Thus as in Jriepeolus

and the trap-line visiting bumblebees, more than a single individual

may come to the same station(s).

Philanthus gibbosus (Fabricius) (det. by H. E. Evans)

Males of this sphecid wasp were observed at study site 2 from 2-8

August 1975. The behavior of male Philanthus has been reported in

some detail for a number of species (Simon Thomas and Poorter,

1972; Alcock, 1975 a, b; Evans 1975). Although Evans (1973) found that

males of P. gibbosus spend the night in nests occupied by their

sisters, he did not discuss male reproductive behavior. | found 2-4

males of this species on each day between 2-8 August at a cluster of

yucca stalks on the earthen bank of the stock tank. Individuals

perched at heights of 2-2/2 m on dried branches or twigs (Fig. 2) and

were separated by about 112-3 m. The wasps regularly flew out from

their perches (usually 2-3 times per min) for an average flight of 5.6

SS Te

Fig.2. A male of Philanthus gibbosus at its territorial perchona yucca stem.

VOL. 54, NO. 3, JULY 1978 219

sec (N = 54; range 1-25 sec) before returning. Upon landing the male

was likely to pump his abdomen up and down while standing on the

perch or walk along the stem dragging the venter of his abdomen on

the substrate for several cm (see Fig. 7, Alcock, 1975a). This behavior

may be associated with the deposition of a sex pheromone (Alcock,

1975a) although during approximately 14 hr of observation no

copulating pairs nor any females appeared in the vicinity of the perches.

Perch owners did attract other male “visitors” (Simon Thomas and

Poorter, 1972; Alcock 1975 b). The resident male flew off his perch at

the approach of a visitor or visitors (up to three intruders were seen

simultaneously at a perch site) and pursued the interloper(s). Males of

P. gibbosus interacted with little or no physical contact, restricting

themselves to simple chase flights. Visitors often remained for

several minutes before finally being driven off or departing of their

own volition. On two occasions a marked resident male was replaced

at its perch by another individual after an interaction between them.

Males ciaimed perch sites in the early morning (0800-0930) and

might remain until midday. Records of marked males show that in-

dividuals held their territories for an average of about 2.2 hrs per day

(with a maximum of 3.5 hrs; N = 7). There was a high daily turnover of

males at the site. Of eight marked individuals only two appeared at

perch for more than one day (one male for two days and another for

four days). Apparently there were many males floating through the

area judging from the number of visitors. On 7 August at midmorning

| removed a resident male from each of four yucca stalks and on the

next day two of the stalks were held by new males.

The social system of male P. gibbosus is similar to that of several other

philanthine wasps (Simon Thomas and Poorter, 1972; Alcock, 1975a; D.

L. Gwynne, pers. communication). Males of these species form small

aggregations of pheromone-applying, territorial individuals that

rarely spend more than a few hours at their perches. Territorial sites

are attractive to other males with several visitors inspecting the perch

on any given day and the same perch occupied by different males on

each of several days in succession. Perch sites are located near

flowering plants or near small diffuse clusters of nesting females.

One active nest of P. gibbosus was found within a few m of the yucca

perch area.

Hemipepsis ustulata ochroptera Stal (det. by P. D. Hurd, Jr.)

Males of this pompilid wasp were studied between 14-19 August

1977 at study site 3. They were found exclusively on three peaks

along a mountain ridge behind Portal, Az. and behaved like a typical

“hilltopping” insect (Shields, 1967). The tarantula hawk males

perched on the tips of pine needles on the stunted, flat-topped pine

trees on the peaks. As many as seven males were seen, each ona

separate tree, during one morning at one peak. Males were active

between at least 0830-1200 and on one afternoon excursion to the site

| found a few males present at 1800.

220 THE PAN-PACIFIC ENTOMOLOGIST

Males flew out from their perches at irregular intervals looping

around the tree several times, usually to inspect or pursue a passing

Hemipepsis from another tree. Normally interactions involved only a

brief period of pursuit in horizontal flight followed by the separation

of males and their return to their respective perches. Despite the

customary absence of overt aggression, this species is clearly ter-

ritorial as shown from the following observations:

(1) Never more than a single male occupied the same tree top (areas

of 3-5 m’).

(2) Three of five males that were marked on 14 August held the same

tree tops on subsequent visits to the area on 16 and 19 August.

(The other two males were not seen again.)

(3) When a male (White) on the highest tree on the peak was captured

and held in a net, its tree top was occupied within 7 min. by an

unmarked male. After 5 min. more, White was released and im-

mediately returned to its perch area. There followed a series of

elaborate aerial duels between the two males. These began with

the horizontal circling chases that occur among established resi-

dent males. The chases quickly led to steep spiral flights with the

two males side-by-side (so close that wing clashing often

occurred). After reaching a height of 10-25 m, the two Hemipepsis

broke off the encounter with first one and then the other diving

back to the perch tree. Within 4 min., White had replaced the

unmarked male although 10-15 additional vertical flights were

seen after White had regained control of the tree.

Transient males were regularly seen. They usually perched in a tree

top for a short time and interacted with nearby resident males. They

then left. Desirable territories were evidently in short supply despite

the availability of unoccupied trees; | removed resident males from

four trees on 19 August and within 4-11 min. three of the trees were

taken by a replacement male. There was some evidence that the

higher the tree, the more desirable it was as a territory:

(1) Males never perched in trees on the lower half of the peaks or on

trees on the saddles between peaks.

(2) The two marked males that did not return to their perches were

captured on the two lowest trees of the five pines that were

occupied on 14 August.

(3) | assume that one of the factors that determines the ability of

males to gain and hold a desirable territory is the size of the

individual. This is true for many territorial invertebrates (e.g.

Potter et al., 1976; Hamilton et al., 1976; Alcock et al., 1977). If this

holds for H. ustulata very small males may be forced to occupy

inferior sites. There was substantial size variation among ter-

ritorial males. The three smallest individuals of 11 collected

males had head-widths of 3.85-4.20 mm vs. 4.50-5.05 mm for the

VOL. 54, NO. 3, JULY 1978 221

Fig. 3. A male of Polistes canadensis navajoe at its territorial perch at the tip of a dead

pine branch.

others; all three were captured on trees well down the slope of the

peak below the other eight males.

No copulations were seen nor were any females observed at the

peaks, although a few were seen searching for prey on mountain

slopes in the Chiricahuas.

Polistes canadensis navajoe Cresson (det. by R. M. Bohart)

This vespid was observed on the same peaks occupied by

Hemipepsis males between 14-19 August 1977. Eberhard (1969) and

Lin (1972) have described territorial behavior in several species of

Polistes, including a different subspecies of P. canadensis, in which

males defended perches near nest sites and hibernation areas. The

males | watched held territorial perches on dead pine branches (Fig.

3), yucca stalks, rocks and patches of foliage on living pines and

other shrubs. The diversity of the perches and the exposed nature of

the peaks makes it unlikely that these wasps were defending hiberna-

tion areas or any other resource of value to females.

