Vol. 51

JANUARY 1975

No. 1

THE

Pan-Pacific Entomologist

NOONAN — Bionomics, evolution, and zoogeography of members of the genus

Dicheirus (Coleoptera: Carabidae) 1

ALLEN — Ephemerella ( Cincticostella ) : a revision of the nymphal stages

(Ephemeroptera: Ephemerellidae) 16

ELLIS — Seasonal abundance and distribution of adult stoneflies of Sashin

Creek, Baranof Island, southeastern Alaska (Plecoptera) 23

HOFFMAN — An arboreal cleidogonid milliped from Chiapas (Chordeumida :

Cleidogonidae) 31

WANGBERG — Biology of the thimbleberry gallmaker Diastrophus kincaidii

(Hymenoptera: Cynipidae) 39

HALL— The North American species of Triploechus Edwards (Diptera:

Bombyliidae) 49

KIRK — Siricid woodwasps and their associated parasitoids in the south-

western United States (Hymenoptera: Siricidae) 57

COOK — A reconsideration of the Nearctic Rhexoza (Diptera: Scatopsidae) .. 62

YOUNG — Observations on the life cycle of Heliconius hecale zuleika (Hewit-

son) in Costa Rica (Lepidoptera: Nymphalidae) 76

LINSLEY — Fifty years of the Pan-Pacific Entomologist (Concluded) 86

SCIENTIFIC NOTE .' 30

BOOK NOTICE 48

RECENT LITERATURE 15, 85

ZOOLOGICAL NOMENCLATURE 38

PROCEEDINGS of the PACIFIC COAST ENTOMOLOGICAL SOCIETY .... 88

SAN FRANCISCO, CALIFORNIA • 1975

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

THE PAN-PACIFIC ENTOMOLOGIST

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The Pan-Pacific Entomologist

Vol. 51

January 1975

No. 1

Bionomics, Evolution, and Zoogeography of Members

of the Genus Dicheirus

(Coleoptera: Carabidae)

Gerald R. Noonan

Department of Entomology, The University of Alberta, Edmonton, Alberta,

Canada T6G 2E3

The genus Dicheirus contains 6 species and subspecies: D. hrunneus

(Dejean), D. dilatatus dilatatus (Dejean), D. dilatatus angulatus Casey,

D. obtusus LeConte, D. piceus (Menetries), D. strenuus (Horn)

(Noonan, 1968). Evolution of the genus, with examination of other

genera of the subtribe Anisodactylina are given by Noonan (1973,

1974). The present paper provides a detailed analysis of the evolu-

tionary and zoogeographic relationships of the members of Dicheirus

and includes additional bionomic information for some species.

Bionomics

The following statements are based on field observations of adults

primarily in northern and central California but also in southern

California, Oregon, and British Columbia.

Members of Dicheirus are adapted to grassy habitats lacking trees

or with scattered trees such as in woodlands or along grassy forest

edges. On rare occasions D. dilatatus dilatatus and D. piceus occur in

moderately moist and possibly slightly brackish grassy areas near ocean

bays.

Species occurring in forest climax areas inhabit natural clearings in-

side forests, the grassy edges of forests, and in man-made clearings. The

clearing of forests to create pastures provides additional habitats for

such species, and adults are more numerous in pastures than in many

natural clearings. Pastures are probably more favorable habitats than

many natural clearings due to: discontinuity and small size of many

natural clearings; unfavorable factors (such as flooding in clearings

formed by streams) present in many natural clearings; and additional

The Pan-Pacific Entomologist 51: 1-15. January 1975

The Pan-Pacific Entomologist

shelter in pastures such as dried cow droppings, fallen fence posts and

other debris. Dried cow droppings are especially beneficial to adults;

pastures with them contain more individuals than similar areas without.

The ground beneath dried cow droppings is often moister and cooler

than that adjacent to them, and adults aggregate in these moister, cooler

microhabitats.

In central and northern California, teneral adults appear from January

to March and are active with adults from the previous year until the

ground and vegetation dry during late April to June. Except for iso-

lated lowland areas and mountainous areas in which the ground and

vegetation dry later in the year or remain moist, adults are uncommon

from approximately June to September. In September or October

teneral adults appear, soon mature, and are active with previously

matured adults.

In central and northern California an early activity period of adults

coincides with winter rains, is reduced during the dry summer, and is

renewed prior to and probably also during fall rains. Rainfall is the

major factor influencing adult activity. During exceptionally dry years

adults of D. strenuus and D. dilatatus dilatatus do not appear in the fall,

and adults of D. piceus are greatly reduced in number. Additional field

studies may reveal that adults of other forms also do not appear or are

reduced in numbers in the fall of dry years.

Specimens of Dicheirus have not been observed flying. However, all

species but D. strenuus have full and apparently functional hind wings.

The winged species may have diurnal spring dispersal flights. I have

noted in the spring while collecting in very small natural clearings that

on warm days adults suddenly appear in areas that had been intensively

examined (debris and top 3 or 4 inches of soil removed) just previously.

Darlington (1943) suggested that flight is advantageous to species of

Carabidae which inhabit relatively large areas with sparse unstable

populations. Before the forest clearing activities of man, species of

Dicheirus presumably were restricted to natural clearings in forest cli-

max areas. Such clearings would have been sparsely distributed, and

often small and unstable. Full-winged species probably remained so

because of the importance of flight as a dispersal mechanism. The lack

of flight records for Dicheirus may be due to flight being diurnal and re-

stricted to early spring. Most Carabid flight records are based on cap-

tures at lights ; if Dicheirus adults fly only during the day, then they will

not aggregate at lights. Furthermore most insect collectors probably do

not begin field work until after early spring.

Vol. 51, No. 1 , January 1975

Table 1. Plesiomorphic and apomorphic character states used in

Fig. 1.

Character

State

No. Character

plesiomorphic

apomorphic

Thorax

1 pronotal

moderate to

glabrous to sparse

pubescence

dense throughout

medially

2 proepisternal

absent or very

large and prominent

punctures

small

Legs

3 venter of

with spongy

glabrous in most

articles 1 to

pubescent

specimens, partial

4 of $ foretarsi

vestiture

to complete spongy

pubescent vestiture

in some specimens

4 mid and hind

without prominent

with prominent

tibia

tubercules at base

of spines along

sides

of spines along sides

5 $ hindtibial

lanceolate shaped

slightly laterally

apical spurs

expanded

a) spatulate shaped

Elytra

6 pubescence

short

varied from short

length

to long

a) long in all

specimens

7 type of

adpressed

erect

pubescence

Male genitalia

8 median lobe

not twisted

twisted

Most forms of Dicheirus occur not only in the same geographic area

(Figs. 2-6) but, at least in the adult stage, in the same microhabitat.

In California I have taken adults together in the same hole beneath

debris on the ground as follows: D. brunneus , D. dilatatus dilatatus, and

D. piceus in Trinity County; and D. strenuus and D. piceus in Kern

and Tulare Counties. Additional collecting will probably demonstrate

that adults of all sympatric forms occur in the same microhabitats.

The Pan-Pacific Entomologist

Evolution

The principles used in evolutionary reconstructions are slightly

modified from those of Hennig (1966) as discussed in detail in Noonan

(1973). Figure 1 and Table 1 present the most probable hypothesis of

evolution of the species and subspecies. All sister taxa in Figure 1 are

linked by synapomorphies. Convergence in a character state has oc-

curred once; all specimens of D. dilatatus angulatus and some specimens

of D. brunneus have long elytral setae. A sequence of evolution other

than that postulated in Figure 1 and Table 1 would not be consistent with

available data.

Contemporary Zoogeography

Species are restricted (Figs. 2-6) to western North America, probably

due to limited adaptation to continental climate. Eastern limits are

represented by specimens of D. piceus from Salt Lake County, Utah.

Northern and southern limits are set by D. piceus in southern British

Columbia and on Guadalupe Island off the western coast of Baja Cali-

fornia. Guadalupe Island has a persistent summer fog belt in the

summit section (Axelrod, 1967) and a milder, moister climate than the

immediately adjacent mainland (Savage, 1967). These climatic factors

probably explain survival of D. piceus on Guadalupe Island, approxi-

mately 220 miles farther south than mainland records of D. dilatatus

angulatus in northwestern Baja California. Dicheirus dilatatus dilatatus

and D. piceus occur on Santa Catalina and San Clemente Islands with

D. piceus also present on Guadalupe Island. Distances of these islands

from the mainland are (Philbrick, 1967) 20, 49 and 157 miles respec-

tively.

Table 2 lists range extents (the linear distance between the 2 most

distant occurrences of each form) of the 6 species and subspecies of

Dicheirus. Previous workers such as Ball and Freitag (1969), Erwin

(1970), Noonan (1973), and Whitehead (1972) used range extents to

elucidate refugia, and centers of evolution, and to discuss probable adapt-

ability of wide ranging versus narrow ranging forms. The small number

of forms in Dicheirus and sympatry of most of them make it impossible

to stipulate a present center of evolution other than the area now oc-

cupied by these forms.

The relatively large range extents of D. piceus , D. obtusus and D.

dilatatus suggest that these 3 species are more tolerant than other forms

to environmental extremes. Only D. piceus and D. dilatatus dilatatus

show a possible tolerance to brackish habitats near ocean bays and are

Vol. 51, No. 1 , January 1975

Fig. 1 . Postulated evolution of species and subspecies of Dicheirus (hollow

rectangles denote plesiomorphic character states, half black rectangles denote

intermediate apomorphic character states, fully black rectangles denote apomorphic

character states) .

also the only forms recorded from islands. Dicheirus piceus also exists

in habitats from which other forms seem excluded.

Dicheirus brunneus has a 450 mile range extent and is apparently

adapted only to mesic environmental conditions in northern California

and Oregon.

Dicheirus strenuus and D. dilatatus angulatus have restricted range

extents of 73 and 30 miles respectively. Dicheirus strenuus is the only

taxon with vestigial hind wings; this condition probably explains in part

its restricted range from the Tehachapi and Greenhorn Ranges, Kern

and Tulare Counties, California. This species may be adapted to only a

narrow range of environmental conditions since it does not follow

woodlands or grasslands down into the San Joaquin Valley or high up

into the Sierras as do all other forms present in central California.

The Pan-Pacific Entomologist

Table 2. Range extents of species of Dicheirus.

Species or Subspecies

Maximum Range Extent

(in miles)

D. piceus

1700 (Guadalupe Island

included)

1200 (mainland only)

D. obtusus

800

D. dilatatus (both subspecies)

750

D. dilatatus dilatatus

650

D. dilatatus angulatus

D. brunneus

450

D. strenuus

Possibly, D. strenuus evolved in its present restricted range. Alterna-

tively, it may formerly have been more widespread, and its low

dispersal ability may have resulted in its elimination from much of its

range during Pliocene and Pleistocene climatic changes.

Dicheirus dilatatus angulatus is known only from coastal areas in

San Diego County, California and northern Baja California. The factors

responsible for such a restricted range are unknown. Possibly D. d.

angulatus occurs further south in Baja California, but has not been

collected.

Historical Zoogeography

Taxa of Dicheirus demonstrate considerable sympatry (Figs. 2-6),

and adults of several species share microhabitats in central and northern

California. This sympatry and ecological similarity suggest that

extant species have existed long enough to acquire isolating mechanisms

enabling them to share the same geographical ranges and, at least as

adults, the same microhabitats. The general lack of orderly patterns of

vicariance suggest that species may have evolved elsewhere than their

present ranges.

Species of Dicheirus are now concentrated from approximately south-

ern California to Oregon, but initial evolution within the genus probably

occurred in Alaska and northern Canada (Noonan, 1973). According

to these ideas, the ancestor of Dicheirus crossed from Eurasia into

North America via the Bering Land Bridge during the early Tertiary,

probably during Eocene when the Bering Land Bridge was present

(Hopkins, 1967), and the climate there was temperate (Dorf, 1960).

Eocene tropical and subtropical climates in the United States (Dorf,

1960) may have prevented initial dispersal southward. However, as

Vol. 51, No. 1, January 1975

Figs. 2-6. Distribution of: (2) D. obtusus ; (3) D. piceus ; (4) D. strenuus ; (5)

D. dilatatus dilatatus (circles) and D. dilatatus angulatus (triangles) ; (6) D.

brunneus.

The Pan-Pacific Entomologist

climates cooled during Oligocene, Miocene and especially Pliocene,

there were probably dispersals south into the area now occupied by

species of Dicheirus. As climates continued to cool, Dicheirus became

extinct in northern Canada and Alaska. The present irregular and

mostly sympatric distribution of species is thus probably due to rela-

tively old species being forced south from their center of evolution.

Eastward dispersal of Dicheirus was probably prevented by in-

tolerance of continental climates and to a lesser extent by the Brooks

Range and Rocky Mountains. The restrictive climatic factor may have

been the past and present seasonal distribution of precipitation. Along

the Pacific Coast precipitation is concentrated in winter, and severe

drought regularly coincides with summer heat; inland there is a strong

summer maximum of precipitation (Wells, 1970). This difference in

climatic rhythm may also be the reason why many carabid species of

interior grasslands (such as those of the subgenus Glanodes, genus

Harpalus , see Ball, 1972) have not extended their ranges to the Pacific

Coast.

The sympatric dilatatus and piceus stocks probably evolved during

Eocene to Miocene north of the present range of Dicheirus. Dicheirus

strenuus and D. dilatatus may be vicariant in that D. dilatatus is not

recorded from the range of D. strenuus. However, D. dilatatus surrounds

the range of D. strenuus on the north, south and west. I know of no past

or present barriers between the known species ranges; additional

collecting will probably establish sympatry.

Dicheirus dilatatus dilatatus and D. dilatatus angulatus are separated

by a gap of approximately 100 miles in southern California (Fig. 5).

Evolution of these subspecies is probably due to the coast corridor

effect described by Peabody and Savage (1958) for amphibians and

reptiles. During Pliocene and early Pleistocene (Fig. 7) the San

Joaquin Valley consisted of a flooded embayment opening to the ocean

by a wide strait in northern Baja California. Populations of D.

dilatatus were probably isolated east of the embayment and evolved

into D. dilatatus angulatus which has several apomorphies suggesting

selection was acting on a small gene pool. Populations west of the

San Joaquin embayment probably exchanged genes northward via the

coast corridor with other populations and were thus part of a relatively

large gene pool. These populations retained plesiomorphic features;

their descendants constitute the nominate subspecies with relatively

plesiomorphic features. This postulate, that the subspecies with apo-

morphic features was derived from a smaller gene pool than the sub-

Vol. 51, No. 1, January 1975

Fig. 7. Pliocene and early Pleistocene paleogeography of California, showing

archipelagic nature of Coast Range region, and presence of strait connecting

Pacific with San Joaquin embayment. Cross hatching indicates flooded areas.

(Adapted from Peabody and Savage, 1958.)

The Pan-Pacific Entomologist

species with plesiomorphic ones, agrees well with Brundin’s (1972)

description of speciation by cleavage.

As the San Joaquin embayment dried, the climate of eastern southern

California probably became drier and warmer; populations of D.

dilatatus angulatus probably migrated westward toward cooler, moister

coastal regions. The Xerothermic Period which lasted from 8,000 to

3,000 years ago constituted a time of warm climate (Axelrod, 1967)

which placed particular stress on organisms along the Pacific Coast

where precipitation is concentrated in the winter and severe drought reg-

ularly coincides with summer heat (Wells, 1970) . Any populations of D.

dilatatus angulatus which remained in eastern southern California after

drying of the San Joaquin embayment probably were eliminated by

stresses of the Xerothermic. The presence of D. dilatatus dilatatus on

San Clemente and Santa Catalina Islands and the absence there of D.

dilatatus angulatus supports the above hypothesis of evolution and dis-

persal. If D. dilatatus angulatus had been present along the coast during

the Pliocene and early Pleistocene, it probably would have migrated to

these 2 islands as did the nominate subspecies.

Species in Carabid genera other than Dicheirus may also have

cleaved into subspecies or sister species due to the coast corridor effect.

Calathus ruficollis Dejean has a distribution similar to that of the

genus Dicheirus and is found in habitats similar to those of Dicheirus

(Ball and Negre, 1972). The coast corridor effect probably explains

the differentiation of C. 7'uficollis ignicollis Casey from the nominate

subspecies.

If the 2 subspecies of D. dilatatus evolved as a result of the San

Joaquin embayment, then they have had approximately 2 million years

since its disappearance in which to become parapatric. All apomorphic

character states of D. dilatatus angulatus but long elytral setae vary

toward the plesiomorphic state found in the nominate subspecies, and

some individuals have the plesiomorphic state for one or more charac-

ters. This variation suggests that the 2 subspecies were parapatric in

the past and had limited gene exchange. Such parapatry probably took

place during cooler, moister (Axelrod, 1967) Pleistocene glacial periods.

My impression of the area now separating the 2 forms (Fig. 7) is that

it does not have favorable habitats for Dicheirus ; D. piceus , the most

widespread species in the genus, is absent from this area but is found

both to the south and north. Mainland records of D. dilatatus dilatatus

north of the Tehachapi Mountains are mostly based on moderate to ex-

tensive series of specimens ; records south of these mountains are mostly

based on only a few specimens per locality. This apparent reduction in

Vol. 51, No. 1, January 1975

abundance south of the Tehachapi Mountains suggests that this sub-

species is not well adapted to conditions in southern California. Finally

D. dilatatus angulatus survives only in a small refugium along the coast

from San Diego County, California to northern Baja California. In-

creasing warmth and aridity after Pleistocene glacial periods (Axelrod,

1967) probably eliminated all Dicheirus from the present gap between

the subspecies of D. dilatatus.

Rates of differentiation in D. dilatatus can be estimated from geologi-

cal dating of the San Joaquin emhayment which existed from approxi-

mately 8 to 2 million years ago (dates personal communication from R.

Fox). Thus, the subspecies are a maximum of 7 million years old and

took 6 million or less years in which to evolve.

Whitehead (1972) calculated the period of time required for specia-

tion in Schizogenius and Brachius by estimating the age of each group

and then counting the number of bifurcations in the reconstructed phy-

logenies. Such calculations are prone to error both from mistakes in the

estimated ages of groups and from errors in the reconstructed phy-

logenies which most often are simplifications of complex processes which

occurred in nature (Noonan, 1973). When fossils are not available, the

most reliable estimates of time required for speciation or subspeciation

are those based on geologic dating of barriers which isolate or once

isolated vicariant sister forms.

Whitehead calculated the average time required for speciation in

North American Schizogenius and Brachinus carabids to be approxi-

mately 3 million years. Geological dating of the San Joaquin emhayment

is not exact; the period of isolation of populations of D. dilatatus may

have been less than 6 million years, but certainly not less than 3 million.

The brunneus and piceus lineages are sympatric as are the species

D. brunneus , D. piceus and D. obtusus ; little can be concluded other

than that these taxa probably evolved during the Eocene to Pliocene

north of the present range of Dicheirus.

Present Utah populations of D. piceus are separated from the main

species range by nearly 300 miles of Great Basin Desert. Precipitation

in lower elevations of this desert is less than 4 inches per year while

January/July temperatures average 44.5°/86°F at Las Vegas, Nevada,

32.5°/71°F at Reno, Nevada, and 30°/77°F at Salt Lake City, Utah

(Morrison, 1965). This dry and moderately warm climate probably

bars dispersal between the main range and the Utah populations, pre-

sumably localized in mountains east of Salt Lake City. However, Mio-

cene and Pliocene climates of the Great Basin were subhumid with 25 to

30 inches per year of estimated precipitation, and the present desert

The Pan-Pacific Entomologist

climate did not begin until early Pleistocene (Morrison, 1965). Pleisto-

cene glacial periods resulted in temperatures 8° to 15° F cooler than now,

more rainfall, and shifts of life zones hundreds of miles north and south

and thousands of feet up and down mountains. Before Pleistocene devel-

opment of desert climate and during Pleistocene cool, moist periods,

populations of D. piceus probably invaded the Great Basin Deserts and

reached Utah. The desert climate of the present and of the interglacial

periods probably eliminated Dicheirus from Great Basin lowlands and

isolated populations of D. piceus in mountains presently moister and

cooler than lowlands.

An alternative explanation for the presence of isolated populations of

D. piceus in Utah is as follows. Before the development of desert climate

and/or during Pleistocene glacial periods, populations of D. piceus (and

possibly other species of the genus) may have dispersed around the

northern periphery of the Great Basin Desert and then south into Utah

via foothills of the Rockies. However, such dispersal would require

greater tolerance to continental climate than now shown by species of

Dicheirus and probably would have resulted in isolated populations

of D. piceus in Montana, Wyoming, and Colorado. Collecting has not

demonstrated the existence of such populations; dispersal directly

across the Great Basin Desert seems more probable than dispersal along

the northern periphery and Rockies.

The effects of Quaternary glacial periods on western North America

are not as well elucidated as for the East. However, various workers

(Crandell, 1965; Martin and Mehringer, 1965; Morrison, 1965; Wahr-

haftig and Birman, 1965) agree that temperatures were cooler, precipi-

tation greater, and that biotic zones shifted downward. Such climatic

changes may have resulted in migrations south into Baja California.

Continental climates probably prevented migrations southeast into main-

land Mexico. If migrations had reached mainland Mexico, relict popula-

tions of Dicheirus would probably occur in the highlands as do other

members of the subtribe Anisodactylina (Noonan, 1973).

The presence of D. dilatatus dilatatus and D. piceus on Santa Catalina

and San Clemente Islands and of D. piceus on Guadalupe Island may

be due to dispersal across present water gaps by wind or by rafting.

Dispersal to the first 2 islands could not have been earlier than their

formation during or after the mid-Pliocene (Valentine and Lipps, 1967).

Ocean barriers between the mainland and these 2 islands were absent

(Clements, 1955, as reported by Valentine and Lipps, 1967) or at least

much narrower during late Pliocene and early Pleistocene (Valentine

and Lipps, 1967). Colonization probably took place then. Migration

Vol. 51, No. 1, January 1975

onto other Channel Islands may also have occurred at that time. The

lack of records of Dicheirus from other California Channel Islands may

be due to lack of collecting or to chance extinction.

Guadalupe Island possibly was never connected to the mainland

(Garth, 1967), and D. piceus probably colonized it by long distance

dispersal over water. However, geological data regarding mainland con-

nections of Guadalupe are somewhat inconclusive, and colonization via

a land bridge remains a possibility.

The above discussion illustrates that species of Dicheirus are mostly

sympatric and probably evolved north of their present ranges. It

also demonstrates the difficulty of elucidating place or time of origin of

forms which do not exhibit vicariance or have a fossil record.

Acknowledgments

I thank G. E. Ball for reading a preliminary draft of this paper,

J. S. Scott for preparing the illustrations, K. Benschop for typing the

final manuscript copy.

Literature Cited

Axelrod, D. I. 1967. Geologic history of the Californian Insular Flora, p. 267-

315. In R. N. Philbrick (ed.), Proceedings of the Symposium on the

Biology of the California Islands. Santa Barbara Botanic Garden,

Santa Barbara, California.

Ball, G. E. 1972. Classification of the species of the Harpalus subgenus

Glanodes Casey (Carabidae: Coleoptera) Coleopt. Bull., 26(4): 179-

204.

Ball, G. E. and R. Freitag. 1969. Zoogeography, p. 174^-190. In R. Freitag,

A revision of the species of the genus Evarthrus LeConte (Coleoptera:

Carabidae). Quaest. entomol., 5: 89-212.

Ball, G. E. and J. Negre. 1972. The taxonomy of the Nearctic species of the

genus Calathus Bonelli (Coleoptera: Carabidae: Agonini). Trans.

Amer. Entomol. Soc., 98: 412-533.

Brundin, L. 1966. Transantarctic relationships and their significance as evi-

denced by chironomid midges, with a monograph of the subfamilies

Podonominae and Aphroteniinae and the austral Heptagyiae. K. svenska

Vetensk. Akad. Handl., (4)11: 1-472.

Brundin, L. 1972. Evolution, causal biology, and classification. Zool. Scripta, 1:

107-120.

Clements, T. 1955. The Pleistocene history of the Channel Island Region,

southern California, p. 311-323. In Essays in the natural sciences

in honor of Captain Allan Hancock. Los Angeles. (Citation from

Valentine and Lipps, 1967.)

Crandell, D. R. 1965. The glacial history of western Washington and Oregon.

p. 341-353. In H. E. Wright and D. G. Frey (ed.), The Quaternary of

the United States. Princeton University Press, New Jersey.

The Pan-Pacific Entomologist

Darlington, P. J., Jr. 1943. Carabidae of mountains and islands: data on the

evolution of isolated faunas, and on atrophy of wings. Ecol. Monogr.,

13: 37-61.

Dorf, E. 1960. Climatic changes of the past and present. Am. Sci., 48(3):

341-364.

Erwin, T. L. 1970. A reclassification of bombardier beetles and a taxonomic

revision of the North and Middle American species (Carabidae:

Brachinida). Quaest. entomol., 6(4): 1-215.

Garth, J. S. 1967. Introduction to Insular Zoology, p. 181-183. In R. N.

Philbrick ( ed. ) , Proceedings of the Symposium on the Biology of the

California Islands. Santa Barbara Botanic Garden, Santa Barbara,

California.

Hennig, W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana,

Illinois. 263 p.

Hopkins, D. M. 1967. The cenozoic history of Beringia — A synthesis, p. 451-

484. In D. M. Hopkins (ed.), The Bering Land Bridge. Stanford

University Press, Stanford, California.

Martin, P. S. and P. J. Meliringer. 1965. Pleistocene pollen analysis and

biogeography of the Southwest, p. 433-451. In H. E. Wright and

D. G. Frey (ed.), The Quaternary of the United States. Princeton

University Press, New Jersey.

Morrison, R. B. 1965. Quaternary geology of the Great Basin, p. 265-285.

In H. E. Wright and D. G. Frey (ed.), The Quaternary of the United

States. Princeton University Press, New Jersey.

Noonan, G. R. 1968. A revision of the genus Dicheirus Mannerheim 1843 (Col.

Carabidae). Opus, entomol., 33: 281-304.

Noonan, G. R. 1973. The Anisodactylines (Insecta: Coleoptera: Carabidae:

Harpalini) : classification, evolution and zoogeography. Quaest. en-

tomol., 9(4) : 266-480.

Noonan, G. R. 1974. Allendia, a new South American genus with notes on its

evolutionary relationships to other genera of Anisodactylina (Coleoptera:

Carabidae: Harpalini). (Manuscript.)

Peabody, F. E. and J. M. Savage. 1958. Evolution of a coast corridor in Cali-

fornia and its effect on the origin and dispersal of living amphibians and

reptiles, p. 159-186. In C. L. Hubbs (ed.), Zoogeography. Publ. 51.

American Association for the Advancement of Science, Washington,

D. C.

Philbrick, R. N. 1967. Introduction, p. 3-8. In R. N. Philbrick, (ed.),

Proceedings of the Symposium on the Biology of the California Islands.

Santa Barbara Botanic Garden, Santa Barbara, California.

Raven, P. H. 1967. The floristics of the California Islands, p. 57-67. In

R. N. Philbrick (ed.), Proceedings of the Symposium on the Biology

of the California Islands. Santa Barbara Botanic Garden, Santa Barbara,

California.

Savage, J. M. 1967. Evolution of the insular herpetofauna. p. 219-227. In

R. N. Philbrick (ed.), Proceedings of the Symposium on the Biology of

the California Islands. Santa Barbara Botanic Garden, Santa Barbara,

California.

Valentine, J. W. and J. H. Lipps. 1967. Late Cenozoic history of the Southern

Vol. 51, No. 1 , January 1975

California Islands, p. 21-35. In R. N. Philbrick ( ed. ) , Proceedings of

the Symposium on the Biology of the California Islands. Santa Barbara

Botanic Garden, Santa Barbara, California.

Wahrhaftig, C. and J. H. Birman. 1965. The Quaternary of the Pacific Moun-

tain System in California, p. 299-340. In H. E. Wright and D. G. Frey

(ed.), The Quaternary of the United States. Princeton University Press,

New Jersey.

Wells, P. V. 1970. Postglacial vegetational history of the Great Plains. Science,

167: 1574-1582.

W hitehead, D. R. 1972. Classification, phylogeny, and zoogeography of Schizo-

genius Putzeys (Coleoptera: Carabidae: Scartini). Quaest. entomol.,

8(3): 131-348.

RECENT LITERATURE

The Cud Beetles of California (Coleoptera : Cud ae) . John F. Lawrence.

Bulletin of the California Insect Survey. Volume 17, 41 pp., 7 figs., 15 maps.

$2.25.

This summary of the biology, systematics and geographic distribution of the

California representatives of an abundant but little-noticed family of fungus-

inhabiting beetles could well be used as a model for similar faunal surveys. The

systematics section is prefaced by concise but informative discussions of bionomics,

host and habitat preference, seasonal occurrence, and patterns of geographic

distribution, and supplemented by tables summarizing information that would have

required many pages if presented as raw data. The taxonomic treatment includes

introductory explanations of important characters, as well as brief differentiations

from similar beetles which occur in bracket fungi and might be confused with

Ciidae. The excellent keys to genera and species utilize features which mostly

show clear, qualitative differences. Difficult characters are quantified, especially

in keys to species. Maps showing characteristic types of distributions are provided

for about one half the species. All maps have insets depicting the entire North

American distribution, a very helpful addition which requires no additional space.

The 37 figures include line drawings of most of the important taxonomic characters,

and excellent half tone illustrations of 14 of the 27 species known or suspected to

occur within the political boundaries of California.

A comprehensive summary of this nature might well alert ecologists to the

opportunities these beetles offer for studies of competition, habitat preference and

partitioning or species diversity and niche breadth. — Editor.

The Pan-Pacific Entomologist

Ephemerella (Cinctico Stella) : A Revision of

the Nymphal Stages

( Ephemeroptera : Ephemerellidae)

Richard K. Allen 1

Department of Biology, California State University

5151 State University Drive, Los Angeles, California 90032

The following treatment includes a characterization of the subgenus

Cinctico Stella, a description, or diagnosis, of the nymphal stages, a key

to the nymphs, and the known distributional limits of each species.

Ephemerella ( Cincticostella ) orientalis Tshernova, 1952, whose nymph

is unknown is included with complete synonymy and known distribution.

Grateful appreciation is expressed to Professors Kazimitzu Kato and

Valentina Zaydman, California State University, Los Angeles, for trans-

lating parts of the papers by Imanishi (1940) and Tshernova (1972),

respectively.

Subgenus Cincticostella Allen

ftigra-group Imanishi 1938: 33.

Cincticostella Allen 1971: 513; Tshernova 1972: 614 (— Asiatella) .

Asiatella Tshernova 1972: 611.

Imanishi (1938) placed E. nigra Ueno, 1928, in the nigra-group of

the genus Ephemerella and Allen (1971) erected the subgenus Cincti-

costella for E. nigra and five other species. He transferred E. levanido-

vae Tshernova, 1952, and described E. castanea, E. delicata , E. imanishii ,

and E. insolta. Tshernova (1972) described the genus Asiatella to

include E. nigra, E. femorata Tshernova, 1972, E. orientalis , E.

levanidovae , and E. tshernovae Bajkova, 1962, to this taxon. In the

same paper, as a footnote, she synonymized Asiatella with Cincti-

costella and placed E. imanishii as a junior synonym of E. tshernovae.

Cincticostella is confined to southeast and eastern Asia. Ephemerella

boja n. sp., E. gosei n. sp., and E. insolta are known from Thailand;

E. castanea from Korea; E. delicata from Japan and Manchuria; E.

femorata from Vietnam; E. levanidovae and E. orientalis from Russia;

E. nigra from Japan; and E. tshernovae from Japan, Korea, and Russia.

The number of described species in the subgenus is ten.

The species of Cincticostella are all known from the nymphal stage

except E. orientalis which was described from male and female sub-

1 The research upon which this report is based was supported by National Sciences Foundation

Grant No. GB-35591.

The Pan-Pacific Entomologist 51: 16-22. January 1975

Vol. 51, No. 1 , January 1975

imagoes. Ephemerella nigra is the only species known from the nymph

and the male imago.

Nymphal Stage . — Body flattened ventrally, convex dorsally. Head round to

quadrangular; genae often truncate; maxillae without apical canines, with apical

tuft of setae; maxillary palpi well developed, vestigial, or absent. Fore femora

with band of transverse spines near middle; denticles on tarsal claws variable in

number. Anterolateral corners of prothorax produced anteriorly; mesothorax ex-

panded laterally in anterior portion. Abdominal terga with paired dorsal sub-

median tubercles; abdominal segments 7-9 with or without well developed pos-

terolateral projections; abdominal terga 3-7 with lamellate, imbricated gills.

Caudal filaments shorter than body.

On the basis of nymphal characters two groups of related species

are recognized. Ephemerella insolta , E. boja, and E. femorata form

the insolta- group. These species are characterized by possessing suboc-

cipital head tubercles, and the second and third pairs of femora are

enlarged with serrated margins and/or protuberances. The nigra-group

includes E. nigra , E. castanea , E. delicata, E. gosei, E. levanidovae, and

E. tshernovae. These species are without head tubercles and the second

and third pairs of femora are narrow, not enlarged, and the margins

are entire. Ephemerella orientalis cannot be placed to species-group

until the nymphal stage is discovered.

The characters most useful for distinguishing the species of Cinctico-

stella nymphs are as follows : ( 1 ) the number and degree of development

of the paired dorsal abdominal tubercles; (2) the degree of expansion,

the presence or absence of marginal serrations, and the presence or

absence of spines and tubercles on the anterior surfaces of the middle

and hind femora; (3) the degree of development of the posterolateral

projections on abdominal segments 7-9; (4) the shape of the genae;

(5) the color patterns on the head and body; and (6) the relative

lengths of the body and caudal filaments.

The following key will serve to distinguish the known nymphs of the

species of Cincticostella. Illustrations are not included but reference

is made to previously published figures which are enclosed in paren-

theses in the couplets. That is, (Allen 1971: fig. 11) in couplet 1 makes

reference to “New Asian Ephemerella with notes (Ephemeroptera:

Ephemerellidae) . Canad. Ent. 103: 512—528.”

Key To The Nymphs Of Ephemerella (Cincticostella)

1. Middle and hind femora expanded and margins serrate; head with sub-

occipital tubercles (Gose 1969: fig. 23; Tshernova 1972: fig. 5)

insolta- group, 2

— Middle and hind femora narrow, and margins entire (Allen 1971: fig.

The Pan-Pacific Entomologist

9) ; head without suboccipital tubercles (Allen 1971: fig. 13)

nigra-group, 4

2 (1). Abdominal terga with paired submedian tubercles on segments 1-10

(Allen 1971: fig. 18) ; fore femora with median hand of tubercles

(Allen 1971: fig. 12) ; head, body and legs with numerous pale spots

(Allen 1971: fig. 14) insolta

— Abdominal terga with paired submedian tubercles on segments 2-10

(Gose 1969: fig. 23) or 4-9 (Tshernova 1972: fig. 5) ; fore femora

without a median band of tubercles; head, body and legs without

pale spots 3

3 (2). Head quadrangular; abdominal terga with paired submedian tubercles

on segments 2-10 (Gose 1969: fig. 23) ; known distributional limits

Thailand boja

— Head round; abdominal terga with paired submedian tubercles on seg-

ments 4-9 (Tshernova 1972: fig. 5) ; known distributional limits Viet-

nam femorata

4 (1). Head with three pale maculae on frons (Gose 1969: fig. 43) ; maxillae

without palpi (Gose 1969: fig. 44) ; known distributional limits Thailand

gosei

— Head without pale maculae on frons; maxillae with palpi; known dis-

tributional limits Manchuria, Korea, Japan, and Russia 5

5 (4). Head with truncate genae (Tshernova 1952: fig. 78) ; middle and hind

femora with an anterior longitudinal ridge levanidovae

— Head with round genae (Allen 1971: fig. 13) ; middle and hind femora

without a longitudinal ridge 6

6 (5). Abdominal terga with paired submedian tubercles on segments 2-9

(Allen 1971: figs. 16-17) ; caudal filaments more than 70% as long

as body (Imanishi 1940: fig. 18) 7

— Abdominal terga with paired submedian tubercles on segments 5-9;

caudal filaments less than 50% as long as body (Imanishi 1940:

fig. 17) tshernovae

7 (6). Abdominal terga with dark sublateral stripes along line of sub-

median tubercles (Imanishi 1940: fig. 18) ; body short, less than 7

mm in length delicata

— Abdominal terga unicolorous, often with a pale median stripe, without

sublateral dark stripes; body moderately long, more than 9 mm in

length 8

8 (7). Abdominal tubercles on segments 2-4 distinct and sharp (Allen 1971:

fig. 17) ; body reddish-brown; known distributional limits Korea

c as tan e a

—Abdominal tubercles on segments 2-4 barely discernible (Allen 1971:

fig. 16) ; body dark brown to black, often with pale median longi-

tudinal stripe; known distributional limits Japan nigra

Ephemerella (Cincticostella) boja, new species

Ephemerella TEA Gose 1969: 132.

This species was described and figured, but not named, from nymphs

collected in Thailand. Ephemerella boja is a distinctive species with a

Vol. 51, No. 1, January 1975

quadrangular head and well developed posterolateral projections on

segments 7-9.

Nymph. — Length: body 11.0-12.0 mm; caudal fliaments 7. 0-8.0 mm. General

color brown. Head quadrangular; maxillary palpi small, two segmented. Thoracic

nota without tubercles or ridges; prothorax much wider than head; anterior

surface of femora smooth, without protuberances; fore femora small, margins

entire; middle and hind femora broadly expanded laterally, margins serrate;

tarsal claws with 3-4 denticles. Abdominal segments 3-9 with well developed

posterolateral projections; terga 2-10 with paired submedian tubercles; tubercles

small on terga 2-3, well developed on 4-10; segments 7-9 with well developed

posterolateral projections. Caudal filaments brown (rewritten from Gose, 1969).

Holotype nymph. — The specimen from which figure 23 in Gose was illustrated

(Nature and Life in Southeast Asia, vol. VI, 1969) from Chanta Buri, Thailand,

20-VI-61, no other data, is designated as the type of the species. The other

nymphal paratopotype is designated as a paratype.

Ephemerella (Cincticostella) castanea Allen

Ephemerella castanea Allen, 1971: 514.

Nymph. — Length: body 10.0-11.0 mm; caudal filaments 7. 0-8.0 mm. General

color reddish-brown with dark brown markings. Head without occipital tubercles;

maxillary palpi with moderately developed palpi. Thoracic nota without tubercles

or ridges. Abdominal terga 2-9 with paired submedian tubercles; tubercles on

segments 2-4 small and sharp, always discernible; tubercles on segments 5-9

moderately developed. Caudal filaments brown with dark brown annulations at

apex of each segment.

Distribution. — South Korea, Kwang Nung and Seoul.

Ephemerella (Cincticostella) delicata Allen

Ephemerella “nay” Imanishi 1940: 206; Tshernova 1952: 274.

Ephemerella delicata Allen 1971 : 517.

Nymph. — Length : body 5. 5-7.0 mm ; caudal filaments 5. 5-7.0 mm. General

color light brown to reddish brown and dark brown. Abdominal terga with paired

submedian tubercles on segments 2-9; tubercles small on segments 2-4 and 9,

well developed on segments 5-8; abdominal terga with dark submedian longitudinal

stripes along line of submedian tubercles, pale medially and laterally, and with

brown lateral maculae on segments 2-8. Caudal filaments with light thick setae

on every second segment (rewritten from Imanishi 1940) .

Distribution. — Kamo River, Kyoto, Yanashiro Prefecture, Japan, and Botanko-

sho, Sekito-ga, Manchuria.

Remarks . — The following discussion is rewritten from Imanishi

(1940) : “Japanese specimens have white line on thorax, but these are

absent on Manchurian collections. Abdominal markings are distinct

on specimens from Manchuria, and indistinct or barely discernible on

the Japanese. This species is almost identical to E. nigra in form and

color, and the only difference is the smaller size and emergence is

The Pan-Pacific Entomologist

slightly later than E. nigra in Japan. The adult is unknown (adult

probably smaller than E. nigra ) and nymph is not as common as E.

nigra”

Ephemerella (Cincticostella) femorata Tshernova

Asiatella femorata Tshernova 1972: 611, 614 (~ Cincticostella) .

Nymph. — Length: body 12.0 mm; caudal filaments 5.0 mm. General color light

brown. Head with paired suboccipital tubercles; genae rounded; maxillary palpi

reduced in size, two segmented. Thoracic nota without tubercles or ridges; pro-

thorax broadly expanded laterally; anterior surface of fore femora without pro-

tuberances, dorsal margin with small protuberances; middle and hind femora

broadly expanded laterally; middle femora with apical serrations; hind femora

with apical and dorsal serrations; femora, tibiae and tarsi margined with setae;

tarsal claws with 2 denticles. Abdominal terga with paired submedian tubercles

on segments 4-9, tubercles increase in length on posterior segments; segments

7-8 with well developed posterolateral projections. Caudal filaments dark brown

at base (rewritten from Tshernova 1972) .

Distribution. — Red River (Song Koi) , Back Thai, North Vietnam.

Ephemerella (Cincticostella) gosei, new species

Ephemerella TEB Gose 1969: 135.

This species, along with E. boja , was described and figured, but not

named, by Gose (1969) from nymphs collected in Thailand.

Nymph. — Length: body 6.0-7. 0 mm; caudal filaments 3. 0-4.0 mm. General

color dark brown. Head round; frons with pale markings mesad to compound

eyes and pale marking over median ocellus; genae truncate; maxillary palpi ab-

sent. Thoracic nota without tubercles or ridges; tarsal claws with 6 denticles.

Abdominal terga 2-9 with paired submedian tubercles; tubercles small on terga

2-3, better developed on 4-9. Caudal filaments light brown (rewritten from Gose

1969) .

Holotype nymph. — The specimen from which figure 38 was illustrated (Nature

and Life in Southeast Asia, vol. VI, Gose 1969) from Chanta Buri, Thailand, 20-

VI-61, no other data, is designated as the type of the species.

Ephemerella (Cincticostella) insolta Allen

Ephemerella insolta Allen 1971: 516.

Nymph. — Length: body 5. 0-6.0 mm; caudal filaments 4.0-5. 0 mm. General

color light brown to brown with numerous small pale spots on head, body and

legs. Head with paired suboccipital tubercles; genae rounded; maxillary palpi

present, but reduced in size. Thoracic nota with ridges; femora expanded laterally

and with spines and protuberances; tarsal claws with 5-8 denticles. Abdominal

terga 1-10 with well developed paired submedian tubercles; segments 7-9 without

posterolateral tubercles. Caudal filaments brown.

Distribution. — The species is known only from the type series collected from

Chiengmai, Thailand.

Vol. 51, No. 1 , January 1975

Ephemerella (Cincticostella) levanidovae Tshernova

Ephemerella levanidovae Tshernova 1952: 274; Tshernova 1958: 76; Edmunds

1959: 545; Allen 1971: 516; Tshernova 1972: 612.

Nymph. — Length: body 12.0 mm; caudal filaments 10.0 mm. General color

brown. Head without occipital tubercles; genae truncate; maxillae with well de-

veloped palpi. Thoracic nota without tubercles or ridges; femora of middle and

hind legs with longitudinal ridge on anterior surface; tarsal claws with 6 denticles.

Abdominal terga 5-8 with well developed tubercles, tubercles small on other

segments. Caudal filaments with spines on every segment (rewritten from

Tshernova 1952) .

Distribution. — This species is known only from the type series collected from

the Hor River in eastern Russia.

Ephemerella (Cincticostella) nigra Ueno

Ephemerella nigra Ueno 1928: 44; Imanishi 1937: 325; Ueno 1950: 128 (male

genitalia) ; Edmunds 1959: 546; Bajkova 1962: 204; Allen 1971: 513; Tshernova

1972: 611.

Chitonophora (?) nigra Ueno 1931: 224.

Nymph. — Length: body 9.5-11.5 mm; caudal filaments 7. 0-8.0 mm. General

color dark brown to black. Head without occipital tubercles; genae round;

maxillary palpi well developed. Thoracic nota without tubercles or ridges. Ab-

dominal terga 2-9 with paired submedian tubercles; tubercles on segments 2-4

barely discernible; tubercles on segments 5-9 moderately developed. Caudal

filaments brown.

Distribution. — This species has been reported from several localities on the

Island of Honshu, Japan.

Ephemerella (Cincticostella) orientalis Tshernova

Ephemerella orientalis Tshernova 1952: 279; Tshernova 1958: 75; Tshernova

1972: 612.

This species was described from a male imago collected in the vicinity of the

Amur River in eastern Russia and the nymphal stage has not been associated at

this time. Tshernova (1972) placed E. orientalis in Cincticostella (as Asiatella)

with the following statement, “The imago described by me as E. orientalis Tshern.

(Tshernova, 1952: 279, table XIV, fig. 99) due to the structure of the genital

appendages is very close to E. nigra Ueno and by all means belongs to the genus

Asiatella .”

Distribution. — The adult of this species was described from Sudhuhynshi

Preserve, USSR and records of additional specimens have not been published.

Ephemerella (Cincticostella) tshernovae Bajkova

Ephemerella “nax” Imanishi 1940: 205; Tshernova 1952: 274.

Ephemerella tshernovae Bajkova 1962: 203; Tshernova 1972: 612, 614 {—imani-

shii ) .

Ephemerella imanishii Allen 1971: 517.

The Pan-Pacific Entomologist

Ephemerella tshernovae nymphs are easily distinguished from those of the other

species in the nigra- group as they possess paired submedian tubercles only on

segments 5-9.

Nymph. — Length: body 10.0-11.0 mm; caudal filaments 4.0 mm. General color

reddish brown to dark brown with purple to purple black markings. Abdominal

terga with large paired submedian tubercles on segments 5-9; abdomen with

posterolateral projections on sgements 4^9; poorly developed on 4-6, well de-

veloped on 7-9; abdominal terga with short heavy setae; abdominal segment 9

with concave lateral margins. Caudal filaments less than one-half body length;

caudal filaments with heavy setae around each segment (rewritten from Imanishi

1940) .

Distribution. — This species is reported by Imanishi (1940) from Gifu Prefecture,

Japan, and four localities in Korea.

Literature Cited

Allen, R. K. 1971. New Asian Ephemerella with notes (Ephemeroptera:

Ephemerellidae) . Canad. Entomol., 103: 512-528, 46 figs.

Bajkova, 0. J. 1962. New species in the genus Ephemerella Walsh (Ephemerop-

tera: Ephemerellidae) from mountain tributaries of the Amur basin.

Izvest. Trkhookeansk. Naurch. Issled. Inst. Rybu. Khoz. i Okeonog.,

48: 202-205, 6 figs.

Edmunds, G. F., Jr. 1959. Subgeneric groups within the mayfly genus Ephe-

merella (Ephemeroptera: Ephemerellidae). Ann Entomol. Soc. Amer.,

52: 543-547.

Gose, K. 1969. Mayflies (Ephemeroptera) from Thailand. Nat. Life SW

Asia, 6: 125-138, 53 figs.

Imanishi, K. 1937. Mayflies from Japanese torrents. VII. Notes on the genus

Ephemerella. Annot. Zook Japon, 16: 321-329.

Imanishi, K. 1938. Mayfiles from Japanese torrents. IX. Life forms and life

zones of mayfly nymphs. Annot. Zool. Japon, 17: 23-26.

Imanishi, K. 1940. Ephemeroptera of Manchouchuo, Inner Mongolia and

Chozen. Rep. Limnobiol. Surv., Kwantung and Manchouchuo, (1940),

pp. 169-263.

Tshernova, O. A. 1952. Mayflies from the River Amur and neighboring waters,

and their part in the nutrition of Amur-fishes. Trudi Amur, ichtiol.

eksped., 1945-1949, 3: 229-360, 141 figs.

Tshernova, O. A. 1958. The geographical distribution of Ephemeroptera and

some peculiar features of the fauna of the Amur Basin. Rev. Entomol.,

37 : 64^84, 5 maps.

Tshernova, O. A. 1972. Some new species of may-flies from Asia (Ephemerop-

tera, Heptageniidae, Ephemerellidae) . Rev. Entomol., 51 : 604-614, 7

figs.

Ueno, M. 1928. Some Japanese mayfly nymphs. Mem. Coll. Sci. Kyoto imp.

Univ., 6: 19-63.

Ueno, M. 1950. Ephemeroptera, pp. 120-130. In: Iconographia Insectorum

Japonicorum, 2nd ed., Tokyo.

Vol. 51, No. 1 , January 1975

Seasonal Abundance and Distribution of Adult Stoneflies

of Sashin Creek, Baranof Island, Southeastern Alaska

(Plecoptera)

Robert J. Ellis

N orthivest Fisheries Center Auke Bay Laboratory

National Marine Fisheries Service, NOAA, Auke Bay, AK 99821

In this paper I record the species of stoneflies in the study area and

describe the relative abundance and seasonal and spatial distributions

of the adults. In addition I record some incidental observations on the

biology of some species. The collections were made intermittently from

1965 to 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).

Sashin Creek originates about 4,000 m from the ocean in a lake 56 ha

in area nested between mountains at 83 m elevation and also receives

waters from several smaller lakes and ponds. Precipitation averages

about 560 cm per year at sea level near the mouth of Sashin Creek.

Snow usually covers the ground from late October to April — usually

to depths exceeding 1 m after November. The annual range in volume

of flow of Sashin Creek is about 0.2 to 20 m 3 /s. This study was con-

centrated in that portion of the stream between its mouth at tidewater

and a 30-m falls about 1,200 m upstream. The gradient of the stream

increases from about 0.1% near tidewater to about 0.7% close to the

falls where the elevation is about 5 m above sea level. The size of the

stream bottom gravel increases similarly — from about 47% cobbles

near tidewater to 81% close to the falls. The stream runs through a

forest of western hemlock ( Tsuga heterophylla [Raf.] Sarg. ) and

Sitka spruce ( Picea sitchensis [Bong.] Carr.) and has relatively

stable banks with overhanging brush, mostly alder (Alrtus) . Also

present are salmonberry (Rubus spectabilis Pursh.), blueberries (Vac-

cinium) , and devilsclub (Oplopanax horridus [Sm.] Miq.). The

stream bottom along the shores frequently contains branches and logs

in various stages of decomposition.

The resident fish of Sashin Creek in approximate order of abundance

are juveniles of coho salmon ( Oncorhynchus kisutch (Walbaum) ) and

rainbow trout (Salmo gairdneri Richardson) and both juveniles and

adults of Dolly Varden ( Salvelinus malma (Walbaum) ) and coastrange

sculpin ( Cottus aleuticus Gilbert). Many pink salmon ( O . gorbuscha

The Pan-Pacific Entomologist 51: 23-30. January 1975

The Pan-Pacific Entomologist

Vol. 51, No. 1 , January 1975

(Walbaum)), and fewer chum salmon (O. keta (Walbaum)) spawn

in the stream, but their progeny go to the ocean soon after they emerge

from the gravel in April, May, or June.

Materials and Methods

I made 204 collections of adult stoneflies intermittently as part of

larger programs. Stoneflies were collected by air-net sweeping and

beating of streamside brush, hand-picking with forceps, and by rearing

nymphs. Specimens were killed and stored in 70% alcohol. A reference

collection of adult stoneflies is maintained at the Auke Bay Fisheries

Laboratory.

In the following list of adult stoneflies from Sashin Creek, each

species occurred throughout the study area below the falls unless other-

wise noted. Most of the collecting was done below the falls and special

mention will be made if I collected the species also above the falls or

in other streams in southeastern Alaska.

Seasonal Distribution and Abundance

I made collections in Sashin Creek that could have yielded adult

stoneflies on 75 days from the first week of March to the first week in

October. Unusually low winter temperatures can cause delay in onset of

emergence of stoneflies in temperate climates (Elliott, 1967; Harper and

Pilon, 1970). Sashin Creek however, has a combination of relatively

high latitude, marine situation, and several tributary lakes or ponds

which greatly reduces within and between season variations in water

flow and temperature. I have therefore assumed that timing and se-

quence of emergence of the species of stoneflies in Sashin Creek is es-

sentially the same from year to year and combined collections from

all years to compare seasonal occurrence and sequence of emergence

of adult stoneflies. The distribution of sampling effort by week is sum-

marized in Figure 2 along with periods of occurrence for each species.

No collections were made between the first weeks of October and March,

but incidental observations have never revealed adult stoneflies at this

time in Sashin Creek.

The several species of stoneflies differed in abundance and length

of period of occurrence and I define seven general categories from

Fig. 1. Map showing location of streams and rivers mentioned in the text.

The Pan-Pacific Entomologist

consideration of the collection data (Table 1 and Fig. 2). The cate-

gories and species involved are:

1. Very abundant for several weeks: Suwallia pallidula

2. Abundant for several weeks: Capnia excavata, Sweltsa exquisita,

Zapada cinctipes

3. Common for several weeks: Sweltsa borealis , Sweltsa oregonensis

4. Very abundant for many days: Capnia nana

5. Common for many days: Capnia melia

6. Abundant for a few days : Zapada frigida

7. Rare for a few days: Doddsia occidentalis, Megarcys signata ,

Despaxia augusta, ParaleuctJ'a occidentalis , Paraleuctra sara,

Podmosta decepta, Zapada oregonensis , Alloperla serrata.

The seasonal progression of emergence and varying lengths of the

periods of occurrence make it likely that four to six species of adult

stoneflies could be collected from Sashin Creek on any day from mid-

March to mid-September. Two of the common seasonal categories of

species are evident in Figure 2 where “spring” ( Capnia , Zapada) and

“summer” ( Alloperla , Sweltsa ) groups appear.

Collection Data for Adult Stoneflies from Sashin Creek

The collection data are summarized in Table 1 and further details

are presented along with the annotations for each species. Jewett

(1959) brought together illustrations of characters for identification

of all the males and most of the females of the Pacific Northwest which

covers all species found in Sashin Creek. Jewett supplies references

for illustrations of the other females that have been described. First

records for Alaska are indicated in Table 1.

Collection Data

Nemouridae. — Four species of Nemouridae were found but only

Zapada cinctipes was abundant. For the entire collection Z. cinctipes

was one of the five most common species in total numbers, number of

collections, and number of days on which it occurred. The other three

species of Nemouridae were among the six least common species (Table

1 ).

Podmosta decepta (Frison). — One of six least common stoneflies

collected at Sashin Creek but found from lowest area to above falls

and in Traitors River, Harris River, and “Middle Creek” (Rowan Bay).

Zapada cinctipes (Banks). — One of the most common stoneflies of

Sashin Creek; occurred from early spring with earliest winter stone-

Vol. 51, No. 1, January 1975

Table 1 . — Number of specimens, collections, and collection days for

each species of adult stonefly collected in Sashin Creek, 1965-72 ( "in-

dicates first record for Alaska).

Number of Specimens

Number

collections

Number

collection

days

Family and Species

Male

F emale

Nemouridae

Podmosta decepta*

Zapada cinctipes

260

233

Zapada frigida

Zapada oregonensis

Leuctridae

Despaxia augusta

Paraleuctra occidentalis

Paraleuctra sara*

Capniidae

Capnia excavata*

445

169

Capnia melia *

Capnia liana *

855

1,221

Taeniopterygidae

Doddsia occidentalis

Perlodidae

Megarcys signata*

Chloroperlidae

Alloperla serrata

Suwallia pallidula

1,007

800

Sweltsa borealis

Sweltsa exqaisita

242

Sweltsa oregonensis

flies of genus Capnia to July. Although Z. cinctipes was found up to

15 m away from stream on snow, it did not seem to be actively migrat-

ing. Z. cinctipes was collected at Auke Creek and the lowest area to

above the falls in Sashin Creek.

Zapada frigida (Claassen). — One of six least common stoneflies col-

lected at Sashin Creek. It was found only above falls and although

The Pan-Pacific Entomologist

found on only 1 day, was then abundant. Type locality for Z. frigida

is across Baranof Island and somewhat north at Sitka, Alaska.

Zapada oregonensis (Claassen). — Although one of six least common

stoneflies collected at Sashin Creek, this species was found both above

and below falls.

Leuctridae. — Adult Leuctridae occurred only rarely — the three

species captured were among the six least commonly captured species

and were never abundant.

Despaxia augusta (Banks). — A late summer-early fall form and the

most abundant of the Leuctridae. The other two species, Paraleuctra

occidentalis and P. sara, are early summer forms. D. augusta was

found both above and below falls.

Paraleuctra occidentalis (Banks). — An early summer form found in

all areas below falls.

Paraleuctra sara (Claassen). — An early summer form, collected both

above and below falls and in Traitors and Harris rivers.

Capniidae. — The three species of Capniidae from Sashin Creek are

winter forms and begin their emergence while the ground is deeply

covered by snow. Two of the three are among the five most abundant

species. Capnia melia and C. nana appeared to actively migrate by

walking over the snow away from the stream and into the forest. I

observed hundreds of individuals, all moving at nearly 90° to the

stream across the snow. A few were found on a steep hillside among

mature spruce and hemlock trees 150 m away from Sashin Creek.

The fate of these migrants is unknown.

Capnia excavata Claassen. — One of most abundant and commonly

present stoneflies of Sashin Creek, although the long period of oc-

currence may be due to a long-lived adult. One female was collected

above falls. C. excavata was also collected at Traitors River.

Capnia melia Frison. — Least abundant of the Capnia in Sashin

Creek and not collected above falls. C. melia was part of cross-snow

migration at Sashin Creek and was also collected on snow near an

unnamed stream on Douglas Island at about 200-m elevation.

Capnia nana Claassen. — Second most abundant of the stoneflies from

Sashin Creek and the major species in migration across snow both

above and below falls. C. nana occurs abundantly with C. excavata ,

C. melia , and Zapada cinctipes on snowbanks in early spring.

Taeniopterygidae. — A single species, Doddsia occidentalis (Banks).

Only one specimen collected, but collection site, weather, and snow

conditions at the time make it likely that this specimen originated in

Sashin Creek.

Vol. 51, No. 1, January 1975

COLLECTION PERIOD

MARCH

APRIL

MAY

JUNE

JULY

AUG.

SEPT.

OCT.

AND SPECIES

12 3 4

COLLECTION PERIODS

ZAPADA CINCTIPES

CAPNIA EXCAVATA

CAPNIA MELIA

CAPNIA NANA

DODDSIA OCCI DENT ALIS

PODMOSTA DECEPTA

PARALEUCTRA OCCIDENTALIS

SWELTSA BOREALIS

—

— —

—

ZAPADA OREGONENSIS

SWELTSA EXQUISITA

PARALEUCTRA SARA

ZAPADA FRIGIDA

SWELTSA OREGONENSIS

SUWALLIA PALLIDULA

MEGARCYS SIGNATA

ALLOPERLA SERRATA

DESPAXIA AUGUSTA

—

Fig. 2. Periods when collections of stoneflies were made in Sashin Creek,

1965-72 (top line), and periods of occurrence of adults of each species.

Perlodidae. — A single species, Megarcys signata (Hagen). Only

one adult was collected from Sashin Creek, but several Megarcys

nymphs were found. All collections of nymphs and the adult are from

above falls.

Chloroperlidae. — Chloroperlid nymphs are a significant part of the

intergravel fauna of salmon spawning beds in southeastern Alaska

(Nicola, 1968; Ellis, 1970) and are also common on logs in Sashin

Creek. Nearly half the adult stoneflies of this study were chloroperlids.

Alloperla serrata Needham and Claassen. — Only one specimen was

collected and that from above the falls. The same collection contained

several Sweltsa borealis, S. exquisita, and Suwallia pallidula.

Suwallia pallidula ( Banks ). — This bright yellow stonefly was the

most abundant species of this study (closely followed by Capnia nana)

and by far the most abundant chloroperlid. S. pallidula collected both

above and below falls, often abundantly — several collections contained

more than 100 specimens. S. pallidula often seen on fruit of both

red and yellow salmonberry along stream.

Sweltsa borealis (Banks) and S. exquisita (Frison)— Collected both

above and below falls in Sashin Creek.

Sweltsa oregonensis (Frison). — Common during summer but never

abundant. S. oregonensis found both above and below falls in Sashin

Creek and at Traitors and Harris rivers and Auke Creek.

The Pan-Pacific Entomologist

Acknowledgments

Stanley G. Jewett, Jr., supplied encouragement and identified or

confirmed the identifications of representatives of all species found in

this study.

Literature Cited

Elliott, J. M. 1967. The life histories and drifting of the Plecoptera and

Ephemeroptera in a Dartmoor stream. J. Anim. Ecol., 36: 343-362.

Ellis, R. J. 1970. Alloperla stonefly nymphs: predators or scavengers on salmon

eggs and alevins? Trans. Am. Fish Soc., 99: 677-683.

Jewett, S. G., Jr. 1959. The stoneflies (Plecoptera) of the Pacific Northwest.

Oreg. State Monogr., Stud. Entomol., 3: 1-95.

Harper, P. P. and Jean-Guy Pilon. 1970. Annual patterns of emergence of

some Quebec stoneflies (Insecta: Plecoptera). Can. J. Zool., 48: 681-

694.

Nicola, S. J. 1968. Scavenging by Alloperla (Plecoptera: Chloroperlidae)

nymphs on dead pink ( Oncorhynchus gorbuschci ) and chum ( 0 . keta )

salmon embryos. Can. J. Zool., 46: 787-796.

SCIENTIFIC NOTE

Trichophaga tapetzella Linnaeus in the Galapagos Islands (Lepidop-

tera: Tineidae ). 1 — Early in 1973 Dr. Pierce Brodkorh (Department of Zoology,

University of Florida) found exuviae of a lepidopterous larva on the skin of a

dead Galapagos flightless cormorant, Nannopterum harrisi (Rothschild), at Punta

Espinosa, Fernandina Island. On March 23, 1973 an adult carpet moth, Trichophaga

tapetzella Linnaeus, emerged. The specimen is deposited in the Florida State

Collection of Arthropods, Division of Plant Industry, Gainesville.

Although cosmopolitan, Trichophaga tapetzella has not previously been recorded

from the Galapagos Islands (Linsley and Usinger, 1966, Proc. Calif. Acad. Sci.

33(7) : 113-196). The moth was reared in 1966 at Barrington Island (DeVries —

Calif. Acad. Sci.) but this was not reported. Host preferences of the carpet moth

when not an economic pest include general detritus and various kinds of animal

matter.

Dr. Donald R. Davis (Department of Entomology, U.S. National Museum of

Natural History, Washington, D.C.) kindly confirmed my determination of the

adult moth from Fernandina Island and also told of captures of moths reared

from owl pellets on the Galapagos Islands as related in correspondence to him,

which could also be Trichophaga tapetzella.