During the major period of activity (1000-1900) dozens of males

were present in the area. They constantly made short forays out from

their perches which were separated by a meter or two. Usually these

flights occurred when another Polistes flew within several meters of

222 THE PAN-PACIFIC ENTOMOLOGIST

their perch, although Hemipepsis males were repeatedly pursued and

struck in flight and even butterflies were approached. Intraspecific

encounters generally involved fairly slow sinuous flight chases

without contact. Because several males often occupied the same

tree, when one male flew up others were likely to join one by one in

the chase forming an aerial conga line with all participants returning

to their home perches within 15-30 sec. Sometimes however wing

clashes occurred. More rarely males butted heads in flight and fell to

the ground to grapple together. Four prolonged struggles with much

biting and wrestling were seen.

Marked territory owners exhibited strong site tenacity. Of 12 males

captured and marked on 14 August, 10 returned to their perches

promptly that day, 9 were seen again on 16 August and 8 were at their

territories on 19 August.

One copulation was observed at 1005. A female was seen after she

had arrived on a branch in a male’s territory. The male quickly

scrambled onto her back. He probed with the tip of his abdomen and

soon achieved copulation which lasted less than 30 sec. The pair

then separated (perhaps disturbed by my approach) and the female

flew off.

Discussion

The reproductive behavior of these five bees and wasps is the

product of selection for (1) the ability of males to compete with one

another for access to females and (2) the ability of males to locate

productive areas in which to search or wait for mates. The competi-

tive component of male behavior is most obvious in the territorial

species in which individuals seek to exclude others from perch sites.

At least some males in the populations of P. gibbosus, H. ustulata and

P. canadensis in the Portal area appear to be forming leks in which

aggregated males compete for ownership of purely ‘‘symbolic”’ terri-

tories. Defended sites in lek species do not in themselves contain

females or a resource that draws females to the area. However

receptive females may visit the lek to select a male that has demon-

strated his dominance abilities in competitive interactions with other

males. Leks may evolve when males are unable to monopolize

females directly (e.g. by guarding an emergence area) or indirectly by

controlling resources (Such as a nest site or nectar sources) that

females require (Emlen and Oring, 1977).

The competition among males of the Nomada and Triepeolus bees

was more subtle. They patrolled such a large area that they could not

possibly exclude conspecific males from their searching route and

they did not attempt to do so. But perhaps the male visitors that came

to an inspection point were exploiting the male that marked the

station by seeking to intercept a female attracted to the area by his

pheromone.

VOL. 54, NO. 3, JULY 1978 223

Males of all bees and wasps presumably compete for the best loca-

tions in which to find potential mates. For many species this means

searching at emergence sites and mating with emerging virgin

females (Alcock et al., in press). But for all five species whose

behavior has been described here, emerging females may not be easy

to locate because they are probably not clumped spatially. Females

of Hemipepsis nest where they can find an appropriate spider burrow

and victim (Williams, 1956). Polistes canadensis are not uncommon but

their nests are scattered and the emergence of future queens occurs

over a period of weeks (Eberhard, 1969). Philanthus gibbosus does

sometimes form dense nesting groups (Evans, 1973) but in the Portal

area the species is uncommon and nests appear to be distributed in

ones and twos over a broad area (Cazier and Mortenson, 1965). The

species of Nomada and Triepeolus are almost certainly sparsely

distributed; females may emerge from whatever host burrow their

mother happened to locate. Therefore in these species, males resort

to alternate patterns of mate location. The wasp species gathered at

conspicuous landmark beacons (yucca stalks on an elevated bank,

rocks and vegetation at the top of isolated hill tops) in the vicinity of

nesting habitat of their species. The Philanthus lek was near a nest of

a conspecific female. Female Hemipepsis were seen on mountain

slopes searching for prey and female Polistes nest in rock crevices on

mountainsides.

The two parasitic bees employed the very different alternate

strategy of cruising around a circular route looking for females

attracted to their pheromone stations. Male Jriepeo/us patrolled trap-

lines in areas with flowering creosote bushes visited by conspecific

females and their hosts. Males of the European species of Nomada

patrol areas in which females of the host species are nesting (Tengo

and Bergstrom, 1977).

The landmark and trap-line strategies are employed by other male

bees and wasps whose females are scarce and widely dispersed

(Alcock et al., in press). Perhaps the landmark strategy evolves in

those species whose scattered females have a tendency to search

elevated areas (be they banks, hills or mountain peaks) for flower

resources, or prey or nesting habitat. Males that gather in such areas

and advertise their presence and competitive ability through

pheromones or conspicuous flight patterns may have better-than-

average chance to encounter a receptive female.

The trap-line method may evolve in species whose females have no

preference for conspicuous topographical features in their environ-

ments. Here the superior strategy may be to spread a net of

pheromone marked stations through habitat likely to be visited by

females searching for food or for potential hosts.

224 THE PAN-PACIFIC ENTOMOLOGIST

Summary

Male mating strategies are described for five species of bees and

wasps found in scutheastern Arizona. This is the first report of male

reproductive behavior for representatives of the genera Jriepeo/us and

Hemipepsis. Males of the wasps Philanthus gibbosus, Hemipepsis

ustulata and Polistes canadensis gather at landmark sites and form lek-

like assemblages in which males compete for control of perch terri-

tories. Males of undescribed species of Nomada and Triepeolus patrol

routes through habitat which may contain flowers visited by potential

mates or nests of the host species parasitized by their mates. Both

strategies may arise because receptive females are _ scarce,

dispersed, and difficult to monopolize directly or indirectly through

control of a localized resource valuable to females.

Acknowledgements

| thank H. E. Evans, R. M. Bohart, R. R. Snelling, and P. D. Hurd, Jr.

for their willingness to identify specimens for me. Douglas Whitman

graciously assisted me in my observations of Jriepeo/us. This work

was supported by National Science Foundation Grant DEB76-04503-

A01.

Literature Cited

Alcock, J. 1975a. Male mating strategies of some philanthine wasps (Hymenoptera:

Sphecidae). J. Kans. Entomol. Soc., 48: 532-545.

Alcock, J. 1975b. Territorial behaviour by males of Philanthus multimaculatus (Hymenop-

tera, Sphecidae) with a review of territoriality in male sphecids. Anim. Behav.,

23: 889-895.

Alcock, J., C. E. Jones and S. L. Buchmann. 1977. Male mating strategies in the bee

Centris pallida Fox (Hymenoptera: Anthophoridae). Amer. Nat., 111: 145-155.

Alcock, J., E. B. Barrows, G. Gordh, L. J. Hubbard, L. Kirkendall, D. W. Pyle, T. L. Ponder,

and F. G. Zalom. In press. The ecology and evolution of male reproductive

behaviour in the bees and wasps. Zool. J. Linn. Soc. Lond.

Cazier, M. A. and M. A. Mortenson. 1965. Studies on the bionomics of sphecoid wasps.

Il. Philanthus gibbosus (Fabricius) and Philanthus anna Dunning (Hymenoptera:

Sphecidae). Bull. So. Calif. Acad. Sci., 64: 171-206.

Eberhard, M. J. W. 1969. The social biology of a polistine wasp. Misc. Publ. Mus. Zool.,

Univ. Mich., 140: 1-101.