Probably an accidental introduction by man, Trichophaga tapetzella may be

firmly established in the Galapagos Islands by indication of the three reports of

its occurrence there since 1966 noted herein. — J. B. Heppner, Department of

Entomology and Nematology, University of Florida, Gainesville, Florida 32611.

1 Florida Agricultural Experiment Station Journal Series No. 5398.

Vol. 51, No. 1 , January 1975

An Arboreal Cleidogonicl Milliped from Chiapas

(Chordeumida: Cleidogonidae)

Richard L. Hoffman

Radford College, Radford, Virginia 24141

During studies on the fauna of bromeliads in the Mexican state of

Chiapas, Mr. Kenneth E. Lucas (California Academy of Sciences) ob-

tained a number of millipeds which were sent to me for identification.

Most of the species were represented only by immature specimens in the

orders Polydesmida and Spirobolida and not identifiable with cer-

tainty, but of exceptional interest was the occurrence in large numbers

of an undescribed species referable to Cleidogona in the present, rather

broad, concept of that genus.

Up until 1943, only a handful of cleidogonids were known from

Mesoamerica, and even these in a very inadequate way. During the

past three decades, however, a considerable number of species and

several genera have been described from that region, and the “Check-

list of the Millipeds of Mexico and Central America” compiled by H. F.

Loomis (1968) accounted 26 species dispersed through nine genera

(literature surveyed through 1964).

In 1972 appeared a detailed monograph on the Cleidogonidae by

William A. Shear, in which our knowledge of this group was systematized

and vastly augmented in an exemplary fashion. Having access to a

large quantity of new material from Middle America, as well as to types

of many of the established species, Dr. Shear was able to combine

the majority of the existing generic names, and listed a total of 47

species of cleidogonids from south of the Rio Grande: two species in

Solaenogona , three in Dybasia, and 42 in Cleidogona. Although a fair

number of Mexican cleidogonids are known to be troglobitic or troglo-

philic, none have so far, to the best of my knowledge, been recorded

from any kind of arboreal habitat, such as bromeliads.

Not only is the habitat of the present species unusual, but the gonopod

structure is likewise disjunct from the pattern apparently normal for

the genus, so that in publishing a name for the use of Mr. Lucas’s

studies, I take the occasion to append some observations on the morphol-

ogy of the genitalia.

Cleidogona Cook

Cleidogona Cook, 1895, Ann. New York Acad. Sci., vol. 5, p. 3. — Shear, 1972,

Bull. Mus. Comp. Zool., vol. 144, no. 4, pp. 195 et seq. [monographic revision].

The Pan-Pacific Entomologist 51: 31-38. January 1975

The Pan-Pacific Entomologist

Vol. 51, No. 1, January 1975

The reader is referred to Dr. Shear’s revision for a complete account

of the generic synonymy, distribution, phylogeny, and species of this

large and dominantly Nearctic genus.

Cleidogona scandens , new species

(Figs. 1-7)

Male holotype and 46 paratypes of both sexes, from 17 km. SE. of San Cristo-

bal de las Casas, 2195 m., Chiapas, Mexico, collected 15 January 1973, by

Kenneth E. Lucas. Deposited in the California Academy of Sciences.

Diagnosis. A moderately small, well-pigmented species in which the

gonopod telopodltes are strongly reduced and the colpocoxites very

greatly elongated but simple in form (Figs. 1, 6) ; coxae widely sepa-

rated from sternum by prominent peristigmal projections; females

lacking postgenital plate, cyphopodal sternum small and obscure (Fig.

4).

Holotype. Adult male, total length ca. 15 mm., maximum diameter (at segment

7) 1.7 mm. Color pattern to the eye appearing dark brown dorsally with a pale

median stripe and pale legs; with magnification each segment is basically light

horn-brown dorsally with a paramedian dark brown area on each side of mid-

line, each such area containing two small round pale spots, the light brown area

laterad to these with a larger and ovoid pale spot which is strongly areolated on

anterior segments; lower sides and legs whitish-gray. Each segment thus has

a transverse row of six pale spots, four small and two large, and a narrow mid-

dorsal pale area. Lateral sides of femora and tarsi brown; both lateral and dorsal

surfaces of postfemora brown. Antennae uniformly light brown except for some

whitish mottling on articles 1 and 2.

External structure typical of the genus; body widest at segments 6 and

7, tapering very gradually caudad, metazona smooth, very slightly larger than

prozona but without trace of paranotal development. Segments with 3-3 dorsal

setae in a transverse row, the outermost on each end set near caudal margin, the

inner two located near midlength of segment. Head evenly convex above antennae,

the frons slightly but evidently concave. Mandibular bases strongly enlarged

and convex. Ocellaria triangular, each with 27 ocelli in 5 rows.

Anterior legs of male unmodified, except first two pairs smaller as usual.

Gonopods enormously elongated, in situ the apices inserted between the sterna

of the 12th and 13th pairs of legs, and of the form shown in Figures 1, 5, 6, and 7.

<r-

Figs. 1-4. Cleidogona scandens. Fig. 1. Gonopods and legs of the 9th pair,

anteroventral aspect, ventral ends of pleuroterga of segments 6 and 7 shown.

Fig. 2. 9th pair of legs and sternum, anterior view. Fig. 3. Cyphopods and

base of 2nd pair of legs, aboral view, apodematic diverticula (seminal vesicles)

indicated by dotted lines. Fig. 4. Cyphopods, oblique posterolateral view, the

reduced sternum (S) shown by stipple.

The Pan-Pacific Entomologist

Vol. 51, No. 1 , January 1975

Sternum distinct, transverse, strongly sclerotized, not produced medially; lateral

end with a long, laminate sternal apodeme f Sta) extending dorsad nearly to

inside of pleuroterga, and with a prominent triangular lobe containing the stigmal

opening (Fig. 5, 7, Psp) . Coxae small, pivoted upon end of the peristigmatic

process and thus far removed from sternum, a small medially directed basal coxal

apodeme for attachment of the tracheo-coxal muscle; colpocoxites long, slender,

apically expanded, with a triangular subapical lobe on posterior edge, the latter

finely laciniate distad to lobe. Lateral surface of coxae concave, with three macro-

setae. Telopodites small, medially in contact but not fused, less than one-third

length of colpocoxites, each with subapical triangular lobe on the anterior side.

Posterior gonopods (9th pair of legs) of the form shown in Fig. 2, the basal

article with an adenostyle near midlength of ventral surface, second article

narrow at base, broadly expanded distally; three small terminal segments.

Median process of 12th sternum elongate, simple, apically blunt, without

peculiarities.

Female paratype. Similar in general to male except slightly larger, 17.2 mm. in

length, 1.8 mm. in width, body not enlarged at segments 6 and 7. Genitalia as shown

in Figs. 3 and 4; no trace of postgenital sclerite, sternum reduced to a narrow

remnant (Fig. 4, S) hidden above the cyphopods. Latter with densely setose

distal margins, the valves containing an intricate system of convoluted seminal

vesicles.

Distribution and habitat. The species is known so far only from

the type locality. A large series of specimens was taken from Tillandsia

violaceae growing on Quercus sp. at elevations of 3, 6, and 12 meters

above the ground. Specimens taken at the same locality in September

1972, were all immature.

Relationships. So far four species of Cleidogona — C. forficula, C.

laquinta, C. conotyloides, and C. decurva — have been recorded from

Chiapas, all named by Shear in 1972 and all from the vicinity of San

Cristobal. C. scandens shows little close affinity with any of them, and

in fact I have been unable to confidently place it in or near any of

the various species-groups recognized by Dr. Shear. In the key to

Mexican species, pages 206 and 207 of his monograph, it runs out

to couplet 24, but in general appearance the gonopods have little re-

Figs. 5-7. Cleidogona scandens. Fig. 5. Right side of gonopods, oblique

anterolateral view. Fig. 6. Right side of gonopods, lateral view. Fig. 7. Right

side of gonopod bases, anterior view. Fig. 8. Cleidogona major, lateral view of

gonopods. Abbreviations: C, coxa; Cd, condyle; Cxa, coxal apodeme; Clp, colpo-

coxite; S, sternum (the transverse element of the sternal apparatus) ; Sta, sternal

apodeme; T, telopodite; Psp, peristigmatic projection of sternum. The capital

letter S in Figs. 6 and 8 is placed in anatomically the same position for the two

species.

The Pan-Pacific Entomologist

semblance to those of C. stolli and C. mirabilis. There appears to be

actually greater similarity between scandens and C. maculata (Verhoeff)

in the shape of both pairs of gonopods, but the considerable geographic

separation of these two taxa (by over 650 km. and the isthmus of

Tehuantepec) would seem to militate against the likelihood of close

phylogenetic affinity.

I think that a case could be made for the allocation of Cleidogona

scandens to a monospecific group of its own, located perhaps near the

“Crucis Group.” The reduction of the gonopod telopodites, and loss

of the cyphopodal postgenital sclerite, suggest a derivative or specialized

status for the species.

Remarks on Gonopod Morphology

Examination of the gonopods of C. scandens following removal of

extraneous tissue provided the occasion to consider this subject in some

detail, and I venture to set forth here a few comments and observations.

To be sure that we were in agreement on basic concepts, I prepared

by trypsin digestion a specimen of Cleidogona major , the species upon

which Shear’s account (pp. 197, 198) and figures 184 and 185 were

based, to use as a standard. As Shear illustrated the gonopod from

anterior and posterior aspects, I provide here a drawing (Fig. 8) made

from the lateral aspect to complete the picture. I found general con-

cordance with his treatment of anatomy, with the following exceptions

(some of which merely reflect personal preferences in the derivation of

terminology) :

(1) In my material, the lateral ends of the sternum were not bi-

furcated and separate from the lateral extension, but actually formed

a continuous unit somewhat broader on the sides than across the middle.

I think that what Shear has called “lateral sternal sclerites” (LSS) may

be regarded merely as a region of the sternum, set off by a vertical fold

reflecting an internal thickening of chitin, and labeled on my drawing

„ £601 5 ?

as dl

(2) The proximomedian processes from the coxae, referred to as

“coxal knobs” (CK) by Shear, I prefer to call “coxal apodemes” since

they are, as he noted, the point of attachment for tracheocoxal muscles

originating on the opposite side of the gonopod complex. On my figures

they are labeled “Cxa.”

(3) As noted by Dr. Shear, the coxae articulate against the sternum

by a proximal lobe, designated by him “basal coxal bar” (CB), and

shown on his Fig. 184. I could verify this structure, but because of

its form (hardly a “bar”) and function, suggest that it be given a more

Vol. 51, No. 1, January 1975

descriptive name, such as “coxal condyle” as it obviously is a hinge

point for movement of the coxa. As seen in lateral view (Fig. 8, Cd) it

lies just internally to a corresponding thickened region of the sternum

which may represent a pivot against which the condyle moves.

(4) Shear mentions a “. . . membranous front plate (FP) of the

sternum ...” which is said to be attached to the coxae, but no such

plate is shown on his drawings. I found that the space between the

colpocoxites and sternum appears to be somewhat sclerotized in un-

treated material, but after removal of the muscles nothing remains in

that area except very thin connective tissue, the appearance being very

correctly shown in Shear’s Fig. 184. Perhaps there is no true “plate” in

the sense of an anatomically discrete sclerite present in any cleido-

gonids, and even without examination of material I doubt that the

sternum is medially produced in any cleidogonid.

Comparison of the cleared gonopod of C. major with that of C.

scandens showed at once some interesting features, such as the absence,

in the latter, of any trace of lateral extension of the sternum (compare

Figures 7 and 8) , and the considerable prolongation of the peristigmal

region into a long projection which effectively functions as a pivot for

the coxa. The normal internal coxal condyle of C. major appears to be

quite wanting from scandens.

These are differences of considerable magnitude, and doubtless of

more basic importance than the sometimes spectacular modifications

of the colpocoxites and telopodites which have occasioned the proposal

of new genera in the past. But as Dr. Shear has shown convincingly

that anatomical continuities can be found between the most bizarre

gonopodal extremes in this genus, I think that any move to set scandens

off taxonomically would be undesirable, if done prior to a general

re-examination of the coxosternal morphology throughout the genus.

Such an enterprise, as might go without saying, would have to be pred-

icated upon the exclusive use of genitalia cleared of all non-sclerotized

tissue by the use of trypsin or dilute caustic solutions.

At the present I have neither the material nor opportunity to enter

into such a project, and can only emphasize the position of C. scandens

as a cleidogonid of considerable interest both as regards its phylogenetic

position, and from the standpoint of its biotope. How many other

bromeliadicolous cleidogonids remain to be found in Middle America?

Literature Cited

Loomis, H. F. 1968. A Checklist of the Millipeds of Mexico and Central

America. Bull. United States Nat. Mus., 266: 1-137.

The Pan-Pacific Entomologist

Shear, William A. 1972. Studies in the Milliped Order Chordeumida ( Di-

plopoda) : A Revision of the Family Cleidogonidae and a Reclassifica-

tion of the Order Chordeumida in the New World. Bull. Mus. Comp.

Zool., 144(4) : 151-352, figs. 1-538, maps 1-18, diagrams 1-10.

ZOOLOGICAL NOMENCLATURE

Announcement A (n.s.) 94

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. Zool. Nom. 31, part 3, 20th September 1974)

1888. Suppression of Cicada cingulata (Fabricius) var. obscura Hudson, 1891

(Homoptera) .

2048. Suppression of Poly gramma Chevrolat, 1837 (Coleoptera) .

2057. Suppression of Coccus sativus Lancry, 1791, Coccus mexicanus Lamarck,

1801 and Coccus silvestris Lancry, 1791 (Homoptera).

2062. Conservation of Aphis pyri Boyer de Fonscolombe, 1841 and the suppression

of Aphis pyri Kittel, 1827 and Aphis pyri Vallot, 1802 and seven other

binominals proposed by Kittel in 1827 (Homoptera).

2091. Designation of type-species for Dactylopius Costa, 1835 and Pseudococcus

Westwood, 1840; proposed suppression of Diaprosteci Costa, 1828 (Homop-

tera) .

(see Bull. Zool. Nom. 31, part 4, 13th January 1975 [exact publication date]).

1884. Supression of Pamalius Rafinesque, 1815 (Rhopalocera) .

2012. Suppression of Calomicrus taeniatus Wollaston, 1867 (Coleoptera).

2025. Striglina Guenee, 1877 to he given precedence over Daristane Walker, 1859

(Lepidoptera) .

2036. Designation of Hydrophorus binotatus Fallen, 1823 as type-species of

Hydrophorus Fallen, 1823 (Diptera).

2060. Suppression of Xiphidium glaberrimum Burmeister, 1838 and Orchelimum

cuticulare Audinet-Serville, 1838 (Grylloptera) .

2063. Designation of a type-species for Kerrichiella Rosanov, 1965 (Hymenoptera) .

2067. Suppression of Thrips rufa Gmelin, 1790 (Thysanoptera) .

2069. Suppression of Phloeotrogus Motschulsky, 1863 (Coleoptera, SCOLYTI-

DAE).

2070. Suppression of Anodius Motschulsky, 1860 (Coleoptera).

2071. Suppression of Leiparthrum Wollaston, 1854 (Coleoptera).

2072. Suppression of Olonthogaster Motschulsky, 1866 (Coleoptera).

2075. Designation of type-species for Megasternum Mulsant, 1844 and Crypto-

pleurum Mulsant, 1844 (Coleoptera).

2078. Designation of type-species of Platyrhacus Koch, 1847 (Diplopoda).

Comments should be sent in duplicate, citing case number, to 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. —

Margaret Green, Scientific Assistant.

Vol. 51, No. 1, January 1975

Biology of the Tliimbleberry Gallmaker

Diastrophus kincaidii

(Hymenoptera : Cynipidae ) 1

James K. Wangberg

Department of Entomology, University of Idaho, Moscow, Idaho 83843

The cynipid wasp, Diastrophus kincaidii Gillette, is a widely dis-

tributed species in western North America and is known to be the

causative agent of a conspicuous gall of thimbleberry ( Rubus parviflorus

Nutt.). These galls, which are characteristic in form, may be extremely

abundant on the host plant. Weld (1957) has reported that R. par-

viflorus is the only known host of D. kincaidii. According to Weld

(in lit.), distribution extends from Los Gatos, California to British

Columbia. Localities are listed for California, Oregon, Washington

and Idaho. It is possible that D. kincaidii occurs outside of this range

and its distribution may be coextensive with that of thimbleberry which

extends from San Diego County, California north to Alaska and east

into Idaho.

Little information on the biology of D. kincaidii has been reported

in the literature, therefore a study was initiated to examine its life

history and habits, its relationship to gall development and to establish

its insect associates. The study was conducted in a redwood forest

habitat east of Humboldt State University, Areata, California, and sup-

plemented with laboratory rearings from September 1970 to May 1973.

The Gall

Mature galls on thimbleberry are abrupt swellings on the stem or

petiole (Fig. 1). They are irregularly swollen or lumpy and glabrous.

Their size and shape vary greatly depending on the number and precise

location of cynipid larvae present in the gall. Larval cavities may be

numerous and usually are situated closely together. The majority are

located in the vascular zone but many extend into the pith and cortex

as well. The tissue immediately surrounding each larval cavity is hard

and woody as are the outer tissues of the gall but the tissue between

cavities is loose and pithy. Each cavity is irregularly ovoid and about

2X3 mm in size.

Gall development proceeds rapidly following oviposition by the adult

1 A portion of a thesis submitted for an M.A. at Humboldt State University, Areata, California

95521.

The Pan-Pacific Entomologist 51 : 39 - 48 . January 1975

The Pan-Pacific Entomologist

Fig. 1. Mature galls on thimbleberry, .5X.

female in the spring. First evidence is a slight swelling on a stem seven

to ten days following egg deposition. The swelling is due to enlarge-

ment of cortical, xylem and phloem cells. At this stage of development

larval cavities are not yet formed and the eggs and first instar larvae

are wedged between plant cells.

In three to four weeks after oviposition larval cavities have developed

and a gall is present as an abrupt and noticeable enlargement of the

stem. The cavities are formed by a proliferation of parenchyma tissue

that surrounds individual larvae. Xylem and phloem tissues of the gall

are disrupted and only the uninfected portions of the stem have normal

vascular tissue.

The tissues of most galls show signs of woodiness by the end of

summer and during the fall all become woody.

Description of Life Stages

Adult

Descriptions of the adult by Gillette (1893) and Fullaway (1911) will serve

in the recognition of this species. The general characters are as follows: the fe-

male (Fig. 3a) ranges in size from 2.0 to 2.5 mm in length and the male is

approximately 1.75 mm in length. The body is black. The antennae, tegulae and

ovipositor sheath are brown to brownish. The legs are brown and the tips of the

tarsi are black. The laterally compressed abdomen is larger in the female, ap-

proximately half the length of the body. The antennae are 13 segmented in the

female and 14 segmented in the male.

Egg

The egg of D. kincaidii is readily recognized by its oblong shape and long

stalk (Fig. 3b). Measurements of ten eggs averaged 0.65 mm (0.59-0.70) in

length and 0.077 mm (0.052-0.104) in width. The stalk of the egg is approximately

1.3 X longer than the elongate oval body. The stalk is narrow along its length

Vol. 51, No. 1, January 1975

■■■■■■I Period of dominance

Fig. 2. Seasonal occurrence of life stages of Diastrophus kincaidii.

with a swelling at the distal end. The body is white and opaque and the stalk

is clear. The chorion is smooth.

Larva

This species has five larval instars that are similar in appearance (Fig. 3, c-e).

The larvae range in size from 0.3 mm to 2.5 mm in the normal curved position.

They are best distinguished by head capsule measurements (Fig. 4). The body

is 13 segmented, white, translucent, tapering posteriorly, and is apodous. Waste

products are visible and become darker with age and are confined to the center

of the abdomen as an oval mass. The integument is smooth and lacks setae. The

antennae are not visible. Mandibles are present but inconspicuous in the first

instar. They are tridentate in the fifth instar (Fig. 3h) .

Pupa

In D. kincaidii there is long-lasting prepupal stage in addition to a shorter

pupal stage (Fig. 3, f-g) . The prepupa is distinguished by the arcuate body and

opaque yellowish color. The body is fatter than in larval stages so the head

capsule is only about half the size of the prothoracic segment in lateral view. In

older individuals the red pigment of the developing compound eyes is visible in

the prothoracic segment and legs (not yet fully formed) can be seen beneath

each thoracic segment. The mandibles of the larva are forced to the surface as

the pupa develops and come to lie fully exposed on the face.

The free pupa is white to yellowish during early development becoming dark

brown to black with age. The antennae and legs extend caudad and reach the

abdomen. The overall shape is robust, especially in females. Pupae range in

size from 2. 0-2. 5 mm.

The Pan-Pacific Entomologist

Fig. 3. Life stages of D.

instar larva, 150 X, (d) 2nd,

kincaidii. (a) adult, 22 X, (b) egg,

3rd and 4-th instar larva, 90X-25X,

68X, (c) 1st

(e) 5th instar

Vol. 51, No. 1, January 1975

Life History

Diastrophus kincaidii is a univoltine species. The adults are relatively

short lived following emergence in the spring. During the summer D.

kincaidii passes through five larval instars and its galls become notice-

able on thimbleberry. A long diapause begins as a prepupa in the fall

and is broken the following spring when pupation occurs within the

gall. The seasonsal occurrence of the life stages of D. kincaidii is sum-

marized in Figure 2.

Adult

Emergence. Adults emerge from galls in the spring and the emer-

gence of all individuals within a gall is well synchronized. However,

males emerge prior to females. Each adult chews through tissues of the

gall to escape. The wings are fully expanded at this time. Chewing

for the most part follows paths of least resistance and many adults

work their way into the pith. Most adults fortuitously use exit holes

created by others. Once the adult is free of the gall it is fully capable

of moving about, but usually a few minutes are spent grooming or

resting on the gall surface or adjacent stems. A striking characteristic

of the adult is its unwillingness to fly. Neither males nor females were

observed to fly in the field or laboratory. Generally, individuals

wander over the surfaces of the host plant throughout adult life.

Courtship and Mating. Courtship, which occurs on various parts of

the host plant, was observed many times in the laboratory and field.

Upon encountering a female a male immediately shows excitement,

hurriedly walks behind her and mounts from the rear without delay.

In many instances males were observed to fan their wings once or twice

prior to mounting. The female’s response to these initial activities is

always the same and is identical to her reaction to any other disturbance.

Once touched by the male she crouches upon the stem or leaf so her

venter touches or nearly touches the surface. Her head and antennae

are held in contact with the surface and she remains motionless in this

position.

The male caresses her thorax rapidly with his palps, usually touching

the prothorax but sometimes caressing the mesonotum and scutellum.

Simultaneously, his antennae move rapidly in front of her face, although

4 -

larva, 17X, (f) prepupa, 13X, (g) pupa, 19X, (h) left mandible of 5th instar

larva, 175 X-

The Pan-Pacific Entomologist

while she is in the crouched position they do not touch her. After three

to five minutes the male may turn and touch her wings and/or abdomen

with his palps; he then returns to his original position and continues

caressing the thorax. It is common for the male to fan his wings at

brief intervals during courtship.

The female usually remains motionless on the stem throughout the

three to five minutes of courting until the male dismounts and walks

away. The female then rises and continues to walk along the plant

surface. Frequently the same male is attracted again and a second mat-

ing attempt may occur. It is also common for a female to simply drop

from the plant to escape from the male.

After frequent encounters the female will begin to walk with a male

in position for mating. He continues to court in the typical manner but

with the female standing, their antennae touch. The antenna of the male

alternately rubs the corresponding antenna of the female from the distal

to proximal end. This movement is rapid and occurs while he con-

tinues to caress her thorax with his palps.

While the female is in the crouched position, the frequent attempts

by the male to copulate are unsuccessful. Once the female is standing,

however, the male is able to initiate copulation. This is accomplished

when the male moves to one side of the female, grasps her wings and

thorax with his forelegs and lowers himself alongside her abdomen.

Their genitalia unite and mating occurs in this position.

Copulation lasts from thirty to sixty seconds. Both sexes are motion-

less during this period except for slight antennal movements and pulsat-

ing movements of the male’s abdomen. Following copulation the male

remains mounted on the female for a few seconds.

Males were often observed courting the same or different females

following mating; however it is not certain that an individual male or

female actually mates more than once. Males also commonly courted

and attempted to mate with other males.

Oviposition. The time interval between insemination and oviposition is

not known. In a single instance a female was observed to oviposit im-

mediately after mating but the possibility of a prior mating could not

be excluded in this case.

The search for a suitable oviposition site begins after the female has

climbed up the stem to the current season’s growth. She immediately

begins an investigation of the green shoot axis and, occasionally, the

petioles with her antennae. While she walks slowly and erratically over

the stem, her antennae are moved up and down at a moderate rate, com-

Vol. 51, No. 1 , January 1975

ing very near the surface but touching it only occasionally. When a

suitable site is located she stops and both antennae are held downward

with the tips directed toward this site. The antennae quiver in this

position for a few seconds, then the female steps forward, arches her

abdomen and inserts her ovipositor. In so doing, the sternites are bent

downward forming a “V” and the ovipositor is inserted in a smooth,

continuous motion. In many instances females partially insert, pause,

and then may withdraw the ovipositor. This apparent testing of the

site may occur once or twice before the female locates a position on the

stem suitable for oviposition. Once the ovipositor is fully extended and

the venter touches the stem oviposition is accomplished.

During oviposition there are pumping movements of the lower abdo-

men while the female remains motionless, although the antennae and

legs may sometimes quiver slightly. Testing of a site with the ovipositor

lasts only a few seconds but actual oviposition lasts two or three minutes.

Although the passage of eggs was never observed, a brown fluid was

clearly seen passing into the plant tissue at the time of oviposition.

This fluid occupies the entire path of the ovipositor in the plant tissues.

The ovipositor is withdrawn in a smooth motion and the female

moves and reinserts it into the stem very near the previous oviposi-

tional site. When a number of insertions have been made, brown

wounds are visible on the plant surface. These wounds occur in clusters

that range in size from 4^400 mm 2 and may be located on a side of

the stem or may completely encircle it.

The number of eggs deposited by a female is not known, although

it must exceed 30 to 50 as this is the number of larvae in an average

sized gall.

The majority of the eggs are deposited in the xylem or phloem but

a few are usually also located in the cortex or outer edges of the pith.

Egg orientation does not appear to follow a regular pattern.

Oviposition was observed throughout the day. Such activity is greatest

with sunshine and warm temperatures. An individual female may ovi-

posit for an entire day and I believe in some instances into the follow-

ing day. By the second day when oviposition is completed, the female

leaves the oviposition site and moves to the tops of the plant. Females

probably do not live very long following oviposition. In some instances

individuals were found dead with the ovipositor still inserted in the

plant.

A gall can result from the ovipositional activities of one female but

it is not uncommon for two or three individual cynipids to oviposit in

the same gall site. An individual will often oviposit in a second area

The Pan-Pacific Entomologist

HEAD CAPSULE WIDTH, mm

Fig. 4. Larval instars of D. kincaidii.

on the stem, usually above the first, but the galls which develop are

consistently smaller and contain fewer larvae.

Larva

First instar larvae are difficult to discern upon dissection of young

galls. The larval cavities are not fully developed so there is little space

between the larva and adjacent plant cells. The gall tissue is very wet

and I believe that during the first instar ingestion of the surrounding

fluid provides nourishment. The mandibles at this stage are barely

detectable at 50 X magnification and movement of mouthparts cannot

be seen.

By the time the second larval instar is attained, the larval cavities

are well developed in a gall. The amount of parenchyma tissue sur-

rounding the larva is extensive but the larva can move within the cavity

it has made. Movement of the mandibles is now clearly visible. No

changes in larval habits were observed from the second through the

fourth instar.

During the fifth larval instar most galls become woody, to some

extent. The fifth instar has larger, more heavily sclerotized mandibles

which are adapted for chewing the dry tissues that line the larval cavity.

These larvae are active within their cavities and move about to feed,

thus their orientation in the cavity is not consistent. During feeding,

it is common for a larva to chew through the tissue wall separating it

Vol. 51, No. 1, January 1975

from an adjacent larval cavity. Apparently no harm is caused when

two individuals occupy a common cavity because two pupae have been

frequently found within one cavity in mature galls.

Pupa

Prior to pupation the characteristic coloration of mature larvae is an

opaque yellowish due to the accumulation of extensive amounts of fat

body. The development of pupal characters is gradual. Pigmentation

of the eyes is most evident in older individuals and the outline of legs

is only vague. D. kincaidii undergoes diapause during the winter in

this prepupal stage. The prepupa is long lasting and one can expect

to find galls containing prepupae 7 months out of the year.

By late April and May most individuals are in the pupal condition. In

the laboratory the pupal condition lasted from one to two weeks at

room temperature; in the field this period is probably longer. The de-

velopment of the pupae within a gall is well synchronized with the others

and the interval passed in this stage is relatively short.

Insect Associates

Ten species of parasitic hymenopterans were found associated with

D. kincaidii on thimbleberry. Most are parasites of D. kincaidii but

some are hyperparasites. Parasitism of D. kincaidii larvae is extensive

during the summer and oftentimes only those individuals deepest within

the gall tissue escape attack. The parasitic species emerge in succession

throughout the summer and D. kincaidii is subject to parasitism for

its entire larval life.

In addition, an inquilinous weevil and an undetermined cecidomyiid

midge were occasionally present within galls.

The relationships of these species and the structure of the gall com-

munity will be discussed in a subsequent paper. The following is a list-

ing of the insect associates inhabiting galls on thimbleberry.

Parasites — Ichneumonidae :

Torymidae :

Pteromalidae:

Eurytomidae :

Eupelmidae:

Ormyridae :

Eulophidae:

Orthopelma califoriiicum Ash.

Torymus fagopirum 0. S.

Torymus solitarius 0. S.

Habrocytus sp.

Artholytus sp.

Eurytoma n. sp. near auriceps

Eupelmella vesicularis (Retz.)

Ormyrus sp.

Tetrastichus sp.

The Pan-Pacific Entomologist

Inquiline — Curculionidae : Rhynchites bicolor (Fabr.)

Uncertain relationship — Cecidomyiidae: undetermined species

Acknowledgments

I wish to thank Dr. J. R. Barron, Dr. R. deRuette, Dr. C. M.

Yoshimoto (Canada Department of Agriculture), Dr. E. E. Grissell

(Florida Department of Agriculture) and Dr. R. E. Bugbee (Allegheny

College) for the identification of the specimens. Appreciation is also

extended to Dr. R. J. Lyon (Los Angeles City College) for the use

of L. Weld’s notes and collection. I am grateful to Dr. W. F. Barr and

Dr. J. M. Gillespie (University of Idaho) for reviewing this manuscript.

Finally, I wish to thank Dr. R. Hurley (Humboldt State University)

for his assistance during the entire course of the study.

Literature Cited

Fullaway, D. J. 1911. Monograph of the gall-making Cynipidae of California.

Ann. Entomol. Soc. Amer., 4: 331-381.

Gillette, C. P. 1893. Two new cynipids from Washington State. Can. Entomol.,

25: 110-111.

Weld, L. H. 1957. Cynipid galls of the Pacific slope. Ann Arbor, Michigan.

80 pp.

BOOK NOTICE

The Biology of Tribolium with Special Emphasis on Genetic Aspects. A.

SOICOLOFF. Oxford University Clarendon Press, Ely House, London W. 1.

Volume 1. Pp. xix -f- 300, 22 tables and 76 figures in text. £12.00.

This is the first of 3 volumes which will form a study of the flour beetles of the

genus Tribolium. Judging by this part they will also be an invaluable reference

for workers on other genera and families of Coleoptera, and for students in other

fields, since the coverage is monographic. The 8 chapter headings are: Introduc-

tion, Taxonomic position and evolutionary trends, Morphology, Internal anatomy

and histology, Electron microscopy, Developmental and post-embryonic studies,

Teratological abnormalities.

Each chapter is fully documented; the figures are nearly all compound (as

many as 30 drawings) , and the names of anatomical parts are fully spelled out.

Keys include Hinton’s on the genera of Ulomini of America north of Mexico and

the species of adult Tribolium (with Sokoloff’s summary of more recent findings),

and to the pupae and mature larvae of Tribolium from Ho. Chapter 8 on

teratological abnormalities, with illustrations of examples from the genus Tribolium ,

is a particularly useful summary and includes a 4-page tabulation of recorded

abnormalities in the Coleoptera.- — Hugh B. Leech, 1435 Howell Mountain Road

N., Angwin, California 94508.

Vol. 51, No. 1 , January 1975

The North American Species of Triploechus Edwards

(Diptera: Bombyliidae)

Jack C. Hall

Division of Biological Control, University of California,

Riverside, California 92502

Prior to Hull’s 1973 studies of bombyliid genera, Triploechus Edwards

was considered by most American workers to be a subgenus of Hetero-

stylum Macquart. Paramonov (1947) had more or less established the

generic validity of Triploechus , but he was never followed.

Perhaps some of the uncertainties of Triploechus as a distinct genus

in North America lie in the fact that the genus was originally described

from Chile. The two North American species were placed at that time in

either Bombylius, Heterostylum or Triplasius. Painter, 1940 possibly

unaware of Edwards’ 1936 description of Triploechus placed all the

North American species in Heterostylum, removed novum Williston

from Triplasius Loew and synonymized Bombylius recurvus Coquillett

with novum.

Paramonov (1947) was the first to point out reliable differences

between Triploechus and Heterostylum and placed Loew’s Triplasius in

synonymy with Bombylius. Hull (1973) considers Triplasius a sub-

genus of Bombylius.

Triploechus was established by Edwards on the basis of the indented

posterior margin of the eye, the short pulvilli and the presence of three

submarginal cells in the wings. These characteristics are possessed by

all of the South American species but not by all of the North American

forms. Hull’s recharacterization of Triploechus is readily available and

need not be reiterated here, but in the light of present knowledge a few

changes need to be made. Possibly of more significance than the number

of submarginal cells is the course of vein R 2+ 3 , the presence of the upper

intercalary vein or medial crossvein at the base of the second posterior

cell, the width of the head and the configuration of the male genital

armature. A study of all the North and South American species of

Heterostylum and T riploechus has led to the conclusion that T riploechus

is a valid genus.

Most of the North American forms in Triploechus will not run to

that genus in Hull’s key to genera primarily because most have only

two submarginal cells. Vein R 2+ 3 is always curved upward apically

or recurved and meets the costa at a right angle or less in Triploechus.

Also in Triploechus the upper intercalary vein is nearly always present

The Pan-Pacific Entomologist 51 : 49 - 56 . January 1975

The Pan-Pacific Entomologist

even though shorter than the r-m crossvein. It is never or rarely present

in Heterostylum. In conjunction with these differences in venation the

head in Triploechus species is never wider than the thorax while in

Heterostylum it is always as wide as or wider than the width of the

thorax. There are also differences to be found in the configuration of the

male genitalia especially in the shape of the epiphallus.

Differences other than the number of submarginal cells between the

North and South American species of Triploechus should be mentioned

in order to establish a broader base for the genus. The North American

species never have the costa tuberculate while in all the South American

species this is the normal condition although a number of specimens of

bellus (Philippi) have a smooth costa. The pulvilli in the North Ameri-

can species are much longer than those found in the South American

forms except novus. Other than novus, the northern species are

much smaller than their southern cousins. These smaller specimens

actually resemble, in habitus, Bombylius more than Heterostylum.

T. novus and the Chilean species heteroneurus (Macquart) are remark-

ably similar but rather easily separated by the male genitalia.

The new species described herein are all similar in habitus. As stated

above they resemble Bombylius species more than Triploechus species.

In some respects they occupy an intermediate position between Bom-

bylius, Heterostylum and Triploechus sharing characteristics with all

three genera. They are placed in Triploechus on the basis of the indented

posterior margin of the eye, the apically upward curvature of vein R.2+3

and the male genitalia, although it should be mentioned that the geni-

talia of sackeni (Williston) approaches that of Heterostylum.

Key to the Males of Triploechus

1. Eyes separated, usually only narrowly so by less than width of median

ocellus 2

- Eyes contiguous; arista distinctly two segmented; pulvilli short, less than

half the length of the claws novus (Williston)

2. Legs mostly fulvous, femora may be dusky at base or middle 3

- Legs black, tibiae may be testaceous at most sackeni (Williston)

3. Black hairs present on underside of hind femora and usually on mid femora;

first antennal segment fulvous with black hairs above stagei new species

- Legs entirely white pilose; basal antennal segment black with white hairs

above luridus new species

Key to the Females of Triploechus

1. Legs black, tibiae testaceous at most 2

- Legs fulvous, femora dusky toward base at most; first antennal segment

fulvous with yellow hair stagei new species

Vol. 51, No. 1, January 1975

2. Hind femora with at most a few bristles on apical half; pulvilli more than

half as long as claws 3

- Hind femora with strong bristles along the ventral surface; pulvilli less

than half as long as claws novus (Williston)

3. Abdominal segments two to seven with black hairs on posterior margin;

body pile in general whitish sackeni (Williston)

- Body pile predominately yellowish, no black hairs on the abdomen

luridus n. sp.

Triploechus SACKENI (Williston), new combination

Comastes sackeni Williston, 1893: 255.

Heterostylum sackeni , Painter, 1930: 3, Painter and Painter, 1965: 411.

Williston’s type female in the Snow Collection at the University of

Kansas is in fair condition, but one wing, four legs and one antenna are

missing; however the body vestiture is quite intact. In general it is a

little larger than most of the specimens before me and the color of the

wings is quite light.

Since the original description is short and incomplete and of the

female only, the male and female of this species are herein redescrihed

completely.

Male. — Small, 6-7 mm. Ground color dull grayish, in some lights with a greenish

cast, head cinereous, tibiae brownish to dark testaceous; occasionally with three

gray longitudinal stripes on thoracic dorsum. Eyes separated by a little less

than width of median ocellus. First antennal segment five times length of second

segment, with black and white hair above, white hair at sides and below, second

segment bare, one-third wider than long; third segment four times longer than

first, linear, gradually tapering from base to apex, arista terminal, small, shorter

than width of third segment at apex. Head white pilose with a few black hairs

on ocellar tubercle and upper half of face on each side of antennae. Proboscis

projecting beyond oral margin for a distance more than length of hind femur.

Palpi short with short white hair. Thorax and scutellum white pilose with scattered

black hair present on former, tomentum wanting, bristles white. Legs with white

hair and tomentum, bristles testaceous; pulvilli more than half as long as claws.

Halter stem testaceous, knob white. Squama brownish with fringe of long white

hair. Widened base of costa with black setulae, costa without tubercles. Wing

hyaline, basal half or less suffused with light brown, outline of color indistinct;

vein R. 2+3 curved upward meeting the costa at a right angle, petiole at tip of first

posterior cell about equal in length to r-m crossvein, upper intercalary vein

present, shorter than r-m crossvein, two submarginal cells, r-m crossvein beyond

middle of discal cell, axillary cell at base wider than anal cell, alula well

developed. Abdomen white pilose and tomentose, posterior margins of all segments

except the first with long scraggly black hair, tomentum dense on apical segments;

venter white pilose. Genitalia small, (Fig. 1), epiphallus thick, curved upward

apically, rounded below, aedeagus short, straight, not projecting beyond epiphallus;

dististylus not strongly hooked apically, concave on outer margin; hasistylus boat-

shaped, lower margin convex.

The Pan-Pacific Entomologist

Female. — Eyes separated by four times width of ocellar tubercle; front with

black hair laterally; tibiae light testaceous; proboscis projecting beyond oral

margin for distance twice length of hind femur; a few white hairs at base of costa,

white tomentum present on mesonotum, pubescence on abdomen much denser

than in male, seventh sternite broadly rounded apically; otherwise as described for

male.

Holotype female from Argus Mts., San Bernardino Co., California, May 1891.

In Snow Collection, University of Kansas. Painter, 1930 designated a male in the

S. J. Hine collection as the androtype (sic).

Specimens have been studied from: California, San Bernardino Co., 17 $2,

Kramer Hills, IV-18-62 (J. C. Hall); 1 $, 5 mi. W. Desert Springs, IV-11-62

(J. C. Hall, E. I. Schlinger) ; 2 9 , 20 mi. N. Adelanto, IV-18-62 (J. C. Hall,

E. I. Schlinger); 4 $ 9, Granite Pass, IV-27-68 (E. I. Schlinger, C. Beesly),

3800-4320 ft.; $2, Phelan, IV-4-71, V-4-66 (T. Plichta, J. C. Hall); 12 $ 9 ,

8 mi. N. Llano, IV-11-62 (J. C. Hall, E. I. Schlinger) ; 9, Little Morongo Cyn. ;

V-l-62 (J. C. Hall). Riverside Co., 7^9, Riverside, IV-9-37, V-l-69, III-15-39,

IV-2-69, IV-15-73 (P. H. Timberlake, J. C. Hall, M. E. Irwin, T. Plichta) ; 3 $ 9 ,

Palm Springs, III-5-33, III-24-35, III-3-71 (P. H. Timberlake, J. C. Hall) ; 4 $ 9 ,

Chuckwalla Mts., III-29-58 (G. M. Nicolls) ; 9, Mecca, III-31-58 (F. Colley) ; 9,

6 mi. S. Cottonwood Spgs., IV-5-66 (F. G. Andrews) ; 2 9 , 11-5 mi. N.W.

Alberhill, V-ll-69 (J. C. Hall); 9, Coyote Creek, III-22-63 (E. I. Schlinger);

1 $, Millard Cyn., IV-26-69 (B. Hunt); 1 9, 3 mi. E. Edom, III-14-37 (P. H.

Timberlake); 9, 3 mi. N. North Palm Springs, 11-25-68 (M. E. Irwin); P. L.

Boyd Desert Research Center, Deep Canyon, 3.5 mi. S. Palm Desert, numerous dates

in April and May (various collectors) . Imperial Co., 2 9 , 6.2 mi. W. Glamis,

III-2-69 (M. E. Irwin). San Diego Co., $ , 4 mi. E. Jacumba, IV-31-66 (J. Wilcox) ;

$, 5 mi. E. Calexico, 1-27-73 (J. Pinto) ; $ , Vallecito, III-15-68 (J. C. Hall);

9, Fallbrook, V-l-60 (G. Goldman). Los Angeles Co., 3 $ 9 , Soledad Cyn., 3 mi.

N. Acton, IV-27-67 (J. C. Hall). Kern Co., 9, 14.4 mi. N. Johannesburg,

III- 27-71 (T. Plichta). Inyo Co., $ 2, Nine Mile Cyn., IV-6-71 (T. Plichta). Ari-

zona, 9, 4 mi. E. Benson, Cochise Co., IV-10-63 (G. L. Stage) ; $ Aztec, Yuma Co.,

IV- 16-41 (J. Wilcox, Jr.).

Triploechus luridus new species

Quite similar to stagei, new species from which it can he separated

by the black basal antennal segments and the absence of black hair on

the head.

Male. — 9-10 mm. Ground color dull black, head cinereous, mesonotum with three

gray stripes, palpi, base and sometimes apex of femora, tibiae, sides of apical

Lateral view of the male genitalia. All drawn at the same scale.

Fig. 1. T. sackeni (Will.)

Fig. 2. T. luridus n. sp.

Fig. 3. T. novus (Will.)

Fig. 4. T. stagei n. sp.

-A

Vol. 51, No. 1 , January 1975

The Pan-Pacific Entomologist

abdominal segments and nearly all of venter fulvous. Eyes separated by little less

than width of median ocellus. Antennae black, first segment at least three times

longer than second segment which is a little wider than long, third segment linear,

tapering from base to acuminate apex, twice as long as two basal segments

combined. Proboscis projecting one and one-half times length of hind femur

beyond oral margin. Head white pilose as is first antennal segment, front with a

narrow median longitudinal bare stripe from anterior ocellus to base of antennae.

Body pile whitish to pale yellow, lighter on pleura, legs and venter. Apical bristles

on hind femur thin, hair-like. Pulvilli more than half length of claws. Squama

brownish with fringe of pure white hair. Halter stem testaceous, knob white.

Basicosta with whitish-yellow hair and black setulae. Wing hyaline, lightly suf-

fused with brown on basal half, outline of color indistinct; apex of vein R 2+3

curved upward meeting costa at right angle; petiole of first posterior cell as long

as r-m crossvein; upper intercalary vein small; r-m crossvein beyond middle of

discal cell; axillary cell wider than anal cell on basal half; alula well developed,

hyaline. Abdomen entirely pale yellowish to white pilose, tomentum of same color

not dense, scattered on apical segments. Genitalia small, (Fig. 2), epiphallus, in

lateral view parallel sided, apex curved upward rounded and slightly lobed below;

aedeagus straight, projecting about half length of epiphallus; dististylus not

apically hooked, curved outward; basistylus narrow, lower margin broadly curved.

Female. — Eyes separated by four times width of ocellar tubercle; front with

yellow tomentum, body pile and tomentum yellowish; wing nearly entirely hyaline,

brown coloring light, legs entirely fulvous. Otherwise as described for male.

Holotype male and allotype from P. L. Boyd Desert Research Center, 3 mi.

S. Palm Desert, Riverside, California, III-28-73, IV-10-73 (A. Tabet). Both

in California Academy of Sciences.

Paratypes. Four topotypic collected with types. 1 $ , Borrego, Riverside Co.,

California, IV-19-41 (J. Wilcox, Jr.); 2 $, 25 mi. N.W. New Cuyama, Santa

Barbara Co., California, VI-8-63 (G. I. Stage).

Triploechus novus (Williston)

Triplasius novum Williston, 1893: 254.

Heterostylum vierecki Cresson, 1919: 186 new synonymy.

Bombylius recurvus Coquill ett, 1902: 100.

Heterostylum novum, Painter, 1940 : 279.

Triploechus novum, Paramonov, 1947: 191; Painter and Painter, 1965: 411.

(as a subgenus of Heterostylum ) ; Hull, 1973: 141.

Triploechus vierecki. Painter and Painter, 1965: 411 (as a subgenus of Hetero-

stylum) .

T. novus is a widespread species commonly found in the deserts of

Arizona and California. Because of the rather wide distribution a con-

siderable amount of variation is present particularly in the distribution

of the black hair on the head and body. Likewise, most of the specimens

studied have only two submarginal cells, some have two in one wing and

three in the other and some have three submarginal cells in each wing.

In nearly all of the specimens with two submarginal cells there is a stump

Vol. 51, No. 1 , January 1975

of a vien projecting downward towards vein R 4 which if continued

would form the third cell.

In general the females have paler hair than the males. The major

differences noted between vierecki and novus are the white hair and

shape of the antennae and more abundant black hair on both the scu-

tellum and the abdomen. In a series of specimens collected at the same

time and place variation in all three characters is evident. The females

usually have mostly white hair on the antennae while in the males the

black predominates. The shape of the antennae varies according to what

angle viewed from. The scutellum may be entirely black haired, with

only one or two black hairs, or any gradation between. Several condi-

tions are found on the abdomen ; black hair on segment two and at apex

only, on segments two and three or on two, three and four as well as at

the apex. The ground color of the fourth and fifth abdominal segments

is also subject to considerable variation. It may be as in vierecki ,

that is, pale except for median spots of black or only one of the

two segments pale or both entirely black. I have two specimens which

have the sides of all the segments except the first pale. A study of the

male genitalia (Fig. 3) leaves little doubt as to the synonymy listed

above.

This species is known throughout southern California, Arizona, New

Mexico and Texas. Most of the specimens were collected in April.

Triploeclius stagei new species

Readily separated from other species by the nearly entirely fulvous

legs and first antennal segment.

Male. — 9-10 mm. Ground color dull black, head cinereous; first antennal

segment, upper side of face somewhat, legs except coxae and trochanters, sides

of fourth and following abdominal segments and genitalia partially fulvous. Eyes

nearly contiguous above, separated by about width of a single ommatidium.

First antennal segment four times longer than square second segment, third seg-

ment linear, gradually tapering from base to apex, about one and one half times

longer than two basal segments combined. Front with black to brown hair becom-

ing yellowish towards side below; ocellar tubercle and first antennal segment above

with black hair, the latter with pale yellow hair at side and below. Face yellow

pilose, with black hair laterally above, white hair on underside of head and on

occiput. Proboscis projecting beyond oral margin for a distance about equal to

length of hind femur. Body pile pale yellow to light yellowish- gray, lighter on

pleura and venter. Legs white to pale yellow pubescent, mid and hind femora

with scattered black hair on antero-ventral surface, bristles on hind femur on

apical half only; pulvilli nearly as long as claws. Squama brownish with fringe of

long pale yellowish hair. Halter stem flavo-testaceous, knob white. Basicosta with

short whitish hair and black setulae; costa not tuberculate. Wing hyaline, basal

The Pan-Pacific Entomologist

half suffused with brown, color not sharply delineated; vein R 2+ 3 curved upward at

apex to meet costa at nearly a right angle; petiole of first posterior cell as long

as r-m crossvein; upper intercalary vein short; r-m crossvein a little beyond middle

of discal cell; axillary cell at base much wider than anal cell; alula well developed.

Abdominal dorsum pale yellowish to whitish pubescent, long black hair across

posterior margin of all segments except the first ; tomentum consists of short, recum-

bent, somewhat curly hair. Venter white pilose, a few black hairs present at apex.

Genitalia small, (Fig. 4), epiphallus curved upward apically, lower portion of tip

of epiphallus somewhat lobed; aedeagus short extending about half length of

epiphallus; dististylus not strongly hooked apically, posterior margin nearly

straight; basistylus narrow, curved below.

Female. — Eyes separated by two and a half times width of ocellar tubercle. Front

entirely yellow pilose, a few black hairs present on each side at vertex, bare stripe

present down front from ocellar tubercle to antennae. Side of abdomen fulvous at

base, last three or four segments entirely fulvous. Wing coloring lighter than in

male. Body pile and tomentum more yellowish, tomentum more dense. Otherwise

as described for male.

Holotype, allotype and seven paratypes from San Felipe, Baja California,

Mexico, III-27-63 (G. I. Stage). Types in California Academy of Sciences.

Literature Cited

Coquillett, D. W. 1902. New Diptera from North America. Proc. U.S. Nat.

Mus., (1903) 25: 83-126.

Cresson, E. T. 1919. Dipterological Notes and Descriptions. Proc. Acad. Nat.

Sci. Philadelphia, 71: 171-194.

Edwards, F. M. 1936. Bombyliidae from Chile and Western Argentine. Rev.

Chilena Hist. Nat., 40: 31-41.

Hull, F. M. 1973. Beeflies of the World. The Genera of the Family Bombyliidae.

U.S. Nat. Mus. Bull., 286, 687 pp.

Painter, R. H. 1930. A review of the Bombyliid Genus Heterostylum (Diptera).

Jour. Kansas Entomol. Soc., 3(1) : 1-7.

Painter, R. H. 1940. Notes on Type Specimens and Descriptions of New North

American Bombyliidae. Trans. Kansas Acad. Sci., 42: 267-301 (1939).

Painter, R. H. and E. M. Painter. 1965. Family Bombyliidae in Stone et al.:

A Catalog of the Diptera of America North of Mexico. Agric. Res.

Serv., U.S. Dept. Agric. Handbk., 276: 407-446.

Paramonov, S. J. 1947. Zur Kenntnis der Amerikanischen Bombyliiden-Gattung

Triploechus Edw. (Diptera). Rev. de Entomol., 18(1-2); 183-192.

Vol. 51, No. 1 , January 1975

Siricid Woodwasps and Tlieir Associated

Parasitoids in the Southwestern United States

(Hymenoptera : Siricidae)

A. A. Kirk

CSIRO Division of Entomology, P.0. Box 1700,

Canberra City, A.C.T., Australia 1

Since 1962 the Division of Entomology, CSIRO, has been involved

in the biological control of Sirex noctilio F which was accidentally

introduced into and has become established in plantations of Pinus

radiata D. Don in southeastern Australia. Insect parasitoids and para-

sitic nematodes of siricids have been collected in the northern hemi-

sphere and consigned to Tasmania for culturing and subsequent release

in infested areas of Tasmania and Victoria. Earlier collecting in North

America has been confined to California and Nevada and eastern

Canada (Taylor 1967; Lloyd 1968, 1970). Ten species of siricids and

six species of hymenopterous parasitoids were known to be present in

Arizona, Colorado, and New Mexico (Cameron 1965), the three states

I surveyed in 1971.

During April to August 1971 dead or dying coniferous trees and

associated branches, logs and stumps were examined for signs of siricid

larvae or galleries. Infested material was cut into one metre lengths

and transported to outdoor cages at Flagstaff, Arizona. The logs were

separated into groups according to locality and tree species. With

limited time available the main emphasis of the work was placed on

obtaining exact identifications of trees and the insects emerging from

them. Precise daily records were made of insect emergence and from

these their flight periods were determined. Insects were reared from

II localities in northern Arizona, 2 localities in New Mexico, and from

the San Juan Mountains in Colorado.

Results

The siricids Sirex cyaneus F., S. juvencus californicus (Ashmead),

S. longicauda Middlekauff, JJrocerus californicus Norton, U. gigas L.,

Xeris morrisoni morrisoni Cresson and X. spectrum L. were reared

from Abies concolor (Gord. & Glend) Lindl, A. lasiocarpa (Hook)

Nutt, Picea engelmanni Parry and Pinus ponderosa Laws. (Table 1).

1 Present address: CSIRO Biological Control Unit, 335 Ave Abbe Paul Parguel, 34000— Montpellier,

France.

The Pan-Pacific Entomologist 51 : 57 - 61 . January 1975

Table 1. Insect emergence from stored material.

The Pan-Pacific Entomologist

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11 All emerged 1972.

Vol. 51, No. 1, January 1975

Fig. 1 . Seasonal emergence of siricids and associated parasitoids from logs

collected at Happy Jack, Arizona.

Nine species which are known to be parasitoids also emerged. These

were the ichneumonids Rhyssa alaskensis Ashmead, R. hoferi Rohwer,

R. persuasoria L. and Megarhyssa nortoni nortoni Cresson; the ibaliids

lbalia leucospoides ensiger Norton, /. niontana Cresson, /. ruficollis

Cameron and I. rufipes rufipes Cresson; and the stephanid Schlettererius

cinctipes (Cresson) (Table 1). The cleptoparasite Pseudorhyssa

maculicoxis (Kreich) was found associated with R. persuasoria at three

localities in Arizona (Table 1). The timber was stored over the winter

of 1971/72 and another 766 insects emerged during 1972 (Table 1).

More than 50% of the insects which emerged were parasitoids, but this

cannot be taken as a true figure for parasitism because insect emergence

had already taken place from some of the material collected.

Discussion

Rather homeostatic conditions prevail in many of the undisturbed

western United States coniferous forests (Hagen et al., 1971), and

epidemic outbreaks of insects are relatively rare (Balch, 1960). No

evidence was found of current siricid epidemic outbreaks during the

The Pan-Pacific Entomologist

Fig. 2. Seasonal emergence of siricids and associated parasitoids from logs

collected at A. San Juan Mountains, Colorado; B. Sandia Crest, New Mexico.

survey. The coniferous forests of the southwestern United States appear

to be very diverse and stable communities, therefore it seems likely

that host specific associations between insects and trees would have

evolved. Evidence that seems to substantiate this is that S. juvencus

calif ornicus was associated only with Pinus spp. and S. longicauda ,

U. calif ornicus and X. morrisoni were associated only with Abies spp.

(Table 1). Less specialized relationships were found with S. cyaneus,

which emerged from both Abies and Picea, and X. spectrum , which

emerged from these as well as Pinus (Table 1). In addition Sprad-

bery and Kirk (unpublished data), after eight years of extensive

survey work and intensive collecting of siricid infested material and

subsequent meticulous rearing, have evidence that there are distinct

siricid coniferous tree associations in Europe and neighbouring areas.

While the apparent siricid host tree associations in the southwestern

United States described above cannot be regarded as conclusive without

further data, there are good reasons for thinking that they are firm

associations.

R. hoferi and I. j-uficollis only emerged from Pinus (Table 1) and

may be associated with S. juvencus californicus. R. persuasoria, M.

nortoni nortoni and S. cinctipes attacked a wider range of siricid

species from different host tree species (Table 1) .

Relationships between siricids and parasitoids are clearly seen when

the flight periods of the three ibaliid species are examined. I. rufipes

Vol. 51, No. 1, January 1975

rufipes and I. montana have early summer flight periods and could

only attack overwintered siricid larvae (Figs. 1, 2). The closely re-

lated I. rufipes drewseni Borries in Europe (Kerrich, 1973) behaves in

the same way (Spradbery, 1970). I. ruficollis has an early autumn

flight period and was observed to attack S. juvencus calif ornicus larvae

late in September. The closely related I. leucospoides leucospoides

(Hochenwarth) , (Kerrich, 1973), from Europe and neighbouring areas

also attacks siricids in autumn, active females being noted in Tunisia

as late as November (Spradbery and Kirk, unpublished data).

Much further work needs to be done to clarify the role of parasitoids

in the suppression of siricid populations in the southwestern United

States, but present evidence indicates considerable importance.

Acknowledgements

I wish to thank the Director of the U.S. Forestry Service for permis-

sion to collect infested timber, Forestry Service personnel for technical

assistance, Dr. J. R. Wick, Chairman, Department of Biological

Sciences, Northern Arizona University, Flagstaff, for providing labora-

tory facilities, Dr. C. D. Johnson for help with the manuscript, Mrs.

Frances B. McAlister for permission to site cages on her land and Mr.

T. D. Center for invaluable assistance during the survey. Funds for

the project were provided by the National Sirex Fund, Australia.

References

Cameron, E. A. 1965. The Siricidae ( Hymenoptera: Siricidae) and their para-

sites. C.I.B.C. Tech. Bull., 5, 31 pp.

Balch, R. E. 1960. The approach to biological control in forest entomology.

Can. Entomol., 92: 291-310.

Hagen, K. S., R. van den Bosch, D. L. Dahlsten. 1971. The importance of

naturally-occurring biological control in the western United States.

Chapt. 11 in Biological Control, C. B. Huffaker (ed.). Plenum Publ.

Corp., N.Y. 511 pp.

Kerrich, G. J. 1973. On the taxonomy of some forms of Ibalia Latreille

(Hymenoptera: Cynipoidea) associated with conifers. Zool. J. Linn.

Soc., 53: 65-79.

Lloyd, D. C. 1968. Report on the collection and shipment of parasites of

Siricidae in eastern Canada in 1967/68. C.I.B.C. Report. 7 pp.

Lloyd, D. C. 1970. Collection and shipment of parasites of Siricidae in the

western United States in 1970. C.I.B.C. Report. 6 pp.

Spradbery, J. P. 1970. The biology of Ibalia drewseni Borries (Hymenoptera:

Ibaliidae) , a parasite of siricid woodwasps. Proc. R. Entomol. Soc.

Lond. (A), 45: 104-113.

Taylor, K. L. 1967. The introduction, culture, liberation and recovery of

parasites of Sirex noctilio in Tasmania, 1962-67. CSIRO Div. Entomol.,

Tech. Paper, 8.

The Pan-Pacific Entomologist

A Reconsideration of the Nearctic Rhexoza 1

(Diptera: Scatopsidae)

Edwin F. Cook

Department of Entomology, Fisheries, and Wildlife

University of Minnesota, St. Paul, Minnesota 55108

The examination of two collections of specimens sent to me for

identification by Dr. H. J. Teskey of the Entomology Research Institute

of the Canadian Department of Agriculture revealed two new species

of the scatopsid genus Rhexoza. This has prompted me to reexamine

the North American species of this group and has also given me the

opportunity to make a decision on the status of the Quatei group of

that genus which I described in 1956. I noted in both 1971 and 1972

that this group is certainly distinct from Rhexoza , sensu stricto.

Swammerdamella , Rhexoza and the species constituting the Quatei

group are closely related, or at least they share characters not present

in other genera. There are also several shared characters with Coboldia,

so these 4 genera should all be included in the Swammerdamellini. This

is a new placement for Coboldia since I previously included it in the

Scatopsini. However, the large, reniform maxillary palpi; the long,

triangular, anterior spiracular plate; the relatively short R complex,

ending only slightly beyond the middle of the wing; and the presence

of lower epimeral setae all indicate a much closer affinity with Rhexoza

than with Scatopse or Reichertella.

The tribe Swammerdamellini, then, consists of Swammerdamella,

Rhexoza, Coboldia and the new genus described here.

The genera in this tribe can be readily discriminated by use of the

following key.

1. Length of costa between Ri and Ra less than or no longer than length

of Ri; fork of M no longer than or shorter than stem. Males with ab-

dominal segment 7 concealed in segment 6 Swammerdamella

Length of costa between Ri and Ra greater than length of Ri; M fork

longer than stem; abdominal segment 7 visible in males 2

2. Male with tergum 7 produced posteriorly as a narrow, spatulate process,

genitalia with elongate, coiled penis; female with small valvifers on

sternum 8, with rounded, bare emargination on posterior margin of

sternum 7 Coboldia

1 Paper No. 8846 Scientific Journal Series, Minnesota Agricultural Experiment Station, St. Paul,

Minnesota.

The Pan-Pacific Entomologist 51: 62-75. January 1975

Vol. 51, No. 1, January 1975

Male with tergum 7 produced posteriorly as a broad triangle, penis small,

never coiled; female without posterior emargination on sternum 7 3

3. Male genitalia laterally compressed, tergum 9 produced ventrally in a

beak-like process (fig. 6) ; female with tergum 8 medially divided

(fig. 4) Quateiella

Male genitalia not laterally compressed, tergum 9 not produced ventrally;

female with tergum 8 entire Rhexoza

QUATEIELLA, New Genus

The existence of a super specific taxon for the species included here

was first recognized by Cook in 1956. The taxon was then treated

simply as the quatei group of the genus Rhexoza. I now believe that

the group is sufficiently distinct to raise to full generic status. It shares

the four characters noted above with the other genera in the Swammer-

damellini but the male genitalia are very distinctive. They, in fact, are

more like those of some Reichertella in the Scatopsini.

Description. — Antennae 10-segmented (8 flagellomeres) , about as long as total

head height; cardo-stipites rather broad, bandlike, sclerites fused posteriorly

and beset with setae on each side; maxillary palpi large, reniform, 0.10-0.13 mm

long; occiput setose and microtrichiose ; eyes with setae between facets. Thorax

with elongate, triangular spiracular sclerite (fig. 1) ; supraalar setae 8-18, in a

single row (fig. 1); preepisternals 7-14; anepisternals 18-28; upper episternals

5-12; subalars 6-12; subspiraculars 3-7; lower epimerals 3-8; pedicellars 1-4;

wing venation like that of Rhexoza with costal-radial complex ending near middle

of wing; M fork long, Ml and M2 diverging to wing margin; setae on costa,

radial complex and posterior wing margin; setae rarely on ventral surface of R3,

always present on dorsal surface; membrane with dense microtrichia. Abdomen

with 7 segments; sterna 1-4 unsclerotized ; abdomen setose and microtrichiose;

tergum 7 of male posteriorly produced, sternum 7 with modified posterior margin.