Emlen, S. T. and L. W. Oring. 1977. Ecology, sexual selection and the evolution of mating

systems. Science, 197: 215-233.

Evans, H. E. 1973. Burrow sharing and nest transfer in the digger wasp Philanthus gibbosus

(Fabricius). Anim. Behav., 21: 302-308.

Evans, H. E. 1975. Nesting behavior of Philanthus albopilosus with comparisons between

two widely separated populations. Ann. Entomol. Soc. Amer., 68: 888-892.

Free) J. B. and C. G. Butler. 1959. Bumblebees. MacMillan, New York.

Hamilton, W. J. Ill, R. E. Buskirk and W. H. Buskirk. 1976. Social organization of the

Namib Desert tenebrionid beetle Onymacris rugatipennis. Can. Entomol., 108:

305-316.

VOL. 54, NO. 3, JULY 1978 225

Kullenberg, B., G. Bergstrom, B. Bringer, B. Carlbert, and B. Cederberg. 1973.

Observations on scent marking by Bombus Latr. and Psithyrus Lep. males

(Hymenoptera, Apidae) and location of site of production of the secretion.

Zoon, Suppl. 1: 23-30.

Lin, N. 1972. Territorial behavior among males of the social wasp Polistes exclamans

Viereck (Hymenoptera: Vespidae). Proc. Entomol. Soc. Wash., 74: 148-155.

Linsley, E. G. and M. A. Cazier. 1972. Diurnal and seasonal behavior patterns among

adults of Protoxaea gloriosa (Hymenoptera, Oxaeidae). Amer. Mus. Novit., 2509:

1-25.

Potter, D. A., D. L. Wrensch and D. E. Johnston. 1976. Aggression and mating success in

male spider mites. Science, 193: 160-161.

Shields, O. 1967. Hilltopping. J. Res. Lepid., 6: 69-178.

Simon Thomas, R.T. and E. P. R. Poorter. 1972. Notes on the behaviour of males of

Philanthus trianguium (r.) (Aymenoptera, Sphecidae). Tijdschr. Entomol., 115:

141-151.

Tengo, J. and G. Bergstrom. 1977. Cleptoparasitism and odor mimetism in bees: Do

Nomada males imitate the odor of Andrena females? Science, 196: 1117-1119.

Williams, F. X. 1956. Life history studies of Pepsis and Hemipepsis wasps in California

(Hymenoptera: Pompilidae). Ann. Entomol. Soc. Amer., 49: 447-466.

Life History of Zygogramma disrupta

in Southeast Texas

(Coleoptera: Chrysomelidae)!

G.L. Piper?

Dept. Entomology

Texas A&M University

College Station, 77843

The ragweeds (Ambrosia spp.: Compositae) are important agri-

cultural and hayfever-inducing weeds throughout much of North

America (NAS 1968, Wodehouse 1971) and Eurasia (Kovalev and

Runeva 1970, Kovalev 1971). The most abundant and widespread

species include common ragweed (A. artemisiifolia L.), western or

perennial ragweed (A. psilostachya DC.), and giant ragweed (A. trifida

L.) (Payne 1970). Surveys of the entomofaunas of these and other

North American ragweeds have revealed the existence of a diverse

assemblage of insects (Hack 1935, Harris and Piper 1970, Piper 1970,

Stegmaier 1971, Goeden and Ricker 1974a, 1974b, 1975, 1976a, 1976b,

1976c).

In southeast Texas, A. psilostachya is attacked by several species

of phytophagous Coleoptera. One such associate is the chrysomelid,

Zygogramma disrupta (Rogers).

The genus Zygogramma Chevrolat contains 14 species in America

north of Mexico (Arnett 1968) and 85 Neotropical species (Bechyne

1952). Linell (1896) provided a key to the Nearctic species. Biological

information is available for only 2 North American species, the sun-

flower beetle, Z. exclamationis (Fabricius) (Walker 1936), and Z.

suturalis (Fabricius) (Piper 1975). Rogers (1856) described and il-

lustrated the adult of Z. disrupta. The occurrence of this beetle on

weeds and willow (Sa/ix sp.) during the summer months in Kansas

was noted by Douglass (1929). The paucity of information on the

biology of Z. disrupta prompted the study reported herein.

Materials and Methods

Studies on the life cycle of Z. disrupta were conducted in the

laboratory and correlated with field observations. The laboratory

colony, started from adult beetles collected from A. psilostachya

growing in or near College Station (Brazos Co.) during 1976 and 1977,

were maintained in the laboratory at 24 + 2°C, 75 + 5% RH, anda

14L:10D photoperiod. Eggs, larvae, and adults were confined to 10.0

‘Approved as Technical Article 13793, Texas Agric. Exp. Stn. Series.

2Present address: Dept. Entomology, Washington State University, Pullman, WA 99163.

The Pan-Pacific Entomologist 54:226-230. July 1978.

VOL. 54, NO. 3, JULY 1978 227

x 1.5 cm petri dishes provisioned with moistened filter paper and rag-

weed foliage. Water and food were renewed at 48 hr intervals. Pupae

were held in petri dishes containing moist peat moss.

Life History

Zygogramma disrupta has been recorded from Nebraska, south to

Mexico, east to Louisiana, and west to Arizona (Rogers 1856, Powell

1932, Wilcox 1975). Based upon descriptions provided by Rogers

(1856) and Linell (1896), adults are broadly oval, convex, with the head

and pronotum dark brown; each elytron cream-colored, with suture

and four narrow discal vittae dark brown; first vitta free, slightly

abbreviated basally and apically; second and third sinuous, ab-

breviated basally and confluent apically; fourth short, confluent with

third basally; three submarginal spots apically; fourth short, con-

fluent with third basally; three submarginal spots apically; epipleuron

cream-colored with dark brown margin. The lengths of 25 males and

25 females averaged 5.75 + 0.06 mm and 6.47 + 0.07 mm, respectively.

Z. disrupta is a bivoltine species in southeast Texas. First genera-

tion overwintered adults began emerging during early June at a time

when A. psilostachya plants were 20-25 cm in height. Peak populations

were attained by mid-June. Adults of the second generation emerged

in late July with peak populations being found during early August.

Both sexes emerged concurrently. Sex ratio was determined by

examining 94 beetles which emerged from laboratory-reared first

generation pupae. The male:female ratio was 1.0:1.3.

The beetles did little flying, remaining closely associated with the

uppermost foliage of the host plant. Rarely were more than three

individuals found per plant in nature. The adults fed upon both older

and newly developed leaves, often consuming leaf laminae in their

entirety except for the midribs and immediately adjacent tissue.

Adults feign death and drop from the plant at the slightest dis-

turbance, their coloration blending imperceptibly with the soil and/or

litter below. The average life span of laboratory-reared adults was 49.3

+ 8.4 (range 12-83) days for males and 58.4 + 5.4 (range 10-111) days for

females.

The premating period was not determined. Mating occurred most

frequently during either the late morning and/or mid- to late after-

noon in both the field and laboratory. All matings observed in nature

occurred on the upper surfaces of leaves or in leaf axils of the host

ptant. The act of copulation in Z. disrupta was similar to that of Z.

suturalis (Piper 1975). The male mounted the female from behind by

moving directly forward and onto her dorsum. The male positioned

himself on the posterior dorsum at a 45° angle with his forelegs

resting upon the female’s elytra near the second vittae. The middle

pair of legs grasped the elytral margins or epipleura near the apex.

228 THE PAN-PACIFIC ENTOMOLOGIST

The male’s hind tarsi flanked the vagina. The tarsal lobes were rubbed

across the vaginal area preparatory to aedaegus extrusion and intro-

mission. The female generally was quite passive after the initiation of

copulation and often fed while in copula. When a female became rest-

less and started to move about on the plant, copulatory activity was

suspended immediately; the male merely rode the female’s dorsum

until another site was selected. Upon the cessation of female move-

ment, the male resumed copulation. The time spent in copula varied

from 15 min to two hr. Repeated matings between the same male and

female occurred frequently in the laboratory both before and after

oviposition had commenced. However, mating declined steadily in

frequency as the female aged. A male was capable of mating with

different females and a female accepted different males.

The preoviposition period, from emergence to first deposition of

eggs, of 25 females averaged 12 + 0.8 (range 9-14) days. The oviposi-

tion period averaged 37.3 + 4.7 (range 7-97) days during which time the

females laid an average total of 262 + 43 (range 27-845) eggs. Daily

egg production was quite variable. Based only on days when females

oviposited, the daily totals ranged from 1 to 40 and averaged 9.3 + 0.7

eggs.

Oviposition occurred most commonly during the late afternoon or

evening. Eggs characteristically were deposited in clusters of 2 to 18

on the undersides of young ragweed leaves. Of 100 clusters

examined, 75 contained two to five eggs, with a mean of five eggs/

cluster. The eggs overlapped one another and stood out from the leaf

at a 45° angle. In the field, first generation eggs were found from mid-

June to late July; second generation eggs from mid-August to late

September.

The egg is ellipsoidal, yellowish-orange and has an asperate

chorion. The mean length and width of 25 eggs were 1.57 + 0.01 mm

and 0.69 + 0.01 mm, respectively. The incubation period of 300 eggs

averaged 6.26 + 0.03 (range five to seven) days. Eggs in the same

cluster usually hatched concurrently.

Recently hatched larvae were pale orange and fed together at the

margin of the leaf on which the eggs had been laid. The pale yellow

second-, third-, and fourth-stage larvae were less gregarious and

distributed themselves among the uppermost foliage. Larvae fed on

the upper surfaces of leaves and the resultant damage resembled

that produced by adult feeding. When touched or disturbed while

feeding, third- and fourth-stage larvae either regurgitated undigested

foodstuffs or voided a droplet of fecal material, a behavioral trait also

characteristic of Z. suturalis larvae (Piper 1975). Confinement of late

stage larvae to potted ragweed usually resulted in extensive defolia-

tion and plant death. However, the damaging defoliation levels pro-

duced in the laboratory were never observed among field populations

of Z. disrupta larvae.

VOL. 54, NO. 3, JULY 1978 229

The maximum lengths attained by 25 first- to fourth-stage larvae

were 2.43 + 0.02 mm, 3.22 + 0.06 mm, 4.25 + 0.07 mm, and 6.00 + 0.13

mm, respectively. Under laboratory conditions, the stadia of larval

development were as follows: first, 5.2 + 0.1 (range four to seven)

days; second, 3.6 + 0.2 (range two to six) days; third, 4.2 + 0.2 (range

three to eight) days; and fourth, 7.3 + 0.4 (range six to ten) days.

During the fifth or sixth day of development, a fourth-stage larva

ceased feeding, descended to the ground directly beneath the plant,

and burrowed five to ten cm into the soil. Having reached a suitable

depth, the larva pushed and compacted the surrounding soil, creating

an ellipsoidal chamber. After completing the earthen cell, the larva

became quiescent and entered the prepupal stage which lasted 6.8 +

0.6 (range five to nine) days.

Pupation occurred within the chamber. The pupa is pale yellow and

the mean length of 25 pupae was 5.87 + 0.06 mm. The duration of the

pupal period averaged 7.8 + 0.1 (range seven to nine) days. After

ecdysis, teneral adults remained underground one to two days before

emerging. Upon emergence, adults are pale and are still quite soft.

Normal coloration and complete sclerotization were attained within

eight to twelve hr. During late September and October, overwintering

of adults produced from second generation pupae commenced

within the pupal cells. In the laboratory, the time required for a

complete life cycle, from egg to adult, was 39 + 1 (range 28-41) days.

Acknowledgment

The author wishes to express his appreciation to R. E. White,

Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv., USDA,

for identifying the chrysomelid; and to P. Allen and J. Loehr for their

assistance with field collections.

Literature Cited

Arnett, R. H., Jr. 1968. The Beetles of the United States. Amer. Entomol. Inst., Ann Arbor,

Mich.

Bechyne, J. 1952. Nachtrage zu den Katalogen von neotropischen echten Chrysomeli-

den. Entomol. Arb. Mus. G. Frey Tutzing Muenchen 3:1-62.

Douglass, J. R. 1929. Chrysomelidae of Kansas. J. Kans. Entomol. Soc. 2:1-15.

Goeden, R. D. and D. W. Ricker. 1974a. The phytophagous insect fauna of the ragweed,

Ambrosia acanthicarpa, in southern California. Environ. Entomol. 3:827-834.

Goeden, R. D. and D. W. Ricker. 1974b. The phytophagous insect fauna of the ragweed,

Ambrosia chamissonis, in southern California. Environ. Entomol. 3:835-839.

Goeden, R. D. and D. W. Ricker. 1975. The phytophagous insect fauna of the ragweed,

Ambrosia confertiflora, in southern California. Environ. Entomol. 4:301-306.

Goeden, R. D. and D. W. Ricker. 1976a. The phytophagous insect fauna of the ragweed,

Ambrosia dumosa, in southern California. Environ. Entomol. 5:45-50.

Goeden, R. D. and D. W. Ricker. 1976b. The phytophagous insect faunas of the ragweeds,

Ambrosia chenopodiifolia, A. eriocentra, and A. ilicifolia, in southern California.

230 THE PAN-PACIFIC ENTOMOLOGIST

Environ. Entomol. 5:923-930.

Goeden, R. D. and D. W. Ricker. 1976c. The phytophagous insect fauna of the ragweed,

Ambrosia psilostachya, in southern California. Environ. Entomol. 5:1169-1177.

Hack L. 1935. Insects of the giant ragweed (Ambrosia trifida L.). Unpub. M.S. Thesis,

Univ. Kans.

Harris, P. and G. L. Piper. 1970. Ragweed (Ambrosia spp.: Compositae): Its North

American insects and the possibilities for its biological control. Common-

wealth Inst. Biol. Contr. Tech. Bull. 13:117-140.