Genital vesica and apodemes of male small; male genitalia rotated 180°, with

only 1 pair of appendages, penis inconspicuous, tergum 9 without processes but

produced more or less “beak-like” ventrally. Female genitalia with tergum 8

medially divided, cerci small, no obvious appendages or valvifers, spermatheca

spherical to broadly elliptical. Type-species of the genus: Rhexoza quatei Cook.

Key to Adults of Quateiella

Tergum 9 of male with dorsally projecting horns; female with sternum

9 bearing 10 or more setae in elongate cluster miniscula

Tergum 9 of male without dorsal horns; female with sternum 9 bearing

5-7 setae near posterior margin only quatei

Quateiella quatei (Cook), New Combination

Rhexoza quatei Cook, 1956: 8-10.

Male. — Total length 1.20-1.55 mm; greyish brown to dark blackish brown,

dull to feebly shining; halteres dark grey; wings clear, shining. Antennae with

The Pan-Pacific Entomologist

each flagellomere bearing 10-13 setae; cardo-stipites fused posteriorly, each

side with 12-15 setae in 2 irregular rows; maxillary palpi large, reniform, 0.11-

0.13 mm long; 2 campaniform sensilla on each side of occiput immediately behind

eyes and below level of antennae, 1 sensilla laterad of each lateral ocellus.

Supraalar setae 8-13; preepisternals 8-12; anepisternals 23-28; upper episternals

5-8; subalars 6-12; subspiraculars 3-5; lower epimerals 4-7; pedicellars 2-3.

Wing length 1.00-1.20 mm; R3 extends 0.49-0.53 of total wing length; Ml and

M2 divergent to wing margin; setae on costa and R, none on M, only on dorsal

side of R3; wing surface with obvious microtrichia. Abdomen with sterna 1-4

unsclerotized, terga all present ; terga and sterna setose and microtrichiose ; 7

pairs of spiracles; tergum 7 (fig. 2) with 2 stout, setose processes arising from

under posterior margin (possibly tergum 8) ; sternum 7 with deep, triangular

emargination posteriorly; genital vesica and apodemes small, 0.12-0.13 mm long.

Genitalia in fig. 5.

Female. — Like male in color, structure and chaetotaxy except rather more

setose. Each flagellomere with about 8 setae; cardo-stipites with 16 setae on each

side. Supraalar setae 8-18; preepisternals 13-14; anepisternals 25-37; upper

episternals 5-12 ; subalars 8-12 ; subspiraculars 3-7 ; lower epimerals 6-8 ; pedi-

cellars 1-2. Wing length 1.30-1.38 mm.

Distribution. — Originally described from California: Davis, Riverside, Coachella,

Rialto, Anaheim, Olive. A long series of additional specimens has become avail-

able from San Bernardino, California, June 6, 1960, reared from Echinocactus

polycephalus. Earlier records included specimens from Citronella bait traps.

The females reared from E. polycephalus seem to he one species only. These

were associated with large numbers of males of quatei and a few males of R.

ryckmani. Also associated were even larger numbers of Coboldia fuscipes (Mg.).

Quateiella miniscula (Cook), New Combination

Rhexoza miniscula Cook, 1956: 10.

Male. — Total length 1.30-1.74 mm; dark grey brown, feebly shining; halteres

dark grey. Antennal flagellomeres each with about 9 setae; cardo-stipites with 11

setae on each side; maxillary palpi 0.10 mm long, reniform; campaniform sensilla

as in Q. quatei. Supraalar setae 9-11 ; preepisternals 7 ; anepisternals 18 ; uppei

episternals 7 ; subalars 8 ; subspiraculars 5-7 ; lower epimerals 3 ; pedicellars 1-4.

Wing length 1.00-1.24 mm; R3 .50 total wing length; setae on costa and R, none

on M; R3 with only dorsal setae; wing with obvious microtrichia. Abdomen

with sterna 1-4 undifferentiated; terga all present; terga and sterna setose and

microtrichiose; tergum 7 (and possibly 8) as in Q. quatei except posterior pro-

cesses stouter (fig. 3). Genital vesica and apodemes 0.18 mm long; genitalia as

in fig. 6.

Female. — Total length 1.55-1.97 mm; wing length 1.10-1.38 mm; colored as

in male. Like the males except some differences in pleural setal counts. Flagel-

lomeres each with 10 setae; cardo-stipites with 14 setae on each side. Supraalar

setae 9-10; preepisternals 7; anepisternals 26; upper episternals 7; subalars 7;

subspiraculars 7 ; lower epimerals 1-2 ; pedicellars 1-2. Genitalia as in fig. 4.

Distribution. — Originally described from Virginia, South Carolina and Texas.

Additional specimens have been seen from Texas and from Nogales, Arizona.

Vol. 51, No. 1 , January 1975

Rhexoza Enderlein

Rhexoza Enderlein, 1936: 55; Cook, 1956: 1-12; 1972: 58-61; 1972: 632-634.

Type-species: Rhexoza zacheri Enderlein — Scatopse subnitens Verrall.

Diagnosis. — A ntennae with 8 flagellomeres ; maxillary palpi large (0.09-0.18

mm), reniform, sometimes apically acute (“slipper-shaped”); cardo-stipites a

band-like sclerite fused posteroinedially, bearing several rows of setae; with

campaniform sensilla as in Quateiella; occiput setose and microtrichiose ; eyes

with setae between facets. Supraalar setae numerous, a dense row of 9 or more;

the usual pleural setae present and some with lower epimeral setae, episternal

setae very evident. Wing venation like that of Coboldia fuscipes (Mg.) with

costal-radial complex ending near middle of wing (0.45-0.57 of total length from

base) ; M fork complete; setae present on costa, R and posterior wing margin

only; membrane with short, abundant microtrichia. Abdomen with 7 evident

segments in both sexes; male with 7 pairs of spiracles, females with 8; all terga

evident, anterior sterna reduced or absent; terga and sterna with microtrichia in

addition to setae; segment 7 of male posteriorly modified either on tergum,

sternum or both; genital vesica and apodemes small. Male genitalia rotated

180°, with only 1 pair of appendages; tergum 9 without processes; penis stout,

conspicuously sclerotized. Female genitalia with tergum 8 large or small, never

completely divided longitudinally, cerci large or small, setose; spermatheca

spherical to elliptical.

Key to the Species of Rhexoza

Males; (males of R. borealis and R. amaryllis unknown)

1. With lower epimeral setae (fig. 1) ; R3 with only dorsal setae 4

Without lower epimeral setae; R3 with dorsal and ventral setae 2

2. Tergum 7 with median notch on posterior margin (fig. 7) grossa

Tergum 7 with posterior margin entire 3

3. Genitalia with tergum 9 short, broadly and shallowly incised posteriorly

(fig. 13) ; genital vesica and apodemes short (0.20-0.24 mm) ; subalar

setae 5-6 incisa

Genitalia with tergum 9 produced posteriorly, apically rounded (fig. 14),

genital vesica and apodemes long (0.35 mm) ; subalar setae 14 iowensis

4. Sternum 7 without posteriorly directed processes; aedeagal plate broadly

spatulate (fig. 18) melanderi

Sternum 7 with posteriorly directed processes (fig. 8) ; aedeagal plate

absent or narrow, spatulate 5

5. Sternum 7 with a pair of bilobed processes on posterior margin (fig. 8)

similis

Sternum 7 with 2 simple processes on posterior margin 6

6. Sternum 7 with 2 short, divergent, apically truncate processes on posterior

margin (fig. 9) ; only 1-2 subspiracular setae aterrima

Sternum 7 with 2 long, slender processes on posterior margin; subspirac-

ular setae 3 or more 7

7. Penis short; long apically bifurcate aedeagal plate also present (fig. 19)

teskeyi

The Pan-Pacific Entomologist

Penis sclerotized, longer than penis valves; aedeagal plate absent (figs.

20, 21) Tyckmani

Females: (female of R. iowensis unknown)

1. With lower epimeral setae; R3 with only dorsal setae 4

Without lower epimeral setae; R3 with dorsal and ventral setae 2

2. Tergum 8 medially separated; with a row of stout setae on anterior margin

(fig. 22) grossa

Tergum 8 deeply emarginate but without anterior setal row 3

3. Tergum and sternum 7 with numerous, small, lobelike processes on pos-

terior margin (fig. 25) ; sternum 8 with apically narrowed valvifers (fig.

24) incisa

Tergum and sternum 7 without posterior processes; sternum 8 with valvifers

broadly rounded posteriorly (fig. 27) borealis

4. With 10 or more lower epimeral setae; valvifers short, rounded somewhat

transverse (fig. 28) ; spermatheca reniform amaryllis

Usually with 6 or fewer lower epimerals; valvifers elongate, not transverse __ 5

5. Subspiracular setae 2-3 ; spermatheca spherical aterrima

Subspiracular setae 6-8 (rarely 4) ; spermatheca elliptical 6

6. Tergum 8 without spiracles; with median, longitudinal suture on pos-

terior half (fig. 31) ryckmani

Tergum 8 with spiracles, no median, longitudinal suture 7

7. Anterior half of sternum 7 densely microtrichiose, devoid of setae teskeyi

Anterior half of sternum 7 with only a narrow band without setae 8

8. Sternum 8 not produced anteriorly; distinct space between valvifers (fig.

30) melanderi

Sternum 8 produced anteriorly as median lobe; valvifers contiguous mesally

(fig. 23) similis

Rhexoza incisa Cook

Rhexoza incisa Cook, 1956: 6-7; 1963: 12.

The original description was fairly extensive but additional characters of value

will be noted in the following brief description.

Male. — Very dark blackish brown, dull to feebly shining; halteres grey; total

length 1.38-1.86 mm. Cardo-stipites with 10-14 setae on each side; antennal

flagellomeres with 12-14 setae in a single irregular whorl; maxillary palpi 0.09-

0.12 mm long. Supraalar setae 16-20; preepisternals 6-7; anepisternals 20-25;

upper episternals 5-6; subalars 5-6; subspiraculars 4—9 ; lower epimerals absent;

pedicellars 1-2. Wing length 1.27-1.60 mm; vein R3 terminates at .53-56 of

Figs. 1-9. Fig. 1. Quateiella quatei, thorax; Fig. 2. Q. quatei , segment 7,

male; Fig. 3. Q. miniscula process of segment 7, male; Fig. 4. Q. miniscula, female

genitalia; Fig. 5. Q. quatei, male genitalia; Fig. 6. Q. miniscula, male genitalia.

Fig. 7. Rhexoza grossa, male segment 7; Fig. 8. R. similis, male segment 7; Fig.

9. R. aterrima, male segment 7.

Vol. 51, No. 1 , January 1975

The Pan-Pacific Entomologist

total wing length; R3 with setae dorsally and ventrally. Abdominal segment 7

with tergum slightly produced posteriorly and sternum posteromedially notched;

genital vesica and apodemes 0.20-0.27 mm long; genitalia in fig. 13.

Female. — As in male in color and otherwise except setae of flagellomere up

to 15; preepisternal setae 4-5; anepisternal setae 17-18; subalar setae 4-9 and

pedicellar setae 2-3; spermatheca nearly spherical; sternum and tergum 7 with

seta bearing lobes on posterior margins (fig. 25) ; genitalia in fig. 24.

Distribution. — Minnesota, Iowa, Illinois and Ontario. Larvae have been found

in wet areas under the bark of decaying cottonwood, elm and boxelder.

Rhexoza grossa Cook

Rhexoza grossa Cook, 1956: 6

A larger species than R. incisa, with a western distribution.

Male. — Dark blackish brown, somewhat shining, halteres grey; total length

2.38-2.60 mm. Cardo-stipites with 24 setae on each side (more than incisa ) ;

each flagellomere with about 18 setae in a single whorl; maxillary palpi 0.15-0.16

mm long. Supraalar setae 12-18; preepisternals 16; anepisternals 30; upper

episternals 6; subalars 14; subspiraculars 10; lower epimeral 0; pedicellars

2-5; wing length 1.90-1.96 mm; R3 with setae both dorsally and ventrally.

Tergum and sternum 7 both with posterior emargination (fig. 7) ; genital vesica

and apodemes 0.29-0.30 mm long. Genitalia in fig. 15.

Female. — As in the male in color and otherwise except cardo-stipital setae

17-22; maxillary palpal length 0.14^0.18 mm; antennal flagellomeres with 14-15

setae. Supraalar setae 11-19; preepisternals 13-17; anepisternals 24-25; upper

episternals 5-7; subalars 6-11; subspiraculars 5-12; lower epimerals 0; R3 ex-

tending 0.52-0.57 of total wing length. Genitalia in fig. 22; spermatheca nearly

spherical.

Distribution. — California, Oregon, British Columbia (Richter Pass Road, 7 mi.

W. Osyoos, B.C., 2- VI-1958) .

Rhexoza iowensis, New Species

Male. — About 1.75 mm long; dark blackish brown, head and thorax somewhat

shining, abdomen dull; halteres grey. Cardo-stipites with 18 setae on each side;

maxillary palpi 0.11 mm long; flagellomeres with about 12 setae arranged in a

single whorl. Supraalar setae 18; preepisternals 8; anepisternals 33; upper

episternals 8; subalars 14; subspiraculars 10; lower epimerals 0; pedicellars 3.

Wing length 1.75 mm; R3 extending 0.57 of total length; R3 with dorsal and

ventral setae. Abdomen with sterna 1-4 undifferentiated, other sterna and terga

present, both setose and microtrichiose ; tergum 7 somewhat produced posteriorly,

Figs. 10-17. Male terminalia of Rhexoza. Fig. 10. R. oivensis, segment 7;

Fig. 11. R. ryckmani, segment 7; Fig. 12. R. teskeyi, segment 7; Fig. 13. R. incisa,

genitalia; Fig. 14. R. iowensis, genitalia; Fig. 15. R. grossa, genitalia; Fig. 16.

R. aterrima genitalia; Fig. 17. R. similis, genitalia.

Vol. 51, No. 1 , January 1975

The Pan-Pacific Entomologist

a broadly truncated triangle with small, shallow posterior emargination medially

(fig. 10) ; sternum 7 with deep posterior emargination; tergum 7 setose but devoid

of microtrichia, sternum with setae and sparse microtrichia. Genital vesica 0.35

mm long. Genitalia in fig 14.

Female. — Unknown.

Holotype male: Ames, Iowa, 18-V-1951, W. L. Downes. Type in the Uni-

versity of Minnesota Collections.

Rhexoza borealis, New Species

Male. — Unknown.

Female. — Very dark grey brown, dull; halteres dark grey, head black; total

length 2.25 mm. Antennae short, flagellomeres wider than long, 8 present, each

flagellomere with about 12 setae in a single, irregular whorl; 2 campaniform

sensilla on occiput on each side below level of antennae, 1 laterad of each lateral

ocellus; cardo-stipites with 12 setae on each side in an irregular double row;

maxillary palpi large, reniform, 0.11 mm long. Supraalar setae 18; preepisternals

12; anepisternals 26; upper episternals 5; subalars 11; subspiraculars 9; lower

epimerals 0; pedicellars 2; wing length 2.00 mm; setae on costa and R but not

on M; setae on both dorsal and ventral surfaces of R3; R3 extends 0.58 of total

wing length. Abdominal sterna 1-3 absent, other terga and sterna present, both

microtrichiose and setose ; tergum 7 with straight posterior margin ; sternum 7

broadly and shallowly emarginate ; genitalia in fig. 27 ; spermatheca spherical,

0.13 mm in diameter.

Holotype female: Umiat, Alaska, 8-VII-1958, J. E. H. Martin. Type in the

Canadian National Collections.

Rhexoza aterrima (Melander)

Rhegmoclema aterrima Melander. 1916: 14 Cook, 1956: 4-5 {Rhexoza).

This species is fairly well described in the references cited but the following

additional characters should be noted.

Males. — Dark blackish brown, feebly shining, halteres dark grey; total length

1.50-1.65 mm. Cardo-stipites with 10-15 setae on each side; maxillary palpi

0.09-0.12 mm long; flagellomeres with 12-13 setae in a single whorl. Supraalar

setae 10-13 ; preepisternals 5-7 ; anepisternals 12-23 ; upper episternals 6-7 ; sub-

alars 5-7; subspiraculars 1-4; lower epimerals 3; pedicellars 2; wing length

1.0-1.50 mm; R3 terminating at 0.48-0.50 of wing length; R3 with dorsal setae

only. Segment 7 of abdomen with tergum somewhat produced medially; sternum

7 with a pair of setose processes on posterior margin (fig. 9) ; genital vesica and

apodemes 0.17-0.20 mm long. Genitalia in fig. 16.

Female. — Color and structure as in male except 1.35-2.00 mm long; wing

length 1.20-1.50 mm. Genitalia in fig. 26.

Distribution. — Early collections were from Idaho, Montana and California.

I now have 1 male, Hempstead Co., Arkansas 18-X-1957, reared from Bk.Btl. logs

[sic] ; 1 female, Constance Bay, Ontario, 3-1-1952, from log of Jack pine (I

originally misidentified this specimen as R. similis) . Some Idaho specimens have

been taken from Pimis ponderosa.

Vol. 51, No. 1 , January 1975

Rhexoza SIMILIS (Beekey)

Scatopse similis Beekey, 1938: 151-154; Cook, 1956: 6; 1963: 11.

Well described in the original description and by Cook, but some additional

characters can be noted.

Male. — Total length 1.80-2.00 mm; dark blackish brown, feebly shining, hal-

teres dark. Cardo-stipites with 12-14 setae on each side; maxillary palpi 0.10-

0.12 mm long; flagellomeres with 14-18 setae each in an irregular whorl. Supra-

alar setae 16-21; preepisternals 10-15; anepisternals 22-23; upper episternals

7-10; subalars 6-11; subspiraculars 3-7; lower epimerals 4-8; pedicellars 2-4;

wing length 1.50-2.00 mm; R3 extending 0.51-0.54 of total wing length; R3 with

dorsal setae only. Tergum 7 of abdomen produced posteriorly, often apically

truncate; sternum 7 with pair of bilobed, posterior projections (fig. 8) ; genital

vesica and apodemes 0.25-0.30 mm long. Genitalia in fig. 17.

Female. — Total length 1.72-2.20 mm; wing length 1.50-1.70 mm; colored as

male; chaetotaxy essentially as in male. Spermatheca elliptical; genitalia in

fig. 23.

Distribution. — Maine, Ontario, Quebec. A single male from Lisadale Lake,

British Columbia, 5-VIII-1960, 4000 ft., has extended the range considerably.

The latter locality is in the northwestern corner of the province. Both R. similis and

R. aterrima probably occur in the central Canadian provinces as well.

Rhexoza teskeyi, New Species

Male. — Total length 1.30-1.70 mm; dark greyish brown, head darkest; thorax

and head somewhat shining, abdomen dull; halteres grey; antennal flagellomeres

each with about 14 setae in a single whorl; cardo-stipites with 10-12 setae on

each side; maxillary palpi 0.11-0.12 mm long, reniform; occiput with usual

campaniform sensilla. Supraalar setae 15-19; preepisternals 14-15; anepisternals

20-32; upper episternals 6-9; subalars 7-8; subspiraculars 3-5; lower epimerals

4-6; pedicellars 2-3. Wing length 1.40-1.50 mm; R3 extends 0.50-0.52 of total

wing length from base; R3 with dorsal setae only. Abdominal sterna 1-3 absent,

other terga and sterna present, both densely setose and microtrichiose; tergum 7

produced posteriorly, triangular, with apex shallowly notched (fig. 12) ; sternum

7 with a pair of long narrow, acute processes posteriorly. Genital vesica and

apodemes 0.19-0.20 mm long; genitalia in fig. 19.

Female. — Total length 1.60-1.95 mm; wing length 1.45-1.46 mm; color identical

with that of male. Flagellomeres with 11 setae each in a single whorl; cardo-

stipites with 16 setae on each side; maxillary palpi 0.12 mm long. Supraalar

setae 14; preepisternals 11-13; anepisternals 27-30; upper episternals 6-8; sub-

alars 8; subspiraculars 6-8; lower epimerals 2-3; pedicellars 2-3; wings as in

male. Abdominal sterna 1-3 absent, other terga and sterna present; terga and

sterna setose and densely microtrichiose; tergum 7 with nearly straight posterior

margin; sternum broadly and shallowly emarginate; spermatheca cylindrical to

elliptical, 0.20 mm long. Genitalia in fig. 29.

Holotype male: Stonecliffe, Ontario, coll. 12, Aug. 1962, em. 80, Sept, 1962. Ex

twigs of red pine. In C.N.C. collections. Paratypes: 6 females, 7 males, 1 larva,

2 pupal skins. Same data as holotype. Types in the Canadian National Collections.

The Pan-Pacific Entomologist

Vol. 51, No. 1, January 1975

Rhexoza amaryllis, New Species

A single series of teneral adult females as well as a few larvae and pupae col-

lected from decaying amaryllis bulbs was recently referred to me by H. J. Teskey.

Males. — Unknown.

Females. — Total length about 2.20 mm (specimens in alcohol) ; color grey

brown, dull; halteres grey. Antennae typical, each flagellomere with 14-15

setae in a single whorl; cardo-stipites with 19 setae on each side; maxillary

palpi 0.13-0.14 mm long, reniform. Supraalar setae 13; preepisternal setae 9-11;

anepisternal setae 35-38; upper episternals 7-9; subalars 8-9; subspiraculars

7-8; lower epimerals 10-13; pedicellars 4. Wings 1.65 mm long; R3 extending

0.53 of total wing length; R3 with setae dorsally only; wing densely microtrichiose.

Abdomen with sterna 1-6 undifferentiated; sternum 7 and all terga evident;

terga and sternum 7 setose and microtrichiose; tergum and sternum 7 with straight

posterior margin; spermatheca reniform, 0.21 mm long. Genitalia in fig. 28.

Holotype female: Pt. Credit, Ontario, Nov. 1966, ex. rotting amaryllis bulbs.

Paratypes: 3 females, 4 larvae, 2 pupae, with above data. Types in the Canadian

National Collections.

Rhexoza ryekmani, New Species

Males. — About 1.50 mm long; greyish brown to dark blackish brown, dull to

feebly shining ; halteres dark grey ; wings colorless, shining. Antennal flagellomeres

with 12-15 setae; cardo-stipites with 12-14 setae on each side; maxillary palpi

large, reniform, 0.09-0.12 mm long. Supraalar setae 11-15; preepisternal setae

9-11; anepisternals 23-30; upper episternals 6-8; subalars 6-9; subspiraculars

6-10; lower epimerals 6-10; pedicellars 2-4; wing length 1.20-1.36 mm; R3

terminates from .50-.56 of total wing length; Ml and M2 diverge to wing margin;

setae usually on dorsal side only of R3, occasionally a very few on ventral surface;

wing microtrichiose. Abdomen with sterna 1-4 unidifferentiated, other sterna and

terga present, setose and microtrichiose; tergum 7 triangular, produced pos-

teriorly (fig. 11) ; sternum with 2 elongate, slender processes posteriorly, a small

notch medially (fig. 11) ; genital vesica and apodemes 0.25-0.27 mm long (longer

than other species in the genus) . Genitalia in figs, 20, 21.

Females. — A single female of this genus was collected in the same locality as

a male of R. ryekmani although about 1 month earlier (San Dimas Canyon, Los

Angeles Co., Calif.). This female is distinct from females of Quateieila quatei

and it is also distinct from a single female collected in the type locality of

R. melanderi. The male, female associations of R. ryekmani and R. melanderi, which

follow are certainly tentative, but the association in the case of Quateieila quatei

would seem to be firmly based.

Generally like the male in size, color and chaetotaxy. Antennal flagellomeres

<C-

Figs. 18-27. Genitalia of Rexoza. Fig. 18. R. melanderi, male; Fig. 19. R.

teskeyi, male; Fig. 20. R. ryekmani, dorsal-ventral aspect; Fig. 21. R. ryekmani,

male, lateral aspect; Fig. 22. R. grossa, female; Fig. 23. R. similis, female; Fig. 24.

R. incisa, female; Fig. 25. R. incisa, female, posterior margin of tergum 7; Fig. 26.

R. aterrima, female; Fig. 27. R. borealis, female.

The Pan-Pacific Entomologist

Figs. 28-31. Female genitalia of Rhexoza. Fig. 28. R. amaryllis ; Fig. 29.

R. teskeyi\ Fig. 30. R. melanderi; Fig. 31. R. ryckmani.

each with 12 setae; cardo-stipites with 18 setae on each side; maxillary palpi

0.15 mm long. Supraalar setae 14; preepisternal setae 10; anepisternals 25; upper

episternals 8; subalars 7; subspiraculars 9; lower epimerals 10; pedicellars 4.

Spermatheca broadly elliptical, 0.20 mm long; genitalia in fig. 31; tergum 8 not

longitudinally divided completely but with a median longitudinal suture on

posterior %•

Holotype male: San Bernardino, San Bernardino Co., Calif, 6-VI-1960, Host

Echinocactus polycephalus, reared, R. E. Ryckman et al. Paratypes: 3 males,

data as above; 1 male, same locality but 24-XI-1957 ; 1 male, Verdemont, Calif.,

l-V-1946; 1 male San Dimas Canyon, Los Angeles Co., Calif., 24-XI-1957 ; 1 female,

San Dimas Canyon, Los Angeles Co., Calif., 10-X-1957. San Bernardino specimens

reared in association with numerous Quateiella quatei males and females and

some Coboldia fuscipes (Mg.). Types in the University of Minnesota Collections.

Rhexoza melanderi, New Species

Males. — Total length 1.50 mm ; head and thorax dark brown, feebly shining ;

abdomen brown dorsally, black ventrally; halteres dark grey brown. Antennal

Vol. 51, No. 1, January 1975

flagellomeres each with 12 setae in a single whorl; cardo-stipites with 11 setae

on each side; maxillary palpi large, reniform, 0.12 mm long. Supraalar setae

14; preepisternals 6; anepisternals 19; upper episternals 5; subalars 8; suh-

spiraculars 8; lower epimerals 4; pedicellars 3; wing length 1.30 mm; R3 termi-

nates at 0.42 of wing length ; R3 with setae on dorsal surface only. Abdomen

with sterna 1-4 undifferentiated, other sterna and terga present, both setose and

microtrichiose ; 7 pairs of spiracles; tergurn 7 produced posteriorly; sternum 7

expanded laterally and dorsally; genital vesica and apodemes shorter than in

preceding species. Genitalia in fig. 18.

Female. — Size and color as in male. Antennal flagellomeres each with 15 setae

in a single whorl; cardo-stipites with 12 setae on each side; maxillary palpi 0.11

mm long. Supraalar setae 11; preepisternal setae 8; anepisternals 22; upper

episternals 7; subalars 9; subspiraculars 4; lower epimerals 6; pedicellars 1-2;

wing length 1.50 mm; R3 extends 0.50 of total wing length. Genitalia in fig.

30. Tergum 8 not divided; spermatheca broadly elliptical, 0.13 mm long.

Holotype male: S. Fk. Santa Ana R. Calif., l-IX-1946, A. L. Melander. Para-

type: female, as above but 18-VI-1945. Types in the U.S. National Museum.

Literature Cited

Beekey, C. E. 1938. The immature and adult stages of a new species of

Scalopse from Maine. Entomol. News., 49: 151-154.

Cook, E. F. 1956. A contribution toward a monograph of the Scatopsidae

(Diptera). Part III. The genus Rhexoza. Ann Entomol. Soc. Amer.,

49: 1-12.

1971. The Australian Scatopsidae. Aust. J. Zool. Suppl. Ser., 8: 1-90.

1972. A synopsis of the Scatopsidae of the Palearctic. Part II. Swammer-

damellini. J. Nat. Hist. 6: 625-634.

Enderlein, D. 1936. Scatopsidae. Die Tierwelt Mitteleuropas, 6, Ins., 3: 53-56.

Melander, A. L. 1916. The Dipterous family Scatopsidae. State Coll. Wash.

Agric. Exp. Sta. Bull., 130:1-21.

The Pan-Pacific Entomologist

Observations on the Life Cycle of Heliconius hecale

zuleika (Hewitson) in Costa Rica

(Lepidoptera: Nymphalidae)

Allen M. Young

Department of Biology, Lawrence University, Appleton, Wisconsin 54911

The neotropical butterfly Heliconius hecale (Fabricius, 1775) is one

of the most widespread and variable species of Heliconius in many re-

gions of Central and South America. In the Pacific dry lowlands of

Guanacaste Province, Costa Rica, the form H. hecale zuleika (Hewitson)

(Fig. 1) is very abundant in pockets of moist secondary and riparian

forests along rivers. In fact, densities of this butterfly approach the

unusually high densities of the closely related species, H. ethilla Godart,

reported in some regions of Trinidad (Ehrlich and Gilbert, 1973).

Along with H. ethilla , this species, H. melpomene and others belong

to the “silvaniform” group of Heliconius (Brown and Mielke, 1972;

Ehrlich and Gilbert, 1973). The widespread occurrence of H. hecale

and it’s regional differentiation into distinct subspecies or races is

probably the result of it’s ability to thrive in a variety of different

habitats, including man-made (Brown, 1972), over a wide geographic

area (Brown and Mielke, 1972). Thus there occur no less than fifteen

distinct subspecies in Amazonian and near “extra-Amazonian” Brazil

alone (Brown and Mielke, 1972). The present paper summarizes some

observations on life cycle, larval food plant, and behavior of the sub-

species zuleika in lowland Guanacaste Province, Costa Rica, including

what is apparently the first published account of its immature stages

and larval host. Such information, especially food plant records, is

crucial for understanding the population dynamics of Heliconius butter-

flies, since at least some populations of H. erato in Costa Rica are limited

by the larval food plant (Benson, 1972).

Materials and Methods

The majority of field observations on adult behavior, larval food

plant, and interactions with other heliconiines were done near the town

of Miramar, along the Inter-American highway in Guanacaste Province,

Costa Rica. The actual study site is the “Barranca site” a patch of semi-

deciduous tropical wet forest in the lowlands. Orians (1969) gives

a good description of the site, and it is supplemented by Janzen (1971) .

The Pan-Pacific Entomologist 51 : 76 - 85 . January 1975

Vol. 51, No. 1, January 1975

III!

(III

1111

llll

llHilill

0 _ 1

Fig. 1 . Adult male Heliconius hecale zuleika (Hewitson), dorsal view. Scale

in mm. This specimen was reared in the laboratory from the egg stage.

Activities of H. hecale were generally limited to the thinned-out periph-

eral regions of the forest.

A total of four eggs were collected July 30, 1973 and five were collected

June 23, 1974. Rearing was accomplished by confining the eggs with

fresh clippings of the plant in a single, large, clear-plastic bag kept

tightly shut. The developmental time and other life history events of

individuals confined to the plastic bag were measured from the egg

stage to eclosion. But during this period the caterpillars were moved

to several different localities in Costa Rica, perhaps distorting the

actual developmental time of this species at the Barranca site resulting

from uncontrolled temperature effects.