Kovalev, O. V. 1971. [Phytophages of ragweeds (Ambrosia L.) in North America and their

application in biological control in the U.S.S.R.]. Zool. Zh. 50:199-209. (In

Russian).

Kovalev, O. V. and T. D. Runeva. 1970. [Tarachidia candefacta Hubner (Lepidoptera,

Noctuidae), an efficient phytophagous insect for biological control of weeds

of the genus Ambrosia L.] Acad. Sci. U.S.S.R. Entomol. Rev. 49:23-36. (In

Russian).

Linell, M. L. 1896. A short review of the Chrysomelas of North America. J. N. Y. Entomol.

Soc. 4:195-200.

National Academy of Sciences. 1968. Weed Control. Pub. 1597. Washington, D. C.

Payne, W. W. 1970. Preliminary reports on the flora of Wisconsin, No. 62. Compositae VI.

The genus Ambrosia — the ragweeds. Trans. Wis. Acad. Sci. Arts and Letters

58:353-371.

Piper, G. L. 1970. The biology and immature stages of certain Coleoptera associated with

common ragweed, Ambrosia artemisiifolia L. Unpub. M.A. Thesis, Kent State

Univ. .

Piper, G. L. 1975. The biology and immature stages of Zygogramma suturalis (Fabricius)

(Coleoptera: Chrysomelidae). Ohio J. Sci. 75:19-24.

Powell, E. F. 1932. The Chrysomelinae of Nebraska (Coleop.: Chrysomelidae). Entomol.

News 43:92-97.

Rogers, W. F. 1856. Synopsis of species of Chrysomela and allied genera inhabiting the

United States. Proc. Acad. Natur. Sci. Phila. 8:29-39.

Stegmaier, C. E., Jr. 1971. Lepidoptera, Diptera, and Hymenoptera associated with

Ambrosia artemisiifolia (Compositae) in Florida. Fla. Entomol. 54:259-272.

Walker, F. H., Jr. 1936. Observations on sunflower insects in Kansas. J. Kans. Entomol.

Soc. 9:16-25.

Wilcox, J. A. 1975. Checklist of the Chrysomelidae of Canada, the United States, Mexico,

Central America and the West Indies. Vol. 1, Pt. 7. (Red version). Biol. Res.

Inst. Amer., Inc., Latham, N. Y.

Wodehouse, R. P. 1971. Hayfever plants. 2nd ed. Hafner Publ. Co., New York, N. Y.

A Mating Aggregation of Dasymutilla foxi in Southem Arizona

(Hymenoptera:Mutillidae)

Donald G. Manley

Dept. of Entomology, University of Arizona, Tucson 85721

and

Stephen Taber, III

U.S. Department of Agriculture

Bee Research Laboratory

2000 E. Allen Road, Tucson, Arizona 85719

In general, velvet ants are most abundant in deserts or similar arid

regions (Hurd, 1951). In the United States, numerous species belong-

ing to the genus Dasymutilla Ashmead occur in the southwestern

states.

Velvet ants are not frequently encountered in large numbers. Fattig

(1936) described seeing unusual mating behavior involving Mutilla

briaxus Blake, in which ‘‘at least” 40 males were involved. Linsley et ai.

(1955) told of an ‘unusually large concentration” of Dasymutilla

formicalia (Rohwer) in which 31 males and 37 females were collected

in a period of about one hour. Other records have been made of speci-

mens in higher densities than usual (Mickel, 1928; 1938), but these

apparently have not involved any true aggregation.

The present study involves a very large aggregation, including both

males and females, of D. foxi(Cockerell). The site was located in Pinal

Co., Arizona, about 40 km (25 miles) north of Tucson (Star Flat Tank,

Black Mt., 7% min. topographic series, about 32°45’00’N/110°52’

30’’W). It was discovered on September 11, 1976, and has now been

under observation for more than 12 months. The site consists of two

reservoirs used for watering cattle, separated by an earthen dam.

Each reservoir is approximately one half hectare in area. The dam is

nearly devoid of vegetation, though some trees of the species

Prosopsis juliflora (Swartz) border the water in the reservoirs. The

dominant plant species in the area is Larrea divaricata Cavanilles.

Vegetation is relatively sparse. Two species of bees, Diadasia rinconis

Cockerell and D. opuntiae Cockerell (identified by E. Ordway), were

nesting at the site.

Materials and Methods

Visits every two to three weeks for the 12 months were made to the

study area for purposes of collection and observation, and for deter-

The Pan-Pacific Entomologist 54:231-235. July 1978.

232 THE PAN-PACIFIC ENTOMOLOGIST

FE TE ee TE EEE Gat

I

RESERVOIR

RESERVOIR

ee High motillid density

=== ~=©Marginal motillid density

Scale: 4

10 m

Fig. 1. A map of the aggregation site at which mutillid activity was observed. There was

an area of high density surrounded by an area of marginal density on and near the earthen

dams.

mining the population size. During a visit on September 29, a map

(Fig. 1) was made in which the aggregation was located. A marginal

area was also noted. Within the location, five one square meter plots

were marked off. The distance between the sample areas was about 6

meters. The number of male and female mutillids in each square

meter was first counted at 1710 MST, and then at 10 minute intervals

for 80 minutes.

Visits were made to the site during all daylight hours to determine

when the mutillids were most active.

It was Suspected that the aggregation may have been for purposes

of courtship and mating, as many pairs were observed in copulation.

To determine whether successful mating was being accomplished,

several pairs of specimens were collected in copula on September 22

and taken to the USDA Bee Research Lab in Tucson for dissection.

Results

The area of high concentration covered approximately 1300 square

meters (Fig. 1). Using the data collected from the one meter squares

VOL. 54, NO. 3, JULY 1978 233

x GOO

Fig. 2. A drawing of the reproductive tract of a female Dasymutilla foxi, with an enlarged

drawing of the spermatheca (A) showing the sperm therein. GNG, ganglion; OVL, ovariole;

SH, sheath; STN, sting; VNMSC, venom sac. (Drawing by R. Schmalzel).

within the area of high mutillid concentration, it was calculated that

there were nearly 5 velvet ants per square meter in the area. Using

this, and the total area of the aggregation, it is estimated that the pop-

ulation consisted of approximately 6000 individuals, with a sex ratio

approaching 1:1.

Activity was observed during all daylight hours, and even shortly

after sunset. Greatest activity, however, was observed from about

1500 to 1900 MST.

It was determined that successful mating was occurring within the

population, as live spermatozoa were observed in spermathecae. A

234 THE PAN-PACIFIC ENTOMOLOGIST

drawing of the female reproductive system and the spermatheca

(enlarged) is shown in Fig. 2.

Discussion and Conclusions

~ Even in the Southwest, where velvet ants are relatively numerous, it

is unusual to see many individuals in any one location. It seems rea-

sonable to assume that the large aggregation observed was not

present at the same location at the same time purely by coincidence.

The aggregation of velvet ants was first observed on September 11,

1976. Subsequent observations were made through October 11, 1977.

The density of the aggregation was greatest from the time of

discovery until about mid-October, at which time the number of

individuals present began to decline. The decline continued through

November, by which time no further specimens could be found.

Much of the area in which the mutillids were numerous, particu-

larly on the slope of the earthen dam, was dotted with burrows

measuring about 5 mm in diameter. Velvet ants, both males and

females, were observed entering and exiting the burrows. It has been

documented that D. fox/ is parasitic upon bees of the genus Diadasia

Patton (Cockerell, 1896). In the fall of 1976, although several burrows

were excavated, nothing was found in them. It seemed probable that

the burrows belonged to the host, but that the burrows were empty

due to the relatively late dates at which they were examined.

No mutillid activity was observed at the site from November until

the following April. At that time there was considerable host nesting

activity. Burrows were again excavated and host pollen balls and

larvae were found. At that time the mutillid population was very light

and limited to females. No males were observed until mid-May. The

mutillid population remained small and steady throughout the

summer and early fall. At no time during 1977 did the population

reach even % the size of the fall 1976 population.

The activity over the course of the year seems to clearly indicate

that the site is being used as a courtship and mating site for the —

mutillids, as well asa host nesting site.

Literature Cited

Cockerell, T.D.A. 1896. Specific characters among the Mutillidae. Nature 54:461.

Fattig, P.W. 1936. An unusual mating of velvet ants (Hymenoptera: Mutillidae). Entomol.

News 47:51-52. :

’ Hurd, Paul D., Jr. 1951. The California velvet ants of the genus Dasymutilla Ashmead. Bull.

_ Calif. Insect Surv. 1:89-116.

Linsley, E.G., J.W. MacSwain, and R.F. Smith. 1955. Observations on the mating habits of

Dasymutilla formicalia Rohwer (Hymenoptera:Mutillidae). Can. Entomol. 87:411-

413.

VOL. 54, NO. 3, JULY 1978 235

Mickel, Clarence E. 1928. Biological and taxonomic investigations on the mutillid wasps.

U.S. Nat. Mus. Bull. 143:1-351.

Mickel, Clarence E. 1938. Photopsoid mutillids collected by Dr. K.A. Salman at Eagle

Lake, California (Hymenoptera). Pan-Pac. Entomol. 14:178-185.

BOOK REVIEW

A Revision of the Subfamily Coelidiinae (Homoptera: Cicadellidae) Il.

Tribe Thagriini. Nielson, M. W. 1977. Pacific Insects Monograph 34.

Published by Entomology Department, Bishop Museum, Honolulu,

Hawaii. 218 pages, generic and specific check lists and keys, 808 text

figures. Price: $12.50 soft cover; $14.00 hard cover.

The leafhopper Subfamily Coelidiinae contains over 600 species,

nearly 100 genera and 6 tribes. Leafhopper species within this sub-

family are primarily inhabitants of tropical areas. Most of the genera

are found in the Neotropical Region while over half of the species are

known from the Oriental Region and another 50 species are repre-

sented in the Ethiopian Realm. Only two or three of more than 600

species are known from the United States and their origins are

thought to be elsewhere.

This kind of worldwide revision of a large subfamily is a

monumental task, in this case necessitating the publication of the

revision in four parts. The first has been published in the Bulletin of

the British Museum (Natural History) Entomology, Supplement 24,

1975 and is entitled ‘“‘A Revision of the Subfamily Coelidiinae (Homop-

tera: Cicadellidae) Tribes Tinobregmini, Sandersellini and Tharrini.’’ The

remaining two parts will be published elsewhere.

Part Il, which concerns us here, covers the large genus Jhagria with

137 species from the Oriental and Australian Regions and the smaller

genus Tahara from New Guinea. Part Il, as with Part |, is a perfect

example of what a revisionary work should be and for this the author

is to be highly commended. Introductory sections are short, concise

and well written. Ample descriptions of all tribes, genera and species

are given and the keys provided appear quite adequate. Text figures

illustrating the genitalic and cephalic characters necessary for speci-

fic identifications are excellent.

Pacific Insects Monograph #34, along with the other three parts of

the revision of the Coelidiinae will make an excellent and important

addition to the Hompterist’s library. R. J. Gill, California Department of

Food and Agriculture, Sacramento, 95814.

Two New Species of Neotropical Culicoides

(Diptera: Ceratopogonidae)'

Willis W. Wirth?

Systematic Entomology Laboratory, {/Bill, Agr. Res. Serv., USDA,

c/o U.S. National Museum, Washington, D.C. 20560

and

Franklin S. Blanton?

Department of Entomology, University of Florida, Gainesville, Florida 32601

In our studies of Neotropical biting midges we have found two new

species of Culicoides that are especially interesting from distribu-

tional and taxonomic aspects, enough to merit their description and

discussion at this time. We wish to thank Mrs. Molly Griffin for

making the illustrations.

Antennal ratio (abbreviated AR) is the combined length of the five

elongated distal flagellomeres (for convenience referred to as

segments) divided by the combined length of the eight shorter pre-

ceding segments. Palpal ratio (PR) is the length of the third palpal

segment divided by its greatest breadth. Proboscis/Head ratio (P/H

Ratio) is the length of the proboscis measured from the distal end of

the labrum-epipharynx to the anterior margin of the tormae, divided

by the distance measured from the anterior margin of the tormae to

the median hair socket between the eyes. Wing length is measured

from the basal arculus to the wing tip; costal ratio (CR) is the length of

the costa measured from the basal arculus to the tip of the second

radial cell divided by the wing length.

Culicoides kuscheli, new species

(Fig. 1)

Female.—Length of wing 1.21 mm; breadth 0.54 mm.

Head: Eyes (Fig. 1d) contiguous to narrowly separated; bare. Antenna (Fig. 1a) with

lengths of flagellar segments in proportion of 30-22-23-23-24-24-24-25-30-30-30-30-50, AR

0.88, five distal segments very little elongated; sensory pattern 3,7-10. Palpus (Fig. 1b)

with lengths of segments in proportion of 15-25-50-20-23, PR 2.1; third segment

moderately swollen, with a moderately large and deep, round sensory pit. Proboscis

1 This investigation was supported in part by U.S. Army Medical Department Contract no. DA-49-193-MD-

2177.

>Research Associates, Florida State Collection of Arthropods, Division of Plant Industry, Florida

Department of Agriculture and Consumer Services.

The Pan-Pacific Entomologist 54:236-240. July 1978.

VOL. 54, NO. 3, JULY 1978 237

moderately long, P/H Ratio 0.88; mandible with 11-12 teeth.

Thorax: Dark brown, with dense dark grayish pollinosity, without prominent pattern.

Legs brown, without pale rings, Knee spots darker, tarsi paler; hind tibial comb with four

spines, the one nearest the spur longest (Fig. 1e).

Wing (Fig. 1c) with pattern as figured; second radial cell in‘a very dark spot, dark to its

distal extremity; pale spot over r-m crossvein prominent, extending broadly from costal

margin to medial stem. Cell R5 with large poststigmatic pale area and a large irregular

pale spot toward apex of cell, the two separated in posterior part of cell by a very dark

area. Cell M1 with two elongate, indistinct pale spots, more or less streaklike, distally not

reaching wing margin. Cell M2 indistinctly pale on basal half, darker distally with a more

distinct distal pale spot more or less meeting wing margin. Cell M4 with a large pale area

more or less filling distal portion of cell. Anal cell with rather indistinct pale area at base

and a more distinct pale spot in distal portion. CR 0.57; radial cells distinct, the second

relatively broad; macrotrichia rather coarse and moderately sparse, confined to distal half

of wing and in anal cell. Halter pale.