Results

Habitat and Food Plant. — In my experience, adults of H. hecale

zuleika are most frequently encountered in dense, old secondary forest

The Pan-Pacific Entomologist

Fig. 2. A habitat of H. heccile zuleika in lowland Guanacaste Province, Costa

Rica. The larval food plant, Pcissiflora vitifolia , is found in the dense

secondary forest at the Barranca site. This photograph was taken during the

wet season (July 1973) . Adult H. hecale exhibit some “promenading” around

Passiflora vines here and visit conspicuous red flowers (inset).

(Fig. 2) and in the thinned-out understory of primary forest. The

butterflies are fond of visiting the understory flowering plant Cephaelis

tomentosa Aubl (Vahl) (Rubiaceae) in many parts of Costa Rica

(Fig. 2, inset). Perhaps the prime reason that the butterflies are found

here is the occurrence of the larval food plant in these places: at the

Barranca site the plant used for oviposition and larval development of

H. hecale is Passiflora vitifolia, and it is the same food plant used by

this species at Finca Tirimbina in the Carribbean wet lowlands of

Costa Rica. This plant occurs as a low, vine sprawling over other

plants, both in sunny and shaded places (Fig. 3). This species of

Heliconius is found along with several other heliconiines on P. vitifolia

at the Barranca site, even to the extent that several species occur to-

gether on the same vine. As I have not done a thorough search for other

food plants of this butterfly at the Barranca site, it is quite possible

that others also occur there.

Vol. 51, No. 1 , January 1975

Fig. 3. Passiflora vitifolia (Passifloraceae) , the larval food plant of H. hecale

over much of Central and South America. Note the low position of the vine on

the herbaceous canopy in secondary forest; a large, lobed, older leaf is apparent

directly above the machete near the center of the photograph (June 1974).

P. vitifolia at the Barranca site occurs in small, scattered clumps,

at least 50 meters apart. Over two years, I have found that the same

clumps of this vine (a total of six examined in different parts of the

forest) are infested with heliconians. Some of these clumps are so

small that I would estimate that about 70% of the green foliage is

destroyed during mid- July by heliconian caterpillars and perhaps other

herbivores.

Life Cycle. — The large, squarish egg (Fig. 4- A) is initially light

yellow, darkening to orangish-yellow within one day. The distribution

and numbers of vertical and horizontal ribs are identical to those of

H. melpomene (Beebe, Crane, and Fleming, 1960). The egg is 1.4 mm

high by 0.9 mm wide at the middle. The egg stage lasts five days.

The first instar larva is about 4.0 mm long at hatching, translucent,

dirty-orange with a light tan head capsule. It is covered with many

fine setae, and very similar to the larva of H. melpomene (see Beebe,

Crane, and Fleming, 1960). The second instar (Fig. 4-B) is similar

The Pan-Pacific Entomologist

Fig. 4. The juvenile stages of H. hecale zuleika. A. egg; B. second instar

caterpillar; C. third instar; D. fourth instar; E. fifth instar; F. pupa; G. eclosion.

Sizes are given in the text.

to the first, but the head capsule is more orangish and margined pos-

teriorly lightly in black. The prothoracic plate is brown and all scoli

and legs are black. The distribution of scoli is set in this instar: the

rows of dorsal, lateral, supralateral, and sublateral scoli are all apparent

(see the excellent diagram for scoli distribution of Heliconius in Beebe,

Vol. 51, No. 1, January 1975

Crane, and Fleming, 1960). The first instar lasts three days and the

second instar lasts four days.

The third instar (Fig. 4-C) is very similar to the fourth instar (Fig.

4-D) but lighter in the orange color of the body. The head capsule

is light orange and the trunk region is light bluish-white above and

orange below. The head scolus is black and about 1.8 mm long, curved

strongly to the posterior end of the body. The anal plate is light orange

and all scoli are black and bear spinules. The third instar reaches a

length of 16 mm in four days. The general appearance is very similar

to that of H. melpomene. The whitish regions of the body become

more pronounced in the fourth instar (Fig. 4-D) and the recurved

head scoli are now about two mm long. All scoli are black and longer

in this instar (Fig. 4-D). The prolegs remain orange as in the previous

instars. This instar attains a length of about 27 mm in four days.

The fifth instar (Fig. 4-E) is strikingly different in appearance

from the previous stages. The body is now chalky-white and the head

capsule glossy light orange. The recurved head scolus is now almost

6 mm long and all scoli are black. All spiracles are black and a lateral

black dot occurs at the posterior edge (fold) of each segment; a second,

less distinct black spot occurs just above the sublateral scolus. The

fifth instar attains a length of about 38 mm in six days, and it is strik-

ingly similar in appearance to the fifth instar of H. cydno (see Young,

1973 1 ) .

The pupa (Fig. 4-F) fits into the general description of “Group C”

Heliconius in Beebe, Crane, and Fleming (1960) ; it is very similar to

the pupae of H. melpomene and H. numata. The pupae of this group

are characterized by (1) spines on the thorax, abdomen, and antennae,

(2) subdorsal flanges on the abdomen, and (3) three to five pairs of

gold-colored spots on the thorax and abdomen. In H. hecale there are

three pairs of gold spots on the abdomen. The length of the pupa is

33 to 34 mm and the general coloration is mottled light and dark brown.

The spines on the ventral side of the abdomen are very long and black.

The subdorsal flanges are very well developed on the first two ab-

dominal segments. The first five abdominal segments bear alternating

long and short spines, a single spine on the first segment, pairs of

spines on succeeding segments. The head region is coarsely sculptured

with a pair of distinct anterad projections. Along the dorsal line of

each wing pad there is a series of short, curved black “hooks” that

continue to the prothoracic midline and to the head (ventrally). The

1 The fifth instar was incorrectly labelled in that publication as the third instar (switch captions

for Figs. 2 B-C, p. 243).

The Pan-Pacific Entomologist

cremaster silk is orange. When touched, the pupa is capable of violent

twitching movements from side to side. It is not known if the pupa

is capable of sound or odor production although Alexander (1961)

mentions that the pupa of the closely related H. melopmene does both.

The pupal period lasts 10 to 14 days, falling within the upper limit

of range reported for Group C pupae (9-10 days). Eclosion (Fig.

4-G) is rapid and the first meconium is brownish; Alexander (1961b)

reports that the first meconium of the related species H. melpomene

ranges from brown to chestnut.

Behavior of Adult Butterflies. — -Oviposition behavior usually

entails the female flying very low through an area where the larval

food plant is found. A female may fly back and forth through the

area many times before actually landing and depositing an egg. I have

noticed similar prolonged pre-oviposition flight habits in Philaethria

dido and other unidentified Heliconius. Eventually an egg is laid

on either a very young (unfolding) leaflet or tendril; in a total of four

oviposition acts observed, the egg was never laid on older leaves and

other structures.

Adults are usually encountered singly during the morning and I did

not determine if nocturnal communal roosting, known for some species

of Heliconius (see summary in Benson, 1971), occurs in H. hecale.

Adults probably visit a variety of flowers, but are known to be attracted

to the bright red flowers of Cephaelis tomentosa (Fig. 2-inset) . At

least two closely related species, H. ethilla and H. melpomene show

strong preferences for red flowers in Brazil (Brown and Mielke, 1972).

Collected adults have been noted with at least two types (colors) of

pollen, also suggesting lack of food plant specificity.

Behavior of Caterpillars. — As with species closely related to

this butterfly (see Alexander, 1961a; Brown and Mielke, 1972), the

caterpillars of H. hecale are solitary, a condition very likely the result

of single oviposition. It often happens that caterpillars in different

instars will be found on the same vine of P. vitifolia, but generally

highly scattered with no apparent interaction. It is likely that a single

vine is exploited repeatedly throughout the year for egg-laying by

this species and several other Heliconius. The youngest caterpillars

are invariably found on terminal and subterminal leaflets and even

tendrils. Older caterpillars also feed on subterminal leaflets (Fig. 4-D),

although fifth instars are generally found on older leaves. Caterpillars

generally feed from undersides of leaves, although I have seen fifth

instar caterpillars on the dorsal side of drooping older leaves of the food

plant. Such individuals are extremely conspicuous. Caterpillars often

Vol. 51, No. 1, January 1975

share adjacent leaflets and leaves with other heliconians on P. vitifolia :

at Finca Tirimbina, H. hecale co-occurs with at least Philaethria dido

(Young, 1974), while at the Barranca site it co-occurs with H. cydno,

H. isemenius , and Dione.

Discussion

As one of the “silvaniform” species of Heliconius , H. hecale displays

close morphological resemblance in the early stages to other members

of this group, such as H. ethilla (formerly numata ) and H. melpomene.

The life cycle characteristics described for H. hecale agree with the

general features of Heliconius biology as summarized by Brown and

Holzinger (1973) : eggs are usually laid on young (“growing meristem”)

structures, and adults collect pollen. The egg is large and yellowish,

and the head capsule does not bear stripes. The pupa usually hangs

vertically.

The fifth instar bears a striking resemblance to H. cydno and it is

interesting to speculate that the two species participate in a Mullerian

mimicry complex as caterpillars, especially since both species occur on

the same food plant and in the same habitats in Costa Rica. Mimicry

has been suggested by Brown (1972) with respect to other Heliconius

in Brazil.

Brown and Mielke (1972) comment that H. hecale is very widespread

in Brazil, not only geographically but also in terms of habitats; the

apparently high degree of distinct sub-speciation in this species in

Brazil is believed to be the result of phenotypic flexibility perhaps

coupled with restricted gene flow. In Central and South America, the

widespread occurrence of H. hecale , clearly far more extensive than

many other species of Heliconius , must be due in part to the wide-

spread occurrence of Passiflora vitifolia and other food plants. Keith S.

Brown, Jr. (pers. comm.) states that P. vitifolia is used by H. hecale

throughout Central America and Colombia and as far as Ecuador.

Young (1974) discussed P. vitifolia as a larval food plant of Philaethria

dido in northeastern lowland Costa Rica. At least three genera of

heliconians and a large number of species (of Heliconius ) use P.

vitifolia , probably for the following reasons: (1) the widespread geo-

graphical distribution of the plant makes it an attractive resource for

heliconians, (2) the plant species occupies several different habitats

within a geographic (regional) zone, and (3) the plant is unusually

low in those defense systems that normally deter heliconians and perhaps

other herbivorous insects. Whatever the precise reasons for it’s abun-

dance over much of tropical America, this and perhaps other species of

The Pan-Pacific Entomologist

Passiflora are easily penetrated (in the evolutionary or ultimate sense

of Ehrlich and Raven, 1964) by local complexes of heliconians.

Acknowledgements

This research is a by-product of National Science Foundation grant

GB-33060 with logistic support from the Costa Rican program of the

Associated Colleges of the Midwest. The butterfly was identified by

Dr. Lee D. Miller (Allyn Museum of Entomology) and the larval food

plant by Dr. Keith S. Brown, Jr. (Rio de Janeiro), who made several

helpful comments regarding his studies of the butterfly in Brazil. From

Brown I have also learned that several researchers are doing extensive

studies on the ecology and genetics of this butterfly.

Literature Cited

Alexander, A. J. 1961a. A study of the biology and behavior of the cater-

pillars, pupae and emerging butterflies of the subfamily Heliconiinae

in Trinidad, West Indies, Part I. Some aspects of larval behavior.

Zoologica, 46: 1-24.

1961b. Part II. Molting, and the behavior of pupae and emerging adults.

Zoologica, 46: 105-125.

Beebe, W., J. Crane, and H. Fleming. 1960. A comparison of eggs, larvae and

pupae in fourteen species of Heliconiine butterflies from Trinidad,

W. I. Zoologica, 45: 111-153.

Benson, W. W. 1971. Evidence for the evolution of unpalatability through

kin selection in the Heliconiinae (Lepidoptera). Amer. Nat., 105:

213-226.

1972. Natural selection for Mullerian mimicry in Heliconius erato in Costa

Rica. Science, 176: 936-939.

Broavn, K. S., Jr. 1972. The Heliconians of Brazil (Lepidoptera: Nymphalidae) .

Part III. Ecology and biology of Heliconius natter eri, a key primitive

species near extinction, and comments on the evolutionary develop-

ment of Heliconius and Eueides. Zoologica, 57: 41-69.

Broavn, K. S., Jr. and 0. H. H. Mielke. 1972. Part II. Introduction and general

comments, Avith a supplementary revision of the tribe. Zoologica, 57 :

1-40.

Brown, K. S., Jr. and H. Holzinger. 1973. Part IV. Systematics and bioloy

of Eueides tales Cramer, with description of a new subspecies from

Venezuela. Zeitschr. Arbeit. Osterr. Entomol., 24: 44-64.

Ehrlich, P. R. and P. H. Raven. 1964, Butterflies and plants: a study in

coevolution. Evolution, 18: 586-608.

Ehrlich, P. R. and L. E. Gilbert. 1973. Population structure and dynamics

of the tropical butterfly Heliconius ethilla. Biotrop., 5: 69-82.

Emsley, M. G. 1965. Speciation in Heliconius (Lep., Nymphalidae) : morphology

and geographic distribution. Zoologica, 50: 191-254.

Gilbert, L. E. 1972. Pollen feeding and reproductive biology of Heliconius

butterflies. Proc. Nat. Acad. Sci., U.S.A., 69: 1403-1407.

Vol. 51, No. 1, January 1975

Janzen, D. H. 1971. The fate of Scheelea rostrata fruits beneath the parent

tree: predispersal attack by bruchids. Principes, 15: 89-101.

Orians, G. H. 1969. The number of bird species in some tropical forests.

Ecol., 50: 783-801.

Young, A. M. 1973. Notes on the biology of the butterfly Heliconius cydno

(Lepidoptera: Heliconiinae) in Costa Rica. Wasmann J. Biol., 31:

337-350.

1974. Further observations on the natural history of Philaethria dido dido

(Lepidoptera: Nymphalidae: Heliconiinae). J. New York Entomol.

Soc., 82: 30-41.

RECENT LITERATURE

Biology, Ecology, and Host Specificity of Microlepidoptera Associated

with Quercus agrifolia (Fagaceae). Paul A. Opler. University of California

Publications in Entomology, Vol. 75, 83 pp., 38 figs., 7 plates, 8 tables. $4.25.

Taxonomically, the microlepidoptera remain one of the most poorly known groups

of insects, as demonstrated by the fact that 57% of the species treated in this study

were undescribed at the time the research was conducted. Nevertheless, this work

is primarily ecological, containing no descriptions of new species, and no keys.

The text contains a wealth of documentary data concerning interrelationships be-

tween the moths, their host, and important climatic and distributional variables.

This descriptive information is balanced with extensive discussion of life cycle

adaptations, host specificity, biogeographic distribution, and evolution. The diversity

of biological adaptations of the moths to avoid unfavorable seasons or to reduce

interspecific competition reveal these small insects, usually ignored by all but

specialized taxonomists, to be highly interesting and significant subjects for

ecological analysis. The evolution of the moths is related to evolution of their host,

emphasizing the fossil record of the oak and the host relationships and present

distribution of the insects. The antiquity of the leafmining habit is documented by

Miocene fossil impressions which are practically indistinguishable from mines of

extant moths. The text is accompanied by useful tables and figures comparing the

chronology of development of different species, host relationships, and details of

geographic distribution. Plates include photographs of examples of larval mines

and shelters and eggs in typical oviposition sites. — Editor.

Revision of the Millipede Family Andrognatiiidae in the Nearctic Region

(Diplopoda: Platydesmida) . M. R. Gardiner. Memoirs of the Pacific Coast

Entomological Society, Vol. 5, 61 pp., 46 text figures. (Available from Pacific

Coast Entomological Society, c/o California Academy of Sciences, Golden Gate

Park, San Francisco, California 94118. $3.00).

Taxonomic study of the five nearctic genera of platydesmid millipeds, including

one new species from northern California.

The Pan-Pacific Entomologist

Fifty Years of the Pan-Pacific Entomologist 1

E. Gorton Linsley

Division of Entomology and Parasitology

University of California, Berkeley 94720

Table of Contents— V

Historical and Biographical Papers

Arnaud, P. H., Jr.

Herbert Ruekes, Sr. 42:156

Carl J. Drake 42:244

Edwin Ralph Leach 47:312

G. Dallas Hanna 48:59

Arnaud, P. H., Jr. and M. N. Wale

Thomas Wrentmore Cook 49:177

Armitage, H. M.

George Ringo Wilson 14:143

Bailey, S. F.

Jones collection of Thysanoptera 13:89

Hinds collection of Thysanoptera 15:91

Dudley Moulton 27:145

Blackwelder, R. E.

Entomological work of Adalbert Fenyes 18:17

Blaisdell, F. E., Sr.

Thomas Lincoln Casey 2:90

Blaisdell, F. E., Sr. and E. C. Van Dyke

Ralph Hopping 18:1

Brown, F. M.

Letters from W. H. Edwards to W. G. Wright 39:198

Eaton, C. B. and G. R. Struble

John Elliot Patterson 40:1

Edmundson, G. F.

Willis C. Day 42:163

Essig, E. 0.

Charles Fayette McGlashan 7:97

Entomology and early development of agriculture 10:1, 49, 97

Charles William Woodworth 17:30

University of California Entomological Society 17:73

Itinerary of Walsingham in California and Oregon 17:97

Sol Felty Light 24:49

Theodore Dru Alison Cockerell 24:117

Edwin Cooper Van Dyke 29:73

Essig, E. O., et al.

History of the Pacific Coast Entomological Society 27:97

Essig, E. O. and R. L. Usinger

Edward Payson Van Duzee 16:145

Fall, H. C.

Adalbert Fenyes 13:145

1 Concluded from vol. 50, p. 415.

The Pan-Pacific Entomologist 51: 86-87. January 1975

Vol. 51, No. 1 , January 1975

Furman, D. P.

William Brodbeck Hermes 25:192

Hottes, F. C.

Little known work of Ph. F. Gmelin 25:83

Hurd, P. D., Jr.

Claude “I” Smith 26:59

Leach, E. R.

F. W. Nunenmacher 24:1

James Edward Cottle 32:19

Leech, H. B.

Harry Phytander Chandler 33:31

Bibliography of G. F. Ferris 35:29

Roy Seymour Wagner 43:134

Linsley, E. C.

Henry Clinton Fall 16:1

James Otis Martin 28:71

Robert Leslie Usinger 45:167

McKenzie, H. L.

G. F. Ferris as a student of scale insects 35:25

Michelbacher, A. E.

Edward Oliver Essig 41:207

Miller, D.

Apropos C. V. Riley 22:28

Miller, D. R., R. M. Bohart and R. F. Wilkey

Howard Lester McKenzie 45:245

Rentz, D. C.

Harold J. Grant, Jr. 42:158

Ross, E. S.

Louis S. Slevin 22:141

Smith, R. F.

Bibliography of E. 0. Essig 41:23

Telford, H. S.

Ralph Ward Dawson 50:209

Tilden, J. W.

Carl Dudley Duncan 43:97

Usinger, R. L.

Brighton Clark Cain 28:125

Howard Madison Parshley 30:1

Gordon Floyd Ferris 35:1

Usinger, R. L. and E. C. Zimmerman

Otto Herman Swezey 36:151

Van Duzee, E. P.

Walter M. Gifford 6:46

Millard Carr Van Duzee 10:90

Van Dyke, E. C.

Frank Ellsworth Blaisdell, Sr. 23:49

Julius George Grundel 10:48

Wiggins, I. L.

G. F. Ferris, the teacher 35:13

Zimmerman, E. C.

Francis Xavier Williams ; 45:135

The Pan-Pacific Entomologist

PACIFIC COAST ENTOMOLOGICAL SOCIETY

C. B. Philip H. V. Daly F. Ennik P. H. Arnaud, Jr.

President President-Elect Secretary Treasurer

Proceedings

Three Hundred and Fifty-Sixth Meeting

The 356th meeting was held Friday, 18 January 1974 in the Morrison Audi-

torium of the California Academy of Sciences, Golden Gate Park, San Francisco,

with President Philip presiding.

Members present (44) : F. G. Andrews, P. H. Arnaud, J. W. Bass, M. M.

Bentzien, L. G. Bezart, F. L. Blanc, R. G. Blair, I. A. Boussy, R. J. Bushnell,

W. Carstens, H. J. Denk, D. G. Denning, F. Ennik, B. Ehreth, B. K. Eya, M. R.

Gardner, A. R. Gillogly, J. Guggolz, K. S. Hagen, R. M. Haradon, Jr., L. Hawkins,

T. W. Hewton, Jr., A. I. Kaplan, C. Y. Kitayama, R. L. Langston, H. B. Leech,

K. L. Lipps, A. C. Mehrer, R. L. Morrison, D. W. Moss, Jr., G. M. Nishida, R. W.

Pemberton, R. L. Peterson, C. B. Philip, M. J. Robey, W. E. Savary, H. I. Scudder,

M. V. Silveira, R. W. Thorp, R. E. Stecker, T. Toren, D. J. Voegtlin, M. Wasbauer

and S. P. Welles.

Visitors present (22) : N. Arnaud, M. C. Beugler, N. Blair, F. Blanc, A. Bush,

J. A. Cate, T. L. Delgman, L. Denning, S. Ferguson, J. Hawkins, K. S. Heston,

A. Hunter, F. 0. Lewis, R. Locatelli, V. Maggi, V. Mayer, M. Mehrer, E. Parker, K.

Taylor, J. T. Thorp, D. Veirs, and J. Wasbauer.

The minutes of the meeting held 14 December 1973 were summarized.

The following persons were elected to membership. Student membership: Don

R. Harris, Ron Locatelli, Mona Lyon, Mark A. Santana. Regular membership:

Kathleen Hesketh, Albert Mehrer and Mike Mehrer.

Dr. W. E. Ferguson exhibited slides of an undescribed thrombidiid mite para-

sitic on camel crickets which is being studied at California State University, San

Jose, by visiting Professor Dr. Pierre Robaux.

A. I. Kaplan presented slides of a dipterous larvae, collected 29 November

1973, from a rotting and liquified fungus found on the University of California,

Berkeley campus. The fungus was tentatively identified as Armillaria sp.,

and the flies were Calotarsa insignis Aldrich, a platypezid, or flat-footed fly.

Laboratory reared adults first emerged 10 January 1974.

The principal speaker of the evening was Dr. William J. Hamilton, III, Pro-

fessor of Animal Ethology, University of California, Davis. His illustrated talk

was entitled, “Behavioral adaptations of the tenebrionid fauna of Africa’s Namib

Desert.”

Coffee and other refreshments were served during a social hour held in the

Trustees Room following the meeting. — F. Ennik, Secretary.

Three Hundred and Fifty-Seventh Meeting

The 357th meeting was held Friday, 15 March 1974 in the Morrison Audi-

torium of the California Academy of Sciences, Golden Gate Park, San Francisco,

with President Philip presiding.

Vol. 51, No. 1, January 1975

Members present (30) : F. G. Andrews, P. H. Arnaud, Jr., J. W. Bass, L. G.

Bezart, R. W. Brown, R. J. Bushnell, H. V. Daly, M. R. Ebertz, J. G. Edwards,

F. Ennik, W. E. Ferguson, B. W. Grushkowitz, R. W. Hall, R. M. Haradon, Jr.,

T. E. Hewton, Jr., C. Y. Kitayama, H. B. Leech, R. G. Locatelli, A. E. Michelbacher,

R. L. Morrison, C. B. Philip, W. E. Savary, R. X. Schick, E. L. Smith, R. E.

Somerby, R. E. Stecker, S. L. Szerlip, M. Wasbauer, C. Williams, and S. C.

Williams.

Visitors present (21) : A. Bush, M. Bush, T. L. Delgman, A. N. Ennik, A. V.

Ennik, S. Ferguson, A. R. Gittins, R. Greek, D. S. Green, D. Grushkowitz, J.

Hawkins, R. Lane, V. Maggi, G. Malleck, M. Michelbacher, Mr. & Mrs. J. J.

Moran, E. Parker, E. J. Rogen, Jr., D. Veirs, and J. Wasbauer.

The minutes of the meeting held 18 January 1974 were summarized.

Two student members were elected to membership: Deborah S. Green and

Donald W. Scott.

Dr. E. L. Smith exhibited an unidentified sawfly of the family Pamphiliidae,

collected in Golden Gate Park, and summarized some of the biology of this

primitive group of Hymenoptera.

The Executive Board decided that the social hour should be held before the

next regular meeting at 7:45 p.m. in the Trustees Room, rather than after, to

expedite meetings. The membership meeting will begin at 8:30 p.m. Dr. R. E.

Stecker stated the opinion that students benefited greatly from exposure to the insect

collections and wanted to know if access to the collections would be possible at

future meetings. Both Dr. Arnaud and Dr. Smith replied that space in the insect

rooms did not really allow study of the collections and socializing at the same

time due to the large attendance of meetings. Access to the collections will be

available before the regular meeting for those who wish, however.

The principal speaker of the evening was Dr. Arthur R. Gittins, Department

of Entomology, University of Idaho. His illustrated talk was entitled, “A brief

history of entomology at Idaho.”

Coffee and other refreshments were served during a social hour held in the

Trustees Room following the meeting.— F. Ennik, Secretary.

Three Hundred and Fifty-Eighth Meeting

The 358th meeting was held Friday, 19 April 1974 in the Morrison Auditorium

of the California Academy of Sciences, Golden Gate Park, San Francisco, with

President-elect Daly presiding.

Members present (34) : J. Anduiza, P. H. Arnaud, Jr., J. W. Bass, L. G. Bezart,

D. Burdick, W. Carstens, P. S. Crane, H. V. Daly, D. G. Denning, M. Eastman,

J. G. Edwards, F. Ennik, B. K. Eya, W. E. Ferguson, L. Green, K. S. Hagen, T. E.

Hewton, Jr., A. I. Kaplan, C. Kitayama, D. H. Kavanaugh, R. L. Langston, V. F.

Lee, K. L. Lipps, R. Morrison, H. G. Neal, R. W. Pemberton, R. L. Peterson,

W. W. Pitcher and wife, M. V. Silveira, J. Spalding, E. L. Smith, D. Whitman, and

S. C. Williams.

Visitors present (24) : J. Bauer, C. Bera, A. Bush, E. J. Bynum, D. W. Caneto,

L. Denning, J. T. Duncan, A. L. Edwards, A. V. Ennik, S. Ferguson, P. Grant,

R. Hansen, N. Hays, A. Horning, C. A. Larrea, G. J. Mallick, M. Martin, G.

Perrier, P. Phillip, R. G. Repke, C. Spencer, J. Stockton, S. Stockton and F. Wray.

The minutes of the meeting held 15 March 1974 were summarized.

The Pan-Pacific Entomologist

The following persons were elected to membership. Student membership: John

H. Benedict, Marion E. Buegler, Marli Eastman, Peter Goodall, Ronald J. Mc-

Ginley, Carol Musgrave, Peggie Phillips, Tom Sparks, and James E. Tobler,

I. Regular membership : David George, Donald S. Leson and W. F. Peregrin.

Dr. Arnaud announced that Memoir Series Volume 4, Autobiography of an

Entomologist by R. L. Usinger is still available at $12.50 to members and $15

to non-members. Approximately 350 copies have been sold.

Dr. Arnaud announced that Dr. David H. Kavanaugh will replace Mr. Hugh

B. Leech, on his retirement, as curator of Coleoptera at the California Academy

of Sciences.

The following note was presented:

A New Record of Mantispid Reared from Spider. — On 24 March 1974,

while collecting along the Kings River, near Piedra, Fresno County, California,

a lycosid spider. Tarantula sp., was seen carrying an egg sac. The spider was

captured and the egg sac opened. Within the sac a pupa of the mantispid

Climaciella brunnea (Say) was found. The pupa was brought to the laboratory,

wrapped in tissue and placed in a rearing cage in constant shade.

On 8 April at 11 a.m. the pupa was seen walking over the tissue, which had a

hole chewed through it, and molted into the adult form by 5 p.m.

The spider and mantispid, with its pupal case, have been preserved and will

be given to the California Academy of Sciences. A brief description of a study

of a European mantispid, that develops within the egg sac of a lycosid spider,

Lycosa sp., is given in Imms, “ A general textbook of entomology ” (0. W.

Richards and R. G. Davies (Eds.), 9th Edition, 1957. pp. 498-9). The ambulatory

behavior of mantispid pupae has been reported for Plega banksi (Werner and

Butler, 1965, Ann. Entomol. Soc. Amer. 58: 66-68). Lloyd D. George and Nelda

L. George.

The principal speaker of the evening was Dr. Karen L. Lipps, Department of

Ecology and Systematics, California State University, San Francisco, whose il-

lustrated talk was entitled, “The evolution of cleaning behavior in insects,

particularly Drosophila.”

Coffee and other refreshments were served during a social hour held in the

Trustees Room prior to the meeting. — F. Ennik, Secretary.

Three Hundred and Fifty-Ninth Meeting

The 359th meeting was the annual field day and picnic held on Saturday, 18

May 1974 at Morton’s Warm Springs, Kenwood, California.

Members present (15) : E. Anderson, P. H. Arnaud, Jr., J. W. Bass, D. G.

Denning, F. Ennik, W. C. Ferguson, R. L. Langston, K. L. Lipps, A. E. Michel-

bacher, W. W. Pitcher and wife, C. B. Philip, W. E. Savary, M. S. Wasbauer

and son.

Visitors present (18) : S. Anderson, N. Arnaud, P. H. Arnaud, S. Arnaud,

J. M. Bass, L. Denning, T. Delgman, J. T. Duncan, A. N. Ennik, J. S. Ennik,

A. V. Ennik, S. Ferguson, M. Michelbacher, G. Philip, P. Robaux, 0. Robaux, C.

Sinclair, and K. Taylor.

Although it was raining lightly at times, picnicking, swimming, and insect

collecting were the main activities.— F. Ennik, Secretary.

Vol. 51, No. 1 , January 1975

Three Hundred and Sixtieth Meeting

The 360th meeting was held Friday, 18 October 1974, in the Morrison Audi-

torium of the California Academy of Sciences, Golden Gate Park, San Francisco,

with President Philip presiding.

Members present (35+) : L. C. Armin, P. H. Arnaud, Jr., J. W. Bass, I. A.

Boussy, T. Briggs, R. W. Brown, D. Durbin, J. G. Edwards, F. Ennik, D. S.

Green, R. M. Haradon, Jr., L. Hawkins, T. E. Hewton, Jr., A. I. Kaplan, D. H.

Kavanaugh, R. L. Langston, H. B. Leech, K. L. Lipps, G. J. Mallick, J. B.

McCarthy, R. L. Morrison, D. W. Moss, Jr., G. M. Nishida, R. W. Pemberton,

C. B. Phillip, W. W. Pitcher, J. A. Powell, H. G. Real, M. J. Robey, E. S. Ross,

W. E. Savary, C. M. Sprenger, S. L. Szerlip, R. W. Thorp, and B. Villegas.

Visitors present (56+): K. Barth, J. M. Bass, R. X. Bettancourt, Jr., J. P.

Boban, K. Brittan, J. Brown, K. W. Brown, P. Brown, B. & P. Buickerood, R. B.

Chorf, J. Crane, C. M. Dixon, J. T. Duncan, A. Edwards, A. N. Ennik, A. V.

Ennik, J. S. Ennik, R. French, T. Friebel, J. C. & K. L. Gerth, T. Gibson, R. D.