Abdomen: Brownish. Spermathecae (Fig. 1f) two plus a rudimentary third and

sclerotized ring; functional spermathecae ovoid, subequal, measuring 0.047 by 0.033 mm

and 0.043 mm by 0.031 mm.

Male.—Unknown.

Distribution. —Chile.

Types.—Holotype, female, Chile, Antofagasta, Paposo, 6 October 1957, G. Kuschel

(Type no. 72237, USNM). Paratypes, 18 females, same data.

Discussion: This species is named for Guillermo Kuschel,

Entomology Division, Department of Scientific and Industrial

Research, Nelson, New Zealand, in appreciation of his intense

interest in the Chilean biting midge fauna and for his important field

collections.

Culicoides kuscheli does not appear to be closely related to any

known Neotropical species, which is less surprising when one con-

siders that it occurs on the extreme southern range of the genus in

South America. Only one species, C. venezuelensis Ortia and Mirsa,

has been recorded from farther south in Chile, where it was described

as pictipennis (Philippi) from Santiago. There is considerable varia-

ED AD ED TS eS o> OG Or

Fig. 1. Culicoides kuscheli, female: a, antenna; b, palpus; c, wing; d, eye separation; e,

femur and tibia of hind leg; f, spermathecae.

238 THE PAN-PACIFIC ENTOMOLOGIST

tion in wing markings in C. kuscheli, and the figure illustrates the

extreme maximum extent of the pale markings (holotype). In most of

the type series the markings are more or less reduced, and in a few

specimens the distinct pale spots are confined to the marking over

the r-m crossvein, the poststigmatic pale spot, and an oblique pale

mark toward the apex of cell R5. Such specimens will key out in Wirth

and Blanton (1959) to C. alahialinus Barbosa (from Ecuador and

Panama). C. alahialinus differs in having the pale mark in the middle of

cell R5 small and faint and confined to the middle of the cell, the

antennal sensory pattern is 3, 8-10, and the mesonotum has a

prominent pattern of small blackish punctiform dots. Nevertheless,

C. kuscheli is perhaps as closely related to C. al/ahialinus as to any

other Neotropical Culicoides species. Discovery of the male would

certainly help clarify the taxonomic position of C. kuscheli.

Culicoides parascopus, new species

(Fig. 2)

Female.—Length of wing-1.71 mm; breadth 0.81 mm.

Head: Eyes (Fig. 2d) narrowly separated; bare. Antenna (Fig. 2a) with lengths of

flagellar segments in proportion of 38-29-27-27-26-26-25-28-60-60-66-70-96, AR 1.65;

sensory pattern 3,4,6,8,10-15. Palpus (Fig. 2b) with lengths of segments in proportion of

15-35-55-19-23, PR 2.1; third segment moderately swollen, with a moderately large, round,

shallow sensory pit. Proboscis moderately short, P/H Ratio 0.70; mandible with 15 teeth.

Thorax: Dark brown, without prominent pattern. Legs brown, knee spots darker; all

tibiae with narrow pale rings; tarsi paler; hind tibial comb with 4-5 spines, the two nearest

the spur longest, subequal (Fig. 2f).

Wing (Fig. 2c) with pattern as figured; second radial cell in a very dark spot, dark to its

distal extremity; pale spot over r-m crossvein moderately large and circular, bearing a

prominent small dark spot in center lying over r-m crossvein itself. A distinct oval pale

spot straddling vein M2 at its midlength. Cell R5 with a small pale spot on anterior margin

just past tip of costa and a second larger, quadrate pale spot halfway between the former

and tip of cell, the second spot not quite reaching vein M1 caudad. Cell M1 with one small

oval pale spot in distal portion, failing by its own length to reach wing margin. Cell M2

with a small pale spot lying immediately in front of mediocubital fork and a moderately

large rounded pale spot at wing margin in apex of cell. Cell M4 with a large rounded pale

spot nearly filling distal portion of cell. Anal cell with two rounded pale spots in distal

portion. Base of wing with a large pale area extending distad in midportion to about half

the distance to mediocubital fork. CR 0.58; radial cells distinct, the second moderately

broad; macrotrichia rather long and moderately dense, extending nearly to base of wing.

Halter pale.

Abdomen: Brownish. Spermathecae (Fig. 2e) two plus rudimentary third and a faintly

sclerotized narrow ring; functional spermathecae subspherical with short, slender necks,

subequal, each measuring 0.058 by 0.041 mm.

Male.—Similar to female with usual sexual differences; antennal plume well developed,

brownish; last three antennal segments with lengths in proportion of 90-80-85. Genitalia

(Fig. 2h): Ninth sternum short with shallow caudomedian excavation, ventral membrane

not spiculate; ninth tergum moderately long, slightly tapered to long, slender, widely

separated apicolateral processes, the caudal margin between them transverse with a

slight median indentation. Basistyle with ventral and dorsal roots slender and elongate;

VOL. 54, NO. 3, JULY 1978 239

dististyle with slender, pointed, distinctly hooked tip. Aedeagus with basal arch rounded,

extending to half of total length of aedeagus, basal arms slender and curved; distal

median process slightly bulbous proximad and tapering to simple slender tip. Parameres

(Fig. 2g) separate; each with well developed basal knob, slender proximal portion abruptly

bent about 120 degrees with the straight, moderately slender, median stem portion; the

latter gradually tapering distally, the tip abruptly bent ventromesad and ending in asimple

filamentous point.

Distribution.—Mexico.

Types.—Holotype, female, allotype, male, Mexico, Michoacan, Puerto Garnica, 2828

meters, 47 km east of Morelia on Hwy 15, 20 August 1964, F. S. Blanton, light trap (Type

no. 72238, USNM). Paratypes, 4 males, 7 females, same data.

Discussion: The name parascopus is a Latin noun taking its name

because of the close relationship of this species to C. scopus Root

and Hoffman. C. scopus is a more widespread species in higher

elevations ranging from Mexico south to Chiriqui, Panama. It differs

in having the female palpus with a slightly smaller sensory pit;

antennal sensory pattern 3,8,10-15; male aedeagus with short, blunt

tip; and male parameres with the distal stem abruptly narrowed

before the slender, ventrally directed, distal portion bearing apical

fringing hairs.

Culicoides parascopus was taken in a light trap operated at the

border to the Insurgente Morelos National Park in a well developed

montane forest of fir and pine in the Sierra Ozumatlan. Cabrera and

Willink (1973) place this area in their biogeographic ‘Provincia Meso-

americana de Montana’, characterized by rather open forests of pine,

fir and oaks.

eR ke ant

Fig. 2. Culicoides parascopus, a-f female, g-h, male: a, antenna; b, palpus; c, wing; d, eye

separation; e, spermathecae; f, femur and tibia of hind leg; g, parameres; h, genitalia,

parameres removed.

240 THE PAN-PACIFIC ENTOMOLOGIST

Literature Cited

Cabrera, A. L., and A. Willink, 1973. Biogeografia de America Latina. Organization of

American States, Washington, D.C. 120 pages.

Wirth, W. W., and F. S. Blanton, 1959. Biting midges of the genus Culicoides from Panama

(Diptera: Heleidae). Proc. U.S. Nat. Mus. 109: 237-482.

ZOOLOGICAL NOMENCLATURE ANNOUNCEMENT A.N.(S.) 106

The required six months’ notice is given of the possible use of

plenary powers by the International Commission on Zoological

Nomenclature in connection with the following names listed by case

number: (see Bull. Zoo/. Nom. 35, part 1, 31 July, 1978).

680 Blatta germanica Linnaeus, 1767 (Insecta, Dictuoptera,

Blattodea): proposal to conserve and to designate it as type-

species of Blatte/la Caudell, 1903.

2143 Proposal to conserve the specific name tenebricola, as

published in Linyphia by Wider, 1834, but in the sense of

; Kulcezynski, 1887 (Arachnida).

‘2213 HESPERIIDAE Latreille, 1809 (Insecta, Lepidoptera): request

for addition to the Official List.

Comments should be sent in duplicate (if possible within six

months of the date of publication of this notice in Bul!. Zool. Nom. 35,

part 1), citing case number to:

R.V. Melville,

The Secretary,

International Commission on Zoological Nomenclature,

c/o British Museum (Natural History),

Cromwell Road,

LONDON, SW7 5BD,

England.

Those received early enough will be published in the Bulletin of

Zoological Nomenclature.

April issue mailed Sept. 5, 1978.

THE PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

Papers on the systematic and biological phases of entomology are favored, including short notes or articles up to ten

printed pages, on insect taxonomy, morphology, behavior, life history, and distribution. Excess pagination must be approved

and will be charged to the author. Papers are published after acceptance in approximately the order that they are received.

Papers of less than a printed page will be published as space is available, in Scientific Notes.

All manuscripts will be reviewed before acceptance.

Manuscripts for publication, proofs, and all editorial matters should be addressed to the editor.

General. — The metric system is to be used exclusively in manuscripts, except when citing label data on type material, or

in direct quotations when cited as such. Equivalents in other systems may be placed in parentheses following the metric,

i.e. 1370 m (4500 ft) elevation”.

Typing. — Two copies of each manuscript must be submitted (original and one xerox copy or two xerox copies are suitable).

All manuscripts must be type-written, double-spaced throughout, with ample margins, and be on bond paper or an

equivalent weight. Carbon copies or copies on paper larger than 8 ¥2 x 11 inches are not acceptable.

Underscore only where italics are intended in the body of the text. Number all pages consecutively and put authors name on

each sheet. References to footnotes in text should be numbered consecutively. Footnotes must be typed on a separate

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Manuscripts with extensive corrections or revisions will be returned to the author for retyping.

First Page. — The page preceding the text of the manuscript must include (1) the complete title, (2) the order and family in

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Names and descriptions of organisms. — The first mention of a plant or animal should include the full scientific name with

the author of a zoological name not abbreviated. Do not abbreviate generic names. Descriptions of taxa should be in

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Figure legends. — Legends should be typewritten double-spaced on separate pages headed EXPLANATION OF FIGURES

and placed following LITERATURE CITED. Do not attach legends to illustrations.

References. — All citations in text, e.g., Essig (1926) or (Essig 1958), must be listed alphabetically under LITERATURE CITED

in the following format:

Essig, E.0. 1926. A butterfly migration. Pan-Pac. Entomol., 2:211-212.

Essig, E.0. 1958. Insects and mites of western North America. Rev. ed. The Macmillan Co., New York, 1050 pp.

Abbreviations for titles of journals should follow the list of Biological Abstracts, 1966, 47(21):8585-8601. For Scientific Notes

the citations to articles will appear within the text, ie... . “Essig (1926, Pan Pac. Entomol., 2:211-212) noted ...”.

Proofs, reprints, and abstracts. — Proofs and forms for the abstract and reprint order will be sent to authors. Major changes

in proof will be charged to the author. Proof returned to the editor without the abstract will not be published.

Page charges. — All regular papers of one to ten printed pages are charged at the rate of $18.00 per page. Private investigators

or authors without institutional or grant funds may apply to the society for a grant to cover a portion of the page charges. In

no case will society grants subsidize more than two thirds of the cost of page charges. Individuals receiving a society

subsidy thus will be billed a minimum of $6.00 per page. Pages additional to the first ten are charged at the rate of $30.00 per

page, without Subsidy. Page charges are in addition to the charge for reprints and do not include the possible charges for

extra pagination or the costs for excessive changes after the manuscript has been sent to the printer.

Reprint costs. — Current charges for reprints are approximately as listed below. These charges are subject to change, and

authors will be billed at the rate in effect at the time of publication.

1-2pp 2%-4 pp 4%-8 pp 8%-12 pp 12%-16 pp

1st 100 22.00 32.00 40.00 70.00 75.00

Additional 100’s 12.00 18.00 25.00 35.00 40.00

PUBLICATIONS

OF THE |

PACIFIC COAST ENTOMOLOGICAL SOCIETY

PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL

SOCIETY.

Vol. 1 (16 numbers, 179 pages) and Vol. 2 Suwa 1-9, 131

pages). 1901-1930. Price $5.00 per volume.

PAN-PACIFIC ENTOMOLOGIST.

Vol. 1 (1924) to Vol. 51 (1975), price $10.00 per volume of 4

numbers, or $2.50 per single issue. Vol. 52 (1976) and subsequent

issues, $15.00 per volume or $3.75 per single issue.

MEMOIR SERIES.

Volume 1. The Sucking Lice by G. F, Ferris. 320 pages. Pub-

lished October 1951. Price $6.00 (Plus 35c postage.)*

Volume 2. The Spider Mite Family ‘Tetranychidae by A. Earl

Pritchard and Edward W. Baker. 472 pages. Published July _—

Price $10.00 (Plus 50c postage. Ip

Volume 3. Revisionary Studies in the Nearctic Decticinae by David

C. Rentz and James D. Birchim. 173 pages. Published July 1968.

Price $4.00. (Plus 25c postage. )*

Volume 4. Autobiography of an Entomologist by Robert L. Usinger.

343 pages. Published August 1972. Price $15.00. (Plus 40c

postage.)* as :

Volume 5. Aevision of the Millipede Family Andrognathidae in the

Nearctic Region by Michael R. Gardner. 61 pages. Published

January 21, 1975. Price $3.00. (Plus 25c postage.)*

*(Add 6% sales tax on all California orders (resident of Alameda, Contra Costaand

San Francisco counties add 612%). Members of the Society will receive a 20%

discount)

Send orders to:

Pacific Coast Entomological Society

c/o California Academy of Sciences

Golden Gate Park

San Francisco, California 94118