Gunk, J. Hjelle, J. & Q. Illige, J. Jacobson, J. Johnson, B. A. Kavanaugh, B.

Lasater, J. Locarnim, G. Low, D. McKay, D. McManus, P. Michelson, W. Moore,

F. Murphey, J. Pang, D. L. Parker, J. B. Parker, P. Philip, P. & K. Pitcher, E. L.

Rhe, G. Sahara, C. Sawley, P. & H. Sherman, P. F. Smith, A. Stephen, W. A.

Sumner, J. T. Thorp, K. Uremorrie, N. Vandenberg, and K. R. Wahl.

The minutes of the annual field trip and picnic, held 18 May 1974, were

summarized.

The following persons were elected to membership. Student membership:

David L. Kulhavy, Ralph J. Michels, Wayne S. Moore, Lee C. Ryker, and

D. Dee Wilder. Regular membership: Paul A. Collins, William E. Gavin, Dennis

Groothuis, George J. Mallick, Dr. William L. Peters and Dr. Thomas J. Zavortink.

Dr. Arnaud announced that Dr. Thomas J. Zavortink has been appointed new

curator of Hymenoptera and alcohol collections at the Academy.

Dr. Arnaud asked that Secretary Ennik send Mrs. Hawley a letter on behalf

of the Society on her speedy recovery from recent surgery.

The following notes were presented:

A Remarkable New Brachypterous, Sand-Dune Moth. — Slides illustrating

an undescribed genus and species of Scythridae from the coastal Santa Maria

dunes in southern San Luis Obispo County, California were exhibited. Bizarre

activities of the flightless, sand-colored moths, including behavioral thermoregula-

tion and self-burial that occurs each evening, as well as the unique larval galleries

which attach to buried parts of various plants in loose sands were shown. The

species and its habits are being described elsewhere. — J. A. Powell, University

of California, Berkeley.

New Record for a Nearctic Deerfly in Baja California. — Early last June

Franklin Ennik and I made a collecting trip into Baja California Norte. Three

females of Chrysops coloradensis Bigot were captured in a small Malaise trap,

COa baited, at midday, 5 June, on a dry lake bed one mile north of Laguna Hanson

in the Sierra Juarez, 5500 ft. altitude. This must be a disjunct colony, as no

previous records are known in Arizona or south of 3 montane localities in Ventura

and San Bernardino Counties in California. The species extends on its southern

boundary eastward through Utah into the Colorado Rocky Mountains, and

northwest into southern British Columbia.

With more adequate collecting, it seems likely that C. coloradensis will also

The Pan-Pacific Entomologist

be taken farther south in the Sierra San Pedro Martir. In the latter mountains,

rare specimens have been taken of other generalized tabanids: Pilimas abaureur

(Phil.), Silvius gigantulus (Lw.), C. coquilletti Hine and C. clavicornis Bren., all

similarly disjunct from the montane California fauna farther north. It appears

not unlikely that these represent relicts from past, more verdent geologic times

when biologic faunal interchange would have been more feasible than it is now.

Only one female of Apatolestes comastes willistoni Bren, was taken at Mike’s

Sky Ranch though we followed a route which Stan Williams and Kathy Blair

had taken the year before about the same dates when they netted 16 specimens in

4 species of this generalized genus of tabanids. We were told by stock owners at

Melling Ranch and elsewhere that the season was late and drier than usual, and

that we appeared to be some two weeks too early for the expected horsefly

pests of their livestock.

Expenses for this field trip were in part defrayed by a grant (no. 1071, Johnson

Fund) from the American Philosophical Society, Philadelphia. — Cornelius B.

Philip, California Academy of Sciences, San Francisco, CA.

The principal speaker of the evening was Dr. Edward S. Ross, Curator, Depart-

ment of Entomology, California Academy of Sciences, San Francisco. His illus-

trated talk was entitled, “Insect photo trophies, 1974: a series of insect and

arachnid slides with annotated biological notes.”

Coffee and other refreshments were served during a social hour held in the

Trustees Room following the meeting. — F. Ennik, Secretary.

Three Hundred and Sixty-First Meeting

The 361st meeting was held Friday, 15 November 1974, in the Morrison Audi-

torium of the California Academy of Sciences, Golden Gate Park, San Francisco,

with President Philip presiding.

Members present (34) : P. H. Arnaud, Jr., F. R. Barnes, J. W. Bass, L. G.

Bezart, K. Corwin, C. Dailey, J. G. Edwards, F. Ennik, K. S. Hagen, T. E.

Hewton, Jr., A. I. Kaplan, D. H. Kavanaugh, R. L. Langston, H. B. Leech, K. L.

Lipps, A. E. Michelbacher, R. L. Morrison, D. W. Moss, Jr., G. M. Nishida,

R. W. Pemberton, C. B. Philip, P. Phillips, W. W. Pitcher, J. A. Powell, M. J.

Robey, E. S. Ross, W. E. Savary, T. N. Seeno, C. M. Sprenger, R. E. Stecker,

S. L. Szerlip, T. Toren, D. D. Wilder, and D. L. Wilson.

Visitors present (33) : J. Alletts, R. X. Bettancourt, Jr., K. Brittan, B. P.

Buickerood, A. Bush, D. Chase, S. Cohen, J. Cramer, C. E. Culley, C. Dailey,

A. Edwards, W. J. Evans, T. Gibson, B. Kavanaugh, P. E. Kenmore, K. A. Kesketh,

K. Nishida, P. Mason, D. McKay, M. Michelbacher, K. Neath, V. A. Nunez, B. C.

Osborne, D. L. Parker, J. B. Parker, K. Pitcher, P. Pitcher, J. Redman, M. D.

Smart, A. Stephen, L. Toren, P. Toren, and C. Wilson.

Two student members were elected to membership: George Erwin and Paul

Mason.

President Philip appointed Hugh B. Leech Chairman of the 1975 Nominating

Committee, to be assisted by Donald Denning and A. E. Michelbacher.

For the Auditing Committee, President Philip appointed H. V. Davis, Chairman,

to be assisted by E. S. Ross.

President Philip appointed an ad hoc committee consisting of E. G. Linsley,

Chairman, assisted by H. B. Leech, E. S. Ross and W. W. Middlekauff, to explore

Vol. 51, No. 1, January 1975

ways the Society could participate in the upcoming joint City of San Francisco-

National Bicentennial celebration in 1976.

J. Gordon Edwards exhibited two live female beetles, Pleocoma conjungens

and P. hirticollis vandykei , collected during recent rains in Quail Hollow, Santa

Cruz Mountains and Coyote, California. The males are attracted to the flightless

females, apparently by a non-species specific pheromone, as the females sit par-

tially buried in the opening of their burrows. After mating, the female digs back

into the ground and deposits her eggs. The larvae eat plant roots during their long

period of development. Because the females do not move from their point of

origin, Pleocoma species have limited geographic ranges.

H. B. Leech stated that Pleocoma beetles are also found at his home in Angwin,

CA.

D. H. Kavanaugh reported on his field and laboratory observations on the carabid

genus Nebria, which are restricted to snow melt trickles or cool mountain stream

margins of the Sierra Nevada above the 11,500 ft. level.

S. L. Szerlip exhibited slides of two species of Diptera found living in the

California pitcher-plant, Darlingtonia calif ornica Torrey, in a Sphagnum bog near

Willow Springs, Nevada County, California, in June, 1974. Larvae of Metriocnemus

edwardsi Jones (Chironomidae) are apparently found only in D. californica,

where they feed on detritus. Larvae of Leptocera sp. (Sphaeroceridae) as well as

pupae and adults, were also found, and appear to be completely adapted to the

living conditions in the pitcher-plants.

The main speaker of the evening was Dr. Ronald E. Stecker, Department of

Entomology, San Jose State University. His illustrated talk was entitled, “Life

in the 300 ft. giant Sequoia.”

Coffee and other refreshments were served in a social hour, held in the Trustees

Room following the meeting. — F. Ennik, Secretary.

Three Hundred and Sixty-Second Meeting

The 362nd meeting was held Friday, 20 December 1974 at 7:45 p.m. in the

Morrison Auditorium of the California Academy of Sciences, Golden Gate Park,

San Francisco, with President Philip presiding.

Members present (39) : R. P. Allen, E. Anderson, L. C. Armin, P. H. Arnaud,

Jr., F. R. Barnes, J. W. Bass, L. G. Bezart, R. M. Brown, P. A. Collins, K. Corwin,

H. V. Daly, H. J. Denk, J. G. Edwards, B. Ehreth, F. Ennik, E. R. Gunderson,

T. E. Hewton, Jr., D. Jamieson, W. Jamieson, A. I. Kaplan, D. H. Kavanaugh, A.

Kitayama, H. B. Leech, K. L. Lipps, R. L. Morrison, C. B. Philip, P. Phillips,

J. A. Powell, H. G. Real, E. S. Ross, T. N. Seeno, R. E. Somerby, R. E. Stecker,

S. L. Szerlip, R. W. Thorp, J. W. Tilden, D. W. Whitman, D. D. Wilder, and

D. L. Wilson.

Visitors present (16) : S. Anderson, G. D. Barhe, P. Brown, A. Bush, D. Harms,

P. Jamieson, R. Kimmick, D. L. Parker, J. B. Parker, D. Rodriguez, A. C. Stanet,

G. Sweeny, J. Thorp, H. Tilden, N. Zackeroff, and G. Zefnah.

The minutes of the meeting held 15 November 1974 were summarized.

Richard A. Hinsdale was elected to membership into the Society.

E. S. Ross exhibited kodachrome slides of small insects mounted on microscope

slides.

Allen Stephen displayed an unusual aberration of a checkerspot butterfly captured

in flight in Alum Rock Park, San Jose. The specimen displayed the coloration

The Pan-Pacific Entomologist

of Euphyclryas chalcedona hemimelanica on the dorsal wing surfaces and E. c.

fusimacula on the ventral wing surfaces.

The following notes were presented:

Observations of the Mating Activities of Rhamphomyia Scaurissima

Wheeler in Northern California. — Rhamphomyia scaurissima Wheeler was

found in the oak-bay woodlands of northern Marin County, California. In March

of 1974 a number of these empidids were observed swarming over Santa Fe Creek,

located south of Novato. This mating swarm consisted of two groups; one larger

group of males approximately three feet in diameter, and a smaller group, perhaps

two feet in diameter, composed entirely of females. The two groups were

separated by only an inch or two. The insects in both groups flew in an apparent

circular or elliptical pattern giving the impression that the female group lay an-

terior to the male group. I observed no overlapping of the two groups.

Individual males would frequently fly into the female swarm, select a mate

and fly up under her until they were touching. Together the pair zigzagged in

tandem, in a direction more or less perpendicular to the main flow of the swarm.

Moments later the pair separated, with the female returning to her swarm and

the male immediately flying down to the water and skiing across the water’s

surface. On close inspection, I observed the male touching its modified middle

tarsi to the water, which left a minute ripple on the surface. After this, the male

returned to the swarming group.

Wheeler first described R. scaurissima (Entomol. News, 7:189-192, 1896) from

specimens collected in Palo Alto, California. Since that time, this empidid has

been largely unknown in collections and unmentioned in literature. When Wheeler

first described the extraordinary, elaborate and large middle tarsi of the male,

he thought perhaps their function was to vibrate in front of the female during

courtship. However, this behavior was not observed in any of the mating swarms

I studied. The only way in which the tarsi were used, were as boats or water

skis. In fact the tarsi do resemble a tiller and rudder, with a flattened cup shaped

tarsal segment as the “boat” or “water ski.” The mating swarms have been

found only during the latter part of March or the early part of April. No adult

individuals have been collected at any other time of the year and the larva is

unknown. — Peggie Phillips, San Francisco State University.

Massive Population Levels of the Cherry Tree Ugly-Nest Caterpillar.

— Extensive silk tenting of Prunus virginiana var. demissa caused by gregarious

larvae of the tortricid, Archipa cerasivoranus (Fitch) were observed in southern

Siskiyou County. During California Insect Survey investigations in early June,

1974, large patches of defoliated Prunus were found along the railroad and

roadsides from McCloud to Ash Creek, 9 miles to the east, and there were small

nests at McBride Springs on the west slope of Mt. Shasta in late June. The

largest colony, where all plants were defoliated in an area about 20 X 5-10 m, was

observed on June 10; larvae were dying in large numbers in all but peripheral

areas where some foliage remained. Two weeks later central portions had col-

lapsed and dried, while new webbing on the expanding periphery had encom-

passed various low herbs and other shrubs, particularly Ceanothus velutinus,

which was defoliated, Arctostaphylos, Purshia and Eriogonum. Four tents of

larvae collected 7 and 9 miles east of McCloud and at McBride Springs showed

no signs of disease in laboratory conditions. More than 550 moths emerged be-

tween July 2-29, but only 2 parasites, tachinids, were reared. These olbserva-

Vol. 51, No. 1 , January 1975

tions are in contrast to those made in 1957-58 in the same region, when only

small colonies of A. cerasivoranus were found, and several parasites emerged,

representing about 10% of some 40 individuals reared (Powell, 1964, U. Calif.

Publ. Ent., 32: 147). — J. A. Powell, University of California, Berkeley.

Confirmation of Occurrence in Amazonas of the Tabanid Tribe

Rhinomyzini. — The third known specimen of the peculiar Betraguia ocellata

Oldroyd previously described in 1970 from 2 (likewise) females from Para

Dist., lower Amazon, Brazil is exhibited. It was taken by Dr. Richard W. Merritt

while a pre-doctoral student at University of California and is labelled Leticia

(Upper Amazonas) Columbia, “kilometer 8,” 2-6 August 1973.

The Tribe Rhinomyzini is discussed by Oldroyd in a British Museum review of

the Ethiopian tabanid fauna, 1957, as an assemblage of “bizarre forms” extending

in the Old World as far as the Orient and related more on biological than on

structural or even phylogenetic similarities. While he envisioned tabanid ancestry

as radiating from a South American origin, no previous specialized members of

the above peculiar tribe have been recorded in the New World. There are,

however, other wasp mimics among Neotropical Tabaninae.

Note in the specimen passed among the audience the obvious resemblance

to some shiny black wasp with elongated antennae and abdomen. The latter is

spatulate and truncated behind, not constricted into a slender ovipositor as in

many of its tree-hole breeding ethiopian relatives. Nevertheless, it appears not

unlikely that this Amazon species also inhabits rot-holes in trees.

The well-developed ocelli and the hind-tibial spurs, though minute, relate

this to the tribe, though the combination of sparsely hairy eyes, closed submarginal

cells of the wings, and discal cells narrowed basally by apical extension of the

2nd basal cells, are unusual.

The discovery recently of only 3 widely-separated specimens in all the col-

lecting in the Amazon region attests to some unusual habitats. But logical

speculation on pathways and direction taken by progenitors is not possible at

present. — Cornelius B. Philip, California Academy of Sciences, San Francisco.

The December meeting is traditionally the business meeting of the year and

President Philip asked for reports of the standing committees.

In the treasurer’s report Dr. Arnaud summarized the financial statement of

the Society. Income is derived from dues and subscriptions, reprints and miscel-

laneous page charges, sales of memoirs, advertising, savings accounts interest and

stock dividends. The financial statement is published here in order for all members

to know the financial condition of the Society.

Dr. Arnaud stated that the treasurer’s office is indebted to Mrs. V. Hawley,

Office Manager of the Academy, for her dedicated attention to the Society’s ac-

counts, to Ms. Roxi Berlin, Entomology Secretary, for her attention to many letters

and mailings of Society publications, and to Mr. H. Vannoy Davis, member of

the firm of J. K. Lasser and Company (Certified Public Accountants), not only

for his review of the Treasurer’s records, but also for the completion of our

Federal and State income tax forms.

Mr. Leech gave no report on behalf of the Historical Committee.

Dr. Williams, of the Membership Committee, reported there were 33 members

(21 student and 12 regular) elected in 1974.

On behalf of the Publication Committee, Dr. Powell summarized results of the

The Pan-Pacific Entomologist

Executive Board meetings regarding publication costs of the journal, which will

increase in the future. A small memoir paper was published by M. R. Gardner.

President Philip announced his appointments to the Publication Committee

for 1975 through 1977 : R. W. Thorp and J. A. Powell.

Mr. Leech, Chairman of the Nominating Committee, presented the 1975 slate of

candidates for office in the Society: President, Howell V. Daly; President-elect,

Fred Andrews; Secretary, Franklin Ennik; Treasurer, Paul H. Arnaud, Jr. There

were no nominations from the floor. The candidates were elected to office in

the Society for 1975 by unanimous vote.

The principal speaker of the evening was Dr. Terry L. Erwin, Curator of

Coleoptera, Smithsonian Institute. His illustrated talk was entitled, “The role of

carabid beetles in the arboreal ecosystem in middle America.”

Coffee and refreshments were served in the Trustees Room following the

meeting. — F. Ennik, Secretary.

HONORED MEMBERS OF THE PACIFIC COAST

ENTOMOLOGICAL SOCIETY

C. P. Alexander

E. G. Linsley

A. E. Michelbacher

R. C. Miller

Vol. 51, No. 1 , January 1975

PACIFIC COAST ENTOMOLOGICAL SOCIETY

Statement of Income, Expenditures and

Changes in Fund Balances

Years Ended September 30, 1974 and 1973

1974 1973

Income :

Dues and subscriptions $ 5,516 $ 5,526

Reprints and miscellaneous 7,349 2,999

Sales of memoirs 1,888 3,748

Advertising _ 28 333

Interest on savings accounts 744 601

Dividends, American Telephone & Telegraph Co. 241 224

Increase (decrease) in value of capital stock

of American Telephone & Telegraph Co. (880) 410

14,886 13,841

Expenditures:

Publication costs — Pan Pacific Entomologist 8,013 9,751

Memoirs 92 199

Reprints, postage and miscellaneous 1,977 2,075

10,082 12,025

Increase (decrease) in fund balances 4,064 1,816

Fund balances October 1, 1973 and 1972 22,035 20,219

Fund balances September 30, 1974 and 1973 $26,839 $22,035

Statement of Assets

September 30, 1974 and 1973

1974 1973

Cash in bank:

Commercial account $ 7,908 $ 3,366

Savings accounts:

General fund 8,861 8,426

Memoir fund 2,457 2,189

Fall fund 2,404 2,173

Life membership fund 1,929 1,721

Total cash in bank 23,559 17,875

Investment in 80 shares of American Telephone &

Telegraph Co. common stock (Life Membership

and Fall Funds), at market value 3,280 4,160

Total $26,839 $22,035

See accompanying notes to the financial statements.

The Pan-Pacific Entomologist

NOTES TO THE FINANCIAL STATEMENTS

Year Ended September 30, 1974

Summary of Significant Accounting Policies

Accounting Method

Income, expenditures and assets are recorded on the cash basis of accounting.

Marketable Securities

American Telephone & Telegraph Co. common stock is carried at market value.

Increases and decreases in value are reflected in income.

Income Tax

The Company is exempt from Federal income and California franchise tax.

Accounts Receivable

As of September 30, 1974 accounts receivable aggregated $301 as follows:

August and September, 1974 billings $211

July billings 9

January and February, 1974 (past due) 42

December, 1973 and prior (past due) 39

$301

As Chairman of the Auditing Committee, and in accordance with its bylaws, I

have reviewed the financial records of the Society.

During the course of this review nothing was noted which indicated any inaccuracy

in the foregoing statements.

H. Vannoy Davis

Chairman of the Auditing Committee

THE PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

Papers on the systematic and biological phases of entomology are favored, including

articles up to ten printed pages on insect taxonomy, morphology, behavior, life history,

and distribution. Excess pagination must be approved and may be charged to the author.

Papers are published in approximately the order that they are received. Immediate publi-

cation may be arranged after acceptance by paying publication costs. Papers of less than

a printed page may be published as space is available, in Scientific Notes.

Manuscripts for publication, proof, and all editorial matters should be addressed to the

Editor , John T. Doyen, Division of Entomology & Parasitology, University of California,

Berkeley, California 94720.

Typing.— Two copies of each manuscript must be submitted (original and one xerox

copy are suitable). All parts of manuscripts must be typed on bond paper with

double-spacing and ample margins. Carbon copies or copies on paper larger than 8%

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First page. — The page preceding the text of the manuscript should include (1) the

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Do not abbreviate generic names. Descriptions of taxa should be in telegraphic style.

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alphabetically under Literature Cited in the following format:

Essig, E. 0. 1926. A butterfly migration. Pan-Pac. Entomol.,

2 : 211 - 212 .

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,

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PUBLICATIONS

OF THE

PACIFIC COAST ENTOMOLOGICAL SOCIETY

PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL

SOCIETY.

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Vol. 51

APRIL 1975

No. 2

THE

Pan-Pacific Entomologist

POWELL — Biological records and descriptions of some little known Epiblema

in the Southwestern United States

PARKER — Nest descriptions and associates of three American bees of the

genus Anthocopa Lepeletier 113

FISHER AND ORTH— The genus Limnia in California 123

NESBITT — Description of two species of Diptera, Cyclorrhapha from the

Ethiopian zoogeographical region 130

PINTO — A description of the first instar larva of Eupompha imperialis and

E. edmundsi 134

MOORE — The larva of Cafius sulcicollis LeConte 140

MARSH — -A new species of Apanteles from South America being introduced

into California 143

KURCZEWSKI — Host records for some species of Pompilidae from South-

western United States and Mexico 147

ARNAUD — A new species of Chersodromia from Mexico 152

PETERS — A new species of Indialis from India 159

CHANDLER — A new Acanthinus from Panama 162

SCIENTIFIC NOTES 165, 166, 167, 168, 169, 170

BOOK REVIEWS 172, 173

RECENT LITERATURE 175

OBITUARY — Donald D. Penny 177

ZOOLOGICAL NOMENCLATURE 178

SAN FRANCISCO, CALIFORNIA • 1975

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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The Pan-Pacific Entomologist

Vol. 51 April 1975 No. 2

Biological Records and Descriptions of Some Little Known

Epiblema in the Southwestern United States

(Lepidoptera : Tortricidae)

Jerry A. Powell

University of California, Berkeley 94720

The genus Epiblema Hiibner is widespread in the northern Hemi-

sphere, comprising nearly 100 described species in the Holarctic Region.

The group extends into southern continents evidently by a depauperate

representation, such as in South Africa, although the fauna of southern

regions probably is largely unworked. Primarily on the basis of male

genitalia Obraztsov (1965a) assigned the Palearctic species to three

subgenera, Epiblema s. str., Cacochroea Lederer, and Notocelia Hiibner.

These taxa had been treated at the generic level by some previous au-

thors (e.g., Bradley, 1959). Larvae of the members of the first two of

these subgenera feed in roots or stems of Compositae, sometimes creating

a gall-forming reaction by the host, while larvae of Notocelia are leaf-

rollers on Rosaceae (e.g., Ford, 1949) .

MacKay (1959) placed Nearctic species currently assigned to Epi-

blema and a related genus, Sonia Heinrich, in three groups on the

basis of larval structures and indicated that these are generic level

categories. Her group 1 included E. praesumptiosa Heinrich and Sonia

canadana McDunnough, and MacKay suggested on the basis of Hein-

rich’s (1923) figures of male genitalia that Epiblema abruptana,

numerosana, grossbecki , praesumptiosa , deflexana, insidiosana and ex-

erbatricana should be combined with Sonia to form a single genus. Her

group 2 corresponds to the remainder of Nearctic Epiblema s. str. for

which larvae were available, that is strenuana Walker, otiosana Clemens,

and in essence the species #7024-7032 of McDunnough ’s list (1939),

nearly all of which are known to be stem borers and/or gall makers.

Finally, group 3 of MacKay is equivalent to Notocelia including the

introduced European species suffusana Zeller and the indigenous cul-

minana Walsingham (probably = #7040-7043 of McDunnough).

I have reared the 3 species of Sonia that occur in California, each

The Pan-Pacific Entomologist 51: 99-112. April 1975

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at a number of localities. All are root borers in woody Compositae

( vovana Kearfott in Gutierrezia, filiana Busck and comstocki Clarke

in Haplopappus) . This suggests that the other members of MacKay’s

group 1 are root-borers and that biologies and larvae of this assemblage

(#7015-7023 of McDunnough) have escaped discovery because Amer-

ican microlepidopterists have not much dug into this ecological horizon

yet. This supposition may apply to the other biologically unknown

Epiblema complex (#7033-7039 of McDunnough), as well as most

Eucosma.

Whether larval characteristics ought to be weighted heavily enough

to provide the basis for generic delineation is debatable, but MacKay’s

opinion should encourage a more comprehensive study of Epiblema

and allies. In this group, as in many Microlepidoptera, too much em-

phasis has been placed on male genitalia and secondary male characters

to the exclusion of other lines of evidence. The rainbow that was repre-

sented by the discovery of male genital characters led microlepidopterists

out of the stormy taxonomic chaos of the turn of the century, but its

pot of gold was so doggedly sought that many workers made the mistake

of attempting to force male genitalia to serve a dual role as species

diagnostic characters and as indicators of relationship in development

of generic concepts. As a result some genera in groups such as

Eucosmini are artificial groupings of convenience rather than taxa

that reflect underlying evolutionary relationships. Probably a complete

reshuffle of American Phaneta, Eucosma , Epiblema , Suleima , and Sonia

will be warranted by a thorough investigation.

Several years ago I prepared a report treating an assortment of

Eucosmini, including three previously undescribed species of Epiblema.

However, after review of the manuscript, it seemed advisable to with-

hold the portion dealing with Epiblema until the work by Obraztsov on

subgeneric relationships in Palearctic members of the genus was com-

pleted. Thus the rest of my study was published (Powell, 1963), and

a review of Nearctic Epiblema in concordance with the European fauna

was anticipated. Unfortunately types of the proposed new species were

distributed to the appropriate museums, but the research was not re-

sumed after the Palearctic classification appeared (Obraztsov, 1965a).

The present descriptions are offered to rectify this situation and to

make the names available for ecological work by C. D. MacNeill at the

Oakland Museum and for more comprehensive taxonomic studies by

Richard Brown who has completed a Master’s degree thesis on Epiblema

at the University of Arkansas.

Vol. 51, No. 2, April 1975

101

Epiblema arizonana, new species

(Figs. 9, 10)

A large moth, resembling E. carolinana (Walsingham) , having a mottled brown-

ish forewing with a central whitish area followed by a darker terminal spot.

Male . — Length of forewing 9.1 to 10.2 mm (reared specimens). Head: labial

palpus short, II segment length about 1.0 vertical eye diameter, slightly upcurved,

thickly clothed with rather appressed scaling which forms a blunt, apical tuft

obscuring III segment; latter about 0.33 the length of II, blunt; pale brownish

exteriorly, whitish interiorly. Antenna slightly less than 0.5 forewing length;

orange-brown, dorsal scaling brownish, darker toward base, scape white below.

Scale tufts of crown dense, elongate, strongly directed mesad, brownish reflecting

purplish; scaling of front short, appressed, whitish. Thorax: dorsum dark brown,

the scales faintly tipped with whitish, more strongly so posteriorly; a tinge of

red-brown in upraised tuft of scutellum; metanotum with dense, lateral hair

brushes which do not entirely obscure scutellum; latter scaled brownish. Under-

side whitish, pro- and mesothoracic legs pale brownish exteriorly, excepting white

apical bands of tarsi and tibiae. Forewing: length about 2.7 times width. A

broad, appressed costal fold from base to beyond middle of costa; costal margin

straight to end of fold, slightly convex beyond; termen slightly concave. Ground

color grayish brown, mottled with darker, vertical strigulae. Costal fold of ground

color with about seven darker bands; basal area heavily mottled with dark and

light scales; a broad whitish area over middle one-third of dorsum, becoming

diffuse above middle of wing, enclosing some pale grayish strigulae; a blackish

spot immediately following pale area, just before tornus, with a black spot above

it in apical area; outer costal and apical areas with some red-brown scaling. Fringe

red-brown above middle, gray at tornus. Underside grayish brown, reflecting

purplish; dorsal area white; latter with several rows of upraised, elongate scales.

Hindwing: broader than forewing; costa with slight concavities before and

following middle; termen very slightly concave, broadly curved to dorsum. Upper-

side dark brown, fringe broad, paler, the scales becoming white at apices. Under-

side dark brownish with paler streaks along veins and near margin; anal area

whitish. Abdomen: (of holotype on slide) dorsal scaling dark brownish, under-

side paler, genital tuft shining pale brownish. Genitalia as in fig. 1 (drawn from

holotype, “C. H. slide 1, 10 July, 1936;” two preparations examined) ; socii

elongate, length equal to width of valva at neck (not evident in figure) ; valva

moderately broad, cucullus length about two times its width, cornuti numerous,

elongate, thin.

Female . — Essentially as described for male. Labial palpus apparently somewhat

shorter. Forewing lacking costal fold and broader, length 2.5-2. 6 times width;

whitish areas more extensive and prominent, the markings, particularly the

vertical strigulae, more distinct; tornal dark area as a nearly rectangular blackish

spot crossed by two diagonal brown lines at veins. White areas of underside of

hindwing more extensive. Genitalia as in fig. 5 (drawn from JAP prep. no. 3884,

two preparations examined) ; anterior lobes of papillae anales produced ventrally;

sterigma a simple, weakly sclerotized ring, ductus bursae with an ill-defined

sclerotized patch above junction of ductus seminalis; signa large, smaller one as

large as sterigma, subrectangular, larger one two times the smaller, distally broad

and rounded ; corpus bursae densely spinulate.

Holotype male: Arizona, Oak Creek Canon, near Flagstaff, Coconino

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County, emerged June 28, 1936, reared from Rudbeckia laciniata (G. P. Engel-

hardt) ; allotype female same data except June 26, 1936. Paratypes (18) : 4 $ $ ,

14 2 2 , same data except June 17 to June 30. One male bears a partially illegible

handwritten label which appears to be: “root and stem borer Rudbeckia amplexi-

caulis.” Seven females (June 24, 1936) are worn specimens and lack the host

data and may be flown individuals. Holotype, allotype and 16 paratypes deposited

in the U. S. National Museum of Natural History, two paratypes in the University

of California, Berkeley collection.

There is little variation among the specimens in good condition. The

species is similar to E. carolinana (Walsingham) , which is known to

range west to Illinois. E. arizonana differs only slightly in appearance,

having more restricted whitish areas and more contrasting dark mark-

ings. The male genitalia differ, the valva being considerably broader

in arizonana. In E. carolinana the cucullus distally is slightly narrower

than the neck of the valva (fig. 2), whereas it is broader in the present

species. The foodplant is the same for both, which suggests that they

may be races of a single species ; but until more information is available,

the differences in male genitalia and the allopatry are bases for species

designation.

Superficially, the new species resembles Eucosma williamsi Powell

(1963), and specimens of the two had been mixed in collections.

Epiblema macneilli, new species

(Figs. 11, 12)

A remarkably distinctive, elongate-winged moth, having a sordid white forewing

marked by large blackish blotches.

Male . — Length of forewing 11.3-14.0 mm. Head: labial palpus large, II seg-

ment length about 1.7 times vertical eye diameter, greatly expanded by a broad

scale brush ventrally, completely obscuring the short, appressed scaled, decumbent

III segment; latter about .25 as long as II; scaling dark brown or blackish, re-

flecting purplish, narrowly tipped with whitish, paler basally and interiorly, scales

of the ventral hair brush almost hairlike. Antenna about 0.4 forewing length,

densely setate ventrally, the erect setae subequal to shaft diameter. Head scaling

dense, elongate both on front and crown, the purplish, white tipped scales strongly

directed mesad, exposing the conspicuous, red ocellus. Thorax: dorsal scaling

concolorous with head, tegulae broadly tipped with white; metanotum posteriorly

with short, broad, gray scales. Underside scaling dark gray, tibiae and tarsi

distinctly banded with white. Forewing: elongate, narrow, not greatly broadened

distally; length 2. 9-3.1 times width; fold less than one-third costa length, rather

narrow, costa beyond nearly straight, curved only slightly in outer third, termen

slightly convex, steeply angled back. Ground color whitish, at times tinged with

pale ochreous; markings dark brown or blackish, mostly distinct, as follows:

basal one-fourth of wing including costal fold, enclosing a round costal spot of

ground color, the patch angled outward at its middle; a rectangular spot on costa

from end of fold to middle, extending to top of cell, followed by four triangulate

Vol. 51, Noj 2, April 1975

103

Figs. 1-4, male genital structures of Epiblema, ventral aspect: 1, E. arizonana

Powell, liolotype; 2, valva of E. carolinana (Walsingham) ; 3, E. macneilli Powell,

holotype ; 4, E. rudei Powell, paratype.

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The Pan-Pacific Entomologist

spots along costa, the last at apex joined to four smaller spots along termen by

the dark fringe; a large, rectangular spot on dorsum, before tornus, extending

into lower, outer corner of cell, its sides concave; an indistinct blotch in preapical

area. Underside dark brown, reflecting purplish, paler towards dorsum; whitish

costal interspaces of upperside reproduced. Hindwing: considerably shorter than

forewing; costa concave in outer half; apex broad, termen strongly angled back,

rather sharply angled with dorsum. Variably whitish basally, tending to brownish

apically; fringe brownish basally, whitish dorsally. Underside whitish, the veins

and fringe brownish. Abdomen: scaling blackish gray, genital scaling paler.

Genitalia as in fig. 3 (drawn from holotype, JAP prep. no. 355, three preparations

examined) ; uncus prominent, cucullus very broad, subangulate on distal curvature,

its length about 1.6 times width; cornuti numerous, elongate, thin.

Female. — Length of forewing 10.6 to 11.7 mm. Essentially as described for male

but smaller with more distinct, contrasting forewing markings and darker hindwings.

Forewing slightly broader than in male, length 2.7-2. 8 times width. Maculation

of forewing generally more extensive so that subtornal blotch often joins median

costal spot and nearly reaches subapical blotch, latter connected to apical marks.

Hindwing dark brown. Solerotized portion of VII sternite relatively narrow.

Genitalia as in fig. 6 (drawn from paratype. Mono Pass, JAP prep. 3866, two

preparations examined) ; sclerotization of VIII tergite elongated and enlarged to

form a nearly tubular extension, sterigma broad, rounded; signa small, broad,

the smaller one hardly pointed; surface of corpus bursae granulose.

Holotype male : California, near Mono Pass, 12,000 feet elevation, Inyo

County, August 11, 1958 (C. D. MacNeill), deposited in the California Academy

of Sciences; allotype female, same locality, 12,500', Sept. 3, 1965 (J. Powell),

deposited in California Academy of Sciences on indefinite loan from the Essig

Museum of Entomology, University of California, Berkeley. Paratypes (26) :

near Mono Pass, 12,600', 6 $ $ , 13 2 2, VIII-30, 31-65 (MacNeill, P. A. Opler,

& H. G. Real), 1 $ , same data except VIII-30-67 (Opler) ; same data as allotype,

1 $ , 3 2 2 (Powell & P. A. Rude) ; Mt. Star, north of Mono Pass, 2 2 2, IX-3-65

(A. J. Slater) ; deposited in California Academy of Sciences, U. C. Berkeley and

U. S. National Museum of Natural History.

One additional male in poor condition which appears to be conspe-

cific, although it has a larger preapical blotch on the forewing, has

been examined: “Summit Mt. Whitney, 14,500', Inyo Co., Cal. VIII-

15-53” (no collector given) (LACM).

With its narrow forewings and large blotched color pattern, E.

macneilli is the most distinctive species of North American Epihlema.

In genitalia form the species is most similar to the walsinghami-infelix

complex, which contains more typical appearing members of the sub-

genus, smaller moths with broad, dark forewings having a white dorsal

patch. Disregarding the pulvinus (or “clasper”) the genitalia resemble

certain Eucosma species (e.g., subflavana Walsingham) ; and macneilli

would seem to be a more likely member of that genus, which is a much

more heterogeneous assemblage, although even in American Eucosma

it would not be closely approached in appearance. Superficially the

Vol. 51, No. 2, April 1975

105

female rather closely resembles that of the Asiatic species Eucosma

victoriana (Kennel). Evidently the moths are diurnal, and at the type

locality seemed to be associated with Hulsea algida (Compositae) , but

I was unable to locate any galls or other evidence of larval work on

above ground parts of this plant in September, 1965.

I take pleasure in naming the species for Don MacNeill, who has

collected many interesting Lepidoptera during his numerous visits to

the arctic-alpine type locality.

Epiblema rudei, new species

(Fig. 13)

A pale moth, resembling E. obfuscana and E. desertana in general appearance, the

forewing having a narrow dark terminal band.

Male . — Length of forewing 8.4 to 10.6 mm (reared from field collected pupae).

Head: labial palpus moderately elongate, appressed scaled (without a broad

scale brush obscuring segment III) , segment II length about 1.15-1.20 eye diam-

eter; III .25-.30 as long as II, scaling dark brown, reflecting a purplish sheen.

Head scaling concolorous, with some pale rust colored scaling intermixed. Thorax:

dorsal scaling dark brown, becoming sordid whitish or gray posteriorly. Ventral

scaling shining whitish gray, femora sordid whitish exteriorly, tibiae and tarsi

dark gray, banded with whitish. Forewing: length 2.7-3. 0 times width; narrow

basally with a narrow costal fold extending nearly to mid costa, wing becoming

broader distally, the apex produced; costa nearly straight, termen strongly angled

back. Ground color sordid whitish or pale grayish, marked by two broad, trans-

verse bands of pale bluish gray, the first covering basal one-third of wing, the

second beyond middle, broad at costa where it encloses three pairs of white

costal dashes, becoming narrower and rectangular across wing at end of cell, ex-

tending to dorsum just before tornus, obscured by ground color in costal one-third;

both bands obscurely strigulate with indistinct, transverse pale lines. Subterminal

area narrowly dark gray blending into the dark brown fringe which is tinged

with red-brown. Underside gray, reflecting purplish, costa and subterminal areas

narrowly cream-white; an indistinct whitish area at mid dorsum. Hindwing:

broader than forewing, costal margin convex, apex rounded, terminal margin

slightly convex. Ground color pale gray basally, irregularly streaked with darker

gray-brown outwardly, becoming brownish on outer half of costal area and distal

margins; fringe paler, gray. Underside white except veins on costal half gray.

Abdomen: dorsal scaling dark gray, becoming sordid whitish on final 2 or 3 seg-

ments and laterally; ventral scaling mostly whitish. Genitalia as in fig. 4 (drawn

from paratype, Colton, JAP prep. no. 233, 2 preparations examined) ; uncus 2

prominent triangular peaks, socii moderately elongate, valva broad, cucullus

broadly rounded, its length about 1.75 width, sacculus only weakly emarginate

before cucullus, cornuti apparently lacking.

Female . — Length of forewing 8.0 to 9.0 mm. Essentially as described for male,

smaller with the forewing appearing broader due to absence of the costal fold.

Fringe more richly rust-brown and hindwing slightly darker than in male.

Genitalia as in fig. 7 (drawn from paratype, Colton, JAP prep. no. 2291, two

preparations examined) ; papillae anales flat, unmodified; sclerotization of sterigma

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The Pan-Pacific Entomologist

Figs. 5-7, female genital structures of Epiblema, ventral aspect: 5, E. arizonana

Powell, paratype ( a , papillae anale, lateral outline) ; 6, E. macneilli Powell, para-

type; 7, E. rudei Powell, paratype. Fig. 8, diagrammatic longitudinal section of

gall caused by E. rudei : e = emergence hole; shaded area indicates zone of

packed frass; approximate position of braconid cocoon indicated by dotted line.

Vol. 51, No. 2, April 1975

107

irregularly emarginate posteriorly, signa small but acutely pointed, the smaller one

sometimes very tiny; corpus bursae surface granulose.

Holotype male and allotype female: California, Big Panociie Creek, at

Fresno-San Benito County line, March 4, 1968, reared from stem galls on

Gutierrezia californica, emerged March 17-19, 1968, JAP 68C7 (J. Powell) ; de-

posited in the California Academy of Sciences on indefinite loan from the Essig

Museum of Entomology, U. C. Berkeley. Paratypes (14) : same data, 1 $, 3 2 2;

same locality: 4 S $ , IV-21-67, r.f. Gutierrezia , emgd. IV-24-67, JAP 67D99 (P.

A. Rude) ,2 $ $ , III-6-69, r.f. Gutierrezia, emgd. III-20, 29-69, JAP 69C1

(Powell), 2 $ $, II-5-70, emgd. 11-23, 26-70, JAP 70B1 (Powell); Colton, San

Bernardino Co., Calif., 1 $ , 2 2 2, HI-13, 17-1910 “ex composit gall” (G. R.

Pilate) ; deposited in the California Academy of Sciences, U. S. National Museum

of Natural History, and U. C. Berkeley.

In external appearance Epiblema rudei most closely resembles the

Solidago-feeders, E. desertana (Zeller) and obfuscana (Dyar) of the

eastern U. S., which differ from the present species by having the basal

patch of the forewing darker than the outer transverse band, and by

having more rectangular shaped forewings (termen less steeply angled

back than in rudei). A ratio of base-to-apex: base-to-tornus distances

is 1.30-1.35: 1 in rudei and 1.1-1. 2: 1 in the two eastern species.

Epiblema obfuscana also differs by having the terminal dark band short,

ending above the tornus, which is white and by having dark brown

hindwings. In male genitalia the new species is distinguished from the

above two in many particulars and appears to be most similar to

abbreviatana (Walsingham) , a smaller moth of quite different external

appearance that ranges across the northern U. S.

This new species is named for Paul A. Rude, who first called my

attention to the galls at Panoche Creek, one of many interesting dis-

coveries that he made during fieldwork as an assistant on my N.S.F.

sponsored project on microlepidoptera biologies. Paul had originally

observed the characteristic galls during a study of dipterous galls on

Gutierrezia at La Mesa, San Diego County, several years previously.

Reexamination of that site in October, 1967, revealed that plant suc-

cession was eliminating the Gutierrezia and there was no evidence of

Epiblema. Other places where we have seen galls presumed to be of

Epibletna rudei are: 2 miles north of Lancaster, L.A. County and 2

miles southeast of Pinon Hills, San Bernardino County (immature

galls in October on Gutierrezia microcephala) ; Jacalitos Canyon near

Coalinga, Fresno County; and the hills 2 miles southwest of Kettleman

City, Kings County. A search in the vicinity of Colton to the La Sierra-

Lake Matthews area, Riverside County, in August, 1968, failed to re-

veal work of this moth, as did extensive late season reconnaissance in

other arid regions of southern California, southern Nevada, and Ari-

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The Pan-Pacific Entomologist

Figs. 9-16, adults and gall of Epiblema: 9, 10, E. arizonana Powell, $, 2 para-

types; 11, 12, E. macneilli Powell, $ , 2 paratypes; 13, E. rudei Powell, $ paratype

(JAP 67D99) ; 14, E. rudei, gall and eclosed pupal shell (Colton, Calif.) ; 15, E.

radicana (Walsingham) , 2, San Bruno Mts., Calif.; 16, E. hirsutana (Walsing-

ham) , $ , Berkeley Hills, Calif.

zona during 1966-1968, in connection with a survey of root-boring

Eucosmini inhabiting various woody Compositae.

Gutierrezia is a disclimax situation plant, and in most of the areas

where we have examined it, colonies grow in sites that are overgrazed

or otherwise affected by man’s activities. By contrast, in the Mojave

and Central Valley where we found populations of this moth, the host-

plant thrives on naturally loose sand substrates.

Vol. 51, No. 2, April 1975

109

Biology . — Adults of Epiblema rudei evidently are nocturnal. One

male (70B1) was retained alive for several days and was inactive

during day, moving only at night.

Presumably full grown larvae overwinter, and pupation occurs in

early spring. Although emergences occurred within 2-3 weeks following

collection of galls at various times in spring, the normal flight period

probably is April. One gall dissected in early February, 4 days fol-

lowing collection, contained a pupa, while 1 larva and 2 pupae were

excised in mid-March, 12 days after collection, and both pupae and

already evacuated galls were taken on April 21.

The galls are elongate, thickened areas of 1.5-3 mm thick stems;

usually mature galls are about 20-23 mm in length, with a maximum

diameter of 6.0-6. 5 mm. The silk-lined pupal chamber varies con-

siderably in size from one gall to another, depending primarily on the

thickness of the gall walls, in diameter from 1.8 to 3.3 mm and in

length from 13-17 mm (figs. 8, 14). Emergence occurs via a 2.5-4

mm aperture which the larva prepares just prior to pupation, leaving

a thin layer of hark for the pupa to dislodge. This covering is more

strongly attached on its lower rim so that the pupal shell hangs over it.

After emergence the pupal shell usually readily falls away and the

hatch recloses, leaving the external appearance of abandoned galls

nearly indistinguishable from occupied ones.

At the type locality there was a high rate of parasitism: 65% of a

1969 random sample consisting of 26 old and active galls had either

braconid cocoons or emergence holes of chalcidoids, and 4 of 6 col-

lected in 1970 were parasitized. The braconids mature after the cater-

pillar has spun its silk pupal chamber lining, while the chalcidoids were

of two types. A larger species (not reared) caused the death of the

Epiblema after construction of the emergence ‘window’ but prior to

cocoon spinning, while a pteromalid affected younger larvae, judging

by gall size, and cut its own emergence hole at varying places in the

gall. At the Kettleman Hills, where the galls were abundant in 1974,

a similar high proportion of parasite emergence holes was evident,

while at Jacalitos Canyon galls were rare in February, 1970, and the

two active ones collected both produced parasites.

Epiblema radicana (Walsingham)

(Fig. 15)

Paedisca radicana Walsingham, 1879, Illus. Lep. Het. Brit. Mus., 4:53 (TL:

Rogue River, Ore.).

Epiblema radicana-, Obraztsov, 1965, Amer. Mus. Novitates, 2213:3 (synonymy).

110

The Pan-Pacific Entomologist

Eucosma vomonana Kearfott, 1907, Trans. Amer. Ent. Soc., 33:90 (TL: Cisco,

Calif.).

Eucosma serangias Meyrick, 1912, Ent. Mo. Mag., 48:35 (invalid repl. name).

The complex synonymy and taxonomic confusion involving this spe-

cies and Griselda radicana Heinrich which was occasioned after Heinrich

misinterpreted Walsingham’s species was disentangled by Obraztsov

(1964, 1965b). Fortunately Obraztsov elected to preserve the name

radicana Walsingham rather than declare it a nomen oblitum , and in

the process eliminated one of Kearfott’s nonsense names and one of

Meyrick’s unnecessary substitution names.

We have discovered two colonies of Epiblema radicana in the San

Francisco Bay area, but efforts to locate the larvae have failed. The

moths fly in moderately early spring and appear to be diurnal, which

probably have been major factors in the century-long preservation of

anonymity of this species. Numerous adults were active during midday

in the first week of April, 1962, below Radio Peak in the San Bruno

Mountains, San Mateo County. In subsequent years a few have been

taken there between mid-March and mid-April: IV- 13-66, III-17-68,

III-28-69, and III- 10-72. The site is a rocky ridge featuring a rich

coastal scrub chaparral, with several woody composites that might in-

clude a likely hostplant of Epiblema. The moths seemed to be asso-

ciated with Eriophyllum (probably E. staechadifolium and E. con-

fertiflorum) , although no precise relationship was observed. No gall-like

structures were discovered and several hours digging in early March,

1972, produced only larvae of Hysterosia (Cochylidae) in the crown

and roots. One female of radicana caged in April, 1966, with Eriophyl-

lum foliage lived only 4 days and produced no eggs.

The second colony was located at Raines Park, Del Puerto Canyon,

20 miles west of Patterson, Stanislaus County, where adults (3 $ <3,

1 $ ) were flushed from Ribes quercetorum bushes during late after-

noon on April 28, 1969. This site is in sparse digger pine ( Pinus

sabinana) woods, a quite different habitat than that of the San Bruno

Mountains, and no woody composite was observed in the immediate

area.

Epiblema hirsutana (Walsingham)

(Fig. 16)

Paedisca hirsutana Walsingham, 1879, Ulus. Lep. Het. Brit. Mus., 4:50 (TL:

Sonoma Co., Calif.).

Epiblema hirsutana ; Heinrich, 1923, Bull. U.S. Natl. Mus., 123:149.

This is another species that remained in seclusion for many years

following Walsingham’s original collection in 1871. No specimens were

Vol. 51, No. 2, April 1975

111

known to Heinrich at the time of his 1923 revision. Later the National

Museum received two from Walsingham’s material via the Fernald

collection (Heinrich, 1929), and these remained the only known speci-

mens in American museum for another 30 years.

We rediscovered E. hirsutana when a colony turned up in the Berkeley

Hills in northeast Oakland, Alameda County. A single male was taken

April 3, 1959 (Chemsak and Powell), and males were common there,

flying at midday April 17, 1962. In subsequent years only a few have

been collected: V-16-63, IV-15-64, V-15-64, and V-16-67. This site,

an east facing slope above the Caldecott Tunnel, used to be characterized

by a mixture of native shrubs and herbs including Lupinus arboreus ,

Phacelia calif ornica , Symphoricarpos alba , Wyethia mollis , Artemisia

calif ornica, and A. douglasiana , etc., but plant succession, especially

invasion of alien grasses, encouraged by a fire in 1968, offroad motor-

bike traffic and fire suppression brush clearance, have greatly reduced

the native vegetation. In 1962 E. hirsutana appeared to be flying in

association with Artemisia douglasiana , but subsequent searches have

failed to reveal any gall-like structures on this plant. Collections of

overwintering larvae in the dry standing floral stalks produced only

Pterophoridae and Mordellidae, and examination of roots in March,

1972, yielded only Hysterosia (Cochylidae) . The Artemisia has been

partly eliminated at this locality, and the colony of E. hirsutana also

may be declining, although only one survey during the flight season

has been conducted during the past 7 years.

Acknowledgements

Sincere thanks is made to collectors and authorities of institutional

collections cited above for use of material they made available, and

especially to J. F. G. Clarke of the Smithsonian Institution, who pro-

vided assistance on this project and the drawings of E. arizonana and

carolinana, which had been prepared by a staff artist at the U. S. Na-

tional Museum of Natural History. The rest of the drawings were

done, in part from my pencil drafts, by Celeste Green, artist of the

Department of Entomological Sciences, U. C. Berkeley. Field studies

during 1965-1970, including assistance by P. A. Rude and A. J.

Slater, was funded by N. S. F. grants GB-4014 and GB-6813X.

Literature Cited

Bradley, J. D. 1959. An illustrated list of the British Tortricidae. Part II:

Olethreutinae. Entomol. Gazette, 10(2): 60-80.

Ford, L. T. 1949. A Guide to the Smaller British Lepidoptera. So. Lond.

Entomol. and Nat. Hist. Soc. ; London; 230 pp.

112

The Pan-Pacific Entomologist

Heinrich, C. 1923. Revision of the North American moths of the subfamily

Eucosminae of the family Olethreutidae. U.S. Natl. Mus., Bull. 123;

298 pp.

Heinrich, C. 1929. Notes on some North American moths of the subfamily

Eucosminae. Proc. U.S. Natl. Mus., 75: 1-23.

MacKay, M. R. 1959. Larvae of the North American Olethreutidae. (Lepi-

doptera). Canad. Entomol., 91, Suppl. 10; 338 pp.

McDunnough, J. 1939. Check list of the Lepidoptera of Canada and the United

States of America. Part II Microlepidoptera. Mem. So. Calif. Acad.

Sci., 2(1) ; 171 pp.

Obraztsov, N. S. 1964. Griclelda radicana Heinrich, 1923 (Insecta: Lepi-

doptera) : Proposed validation under the plenary powers. [Z.N. (S).

1612]. Bull. Zool. Nomencl., 21: 144-145.

Obraztsov, N. S. 1965a. Die Gattungen der Palaearktischen Tortricidae II. Die

Unterfamilie Qlethreutinae. 6 Teil. Tijd. voor Entomol., 108: 365-387.

Obraztsov, N. S. 1965b. On Paedisca radicana and the type species of the

genus Griselda (Lepidoptera, Tortricidae, Qlethreutinae). Amer. Mus.

Novitates, 2213: 1-16.

Powell, J. A. 1963. Records and descriptions of some interesting species of

Eucosma in California (Lepidoptera: Tortricidae). Proc. Biol. Soc.

Washington, 76: 235-246.

Nest Descriptions and Associates of Three American

Bees of the Genus 44 Anthocopa ” Lepeletier

(Hymenoptera: Megachilidae)

Frank D. Parker

Bee Biology and Systematics Laboratory, ARS, USDA

Utah State University, Logan, Utah 84322

“ Anthocopa ’ Lepeletier is a large group of megachilid bees found

in North America, Eurasia, and Africa. This group has been treated

as a genus in North America, but European authors have included

Anthocopa in Hoplitis Klug. Recently Michener (1968) suggested that

Anthocopa be synonymized with Hoplitis. For the purposes of this

paper, I an using Anthocopa because of its familiarity and its current

usage in the catalog of Hymenoptera of America (Muesebeck et al.

1951).

The North American species of Anthocopa are grouped into 4 sub-

genera containing 34 species. The nesting habits of the Nearctic species

are undescribed, but Parker and Bohart (1966, 1968) recorded 2

species from trap stems. This paper describes nests of 3 species in-

cluding the 2 from trap stems. Although these reared species belong to

different subgenera and all have dissimilar nesting habits, I feel it is

premature to propose that these behavior patterns are indicative of

subgeneric categories because so few species are known biologically.

Anthocopa (Eremosmia) hypostomalis Michener

(Figs. 1-6)

Nesting Site : All nests (98 containing 305 cells) were recovered

from prebored elderberry trap stems each with a bore diameter of

6-3 mm (%-% in.) ; only the end holes were utilized though side holes

with similar diameters were present on the same stems. The trap stems

selected by these bees were placed in the following localities in southern

California following the procedures of Parker and Rohart (1966) :

Deep Canyon and White Water Canyon, Riverside Co.; Kramer Hills,

San Bernardino Co.; Palm Canyon (Anza-Borego State Park), San

Diego Co.; and Glamis, Imperial Co. The best trapping site was White

Water Canyon where 92% of the nests studied were recovered. In

1964, this species occupied 28% of the recovered trap stems; the next

season it utilized 53%.

Nest Construction : Nests were made by building complete cells one

on top of the other (Fig. 1) ; nests averaged 3.1 cells with a range of

The Pan-Pacific Entomologist 51 : 113 - 122 . April 1975

114

The Pan-Pacific Entomologist

1—9. Each cylindrical cell was made by combining gravel and masti-

cated plant parts and lining the burrow walls with this material. In

smaller holes, the cells were long and narrow (5 X 13mm) and placed

vertically. In larger borings, the cells were stout (3 X 9mm), and the

cell series was oblique. Pith was removed from the sides of the burrow

to accommodate the oblique cells. Also, because of the softness of the

pith in the stems, the drilled holes were occasionally larger than the

size stated earlier. The inner cell walls were smooth, polished, and

composed of finely masticated plant parts combined with a salivary

substance. The outer cell surface was roughened by protruding parti-

cles of sand and pieces of plant material (Figs. 2, 3). The cell cap was

composed of a thin disc of masticated plant parts with a layer of sand

appressed to the outer surface. The cap was concave and smooth

initially; but later the concave area was filled with small pieces of

gravel stuck together with masticated plant parts. The inside of the

cell cap was rough and without any apparent design or openings. The

nest entrance and burrow above the cells were plugged with fine gravel.

The gravel in the entrance was held together with a salivary substance.

One stem was plugged with a thin disc of resin, but this plug may have

been applied by another species of bee.

Provisions : The size and shape of the pollen mass was not recorded.

Hurd and Michener (1955) listed Dalea and Cryptantha as host plants

for A. hypostomalis. Pollen found in the reared cells was Leguminosae,

probably Dalea ; but one sample contained pollen from a hydrophyl-

laceous flower.

Feces: The walls of the cells were lined with feces; those at the top

were loose, but those on the sides were flattened against the cell walls.

The dark, loose fecal particles were fairly uniform in length (0.5mm) ,

tapered, and oblong; there was a faint linear depression on one surface.

Cocoons: The cocoons were formed by lining the cell walls from the

bottom to nearly the top with finely spun silk. In shorter cells, the

cocoon filled the cavity; in longer cells, it filled the lower % of the cell

(Fig. 4). The cocoon was thick, dark brown, and oval and had a nipple

at the top (Fig. 4). The outer lateral surface of the cocoon below the

nipple usually possessed several rings of loosely spun silk that in some

cells extended to the cell walls. The nipple was loosely spun and of

various shapes; but the inner cocoon surface opposite the nipple

always had many strands of loose silk. The cocoon was tightly ap-

pressed to the cell walls.

Sex Ratio: More females than males emerged, and the ratio females

to males was 1.2: 1.0. The ratios of females to males in each cell

Vol. 51, No. 2, April 1975

115

Fig. 1. Two cells of A. hypostomalis in a trap stem, bottom cell with larva.

Fig. 2. Cell of A. hypostomalis showing material used to construct the cell.

The nest entrance plug is attached to the top of the cell.

Fig. 3. Female and cell of A. hypostomalis.

Fig. 4. Cell B in fig. 1 opened to illustrate the white nipple at the top of the

cocoon. Note rings of silk at the top.

116

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beginning with the first (bottom) cell was 1.6, 1.3, 1.0, 1.0, 0.5, 1.3,

1.0, 0.4, 0.0. Thus, the placement of the sexes in the cells followed the

usual pattern of females at the bottom and males above; but in 7% of

the nests, the sexes were intermixed, and in another 10% only males

were present.

Nest Associates : The four insect species associated with nests of A.

hypostomalis accounted for a 15% cell loss. An undescribed species of

the parasitic bee, Stelis (Megachilidae) (Fig. 5), was the most common

parasite, and it accounted for 62% of all parasitism. Like its host,

more females than males (ratio 1.5:1) of this parasite emerged.

Cocoons of Stelis also possess nipples (more elongate than the Antho-

copa ) , but these cocoons are smaller and lack the rings of silk at the

top (Fig. 6). A clerid beetle, Cymatodera sp., was the second most

abundant nest associate. Larvae of these beetles destroyed 28% of the

depredated cells, and as many as 3 cells in a series were entered and

destroyed by one larva. Leucospis affinis Say, a common chalcid

parasite of bees, was found in 2 cells, and a male mutillid, Sphaerop-

thalma amphion amphion (Fox), was reared from one cell. Two nests

were destroyed by an unknown species of woodpecker, and cell losses

due to unknown causes totalled 23.2% with 13.7% occurring in the pre-

cocoon stages and 9.5% in later stages.

Anthocopa (Hexosmia) copelandica (Cockerell)

(Figs. 7-9)

Two of the 3 recognized subspecies of A. copelandica were

reared. The observations recorded below apply to both subspecies

except where differences are noted.

Nesting Site : Nests of this species were collected from prebored

elderberry stems set out at 6 western localities. At 3 locations, Boca

and Camelian Bay (Nevada Co.), California, and Craters of the Moon

National Park (Butte Co.), Idaho, 32 nests of the subspecies c.

albomarginata (Cockerell) were found. At 3 southern Californian

localities, Brown Canyon, Kramer Hills (San Bernardino Co.), and

Deep Canyon (Riverside Co.), 20 nests of the subspecies c. arefacta

(Cockerell) were found. Three sizes, 6, 3, and 1.5mm (%, Vs, Vi6 in.)

of end and side holes were utilized by these bees.

Nest Construction : The species was versatile in nest building. In

wide burrows (6mm), cells were grouped with cell walls fashioned

mostly from nest building material (Fig. 7) ; in narrow holes (2-4mm) ,

only thin partitions (l-2mm) separated the cells. Cell size ranged

from 4-9mm long and 3-4mm wide. The placement of cells in burrows

Vol. 51, No. 2, April 1975

117

Fig. 5. A male of Stelis sp., a parasite of A. hypostomalis.

Fig. 6. Cocoon of Stelis sp. in cell of A. hypostomalis.

was highly variable, but there was often an empty space (avg length

28mm) between the last cell and the entrance plug, usually enough

space for more cells. Nest building material was coarsely chewed

plant parts; but in most nests of c. arefacta , sand and masticated plants

were used in cell partitions and entrance plugs. Cell partitions were

formed into thin discs, and the edge of some partitions were bent down-

ward. In larger burrows, some of the upper partitions were partially

constructed (except for the small entrance hole) prior to cell provision-

ing. The cell partitions were smooth above and roughened below.

Nests were capped with discs of chewed plant parts, but occasionally

some pith from the sides of the stem was incorporated into the plugs,

which averaged 2-9mm long. Some plugs were composed of as many

as 8 compact discs, and they were often situated below the nest entrance

(avg 4.5mm). Thirty -two nests of the subspecies albomarginata were

examined; they held 79 cells (1-10, avg 2.4/nest). Twenty nests of the

subspecies arefacta were recovered; they held 87 cells (1-11, 4.3 /nest).

Provisions: Shape and size of pollen masses were not recorded.

Pollen samples in cells from Idaho were all Hydrophyllaceae.

Feces: The feces were light tan in cells of c. arefacta (Fig. 8) and

dark brown in cells of c. albomarginata (Fig. 9). Neither had surface

113

The Pan-Pacific Entomologist

y '

Vol. 51, No. 2, April 1975

119

depressions, and some particles were tapered and others were blunt.

Most particles were deposited at the top of the cell or around the walls,

but often they were flattened against the sides.

Cocoon : The cell wall was lined by thick layers of transparent silk

that fit snugly against the wall. Inside this layer but closely attached

to it a dark amber cocoon was spun. The amber cocoon was often

smaller than the outer one in longer cells ( % ) , but in shorter cells it

filled the transparent cocoon. The cocoon did not have a protruding

nipple, but at the top it was slightly raised with many strands of loose

silk. The inner surface of the cocoon was shiny, but the silk strands

were evident.

Sex Ratio : More females than males emerged, 2.4:1, and only 2 nests

contained series with the sexes mixed. In all other nests, the females

were at the bottom, and the male cells were above.

Nest Associates'. Five species of parasites and predators were re-

covered from A. copelandica cells. Total parasitism was low, only 6.6%.

The most common parasite was the chrysidid, Chrysura sonorensis

Cameron, which occupied 6 cells of c. albomarginata ; another wasp,

Sapyga pumila Cresson, was found in 2 cells of this subspecies. The

other 3 parasitized cells each contained one of the following species:

a male leucospid wasp, Leucospis af finis Say, from a c. arefacta cell;

a pteromalid, Epistenia sp., and a meloid beetle, Neniognatha scutel-

laris LeConte, each from a c. albomarginata cell. Three cells in one

nest of c. arefacta were destroyed by a clerid larva belonging to the

genus Cymatodera. Cell mortality due to unknown causes was 4.8%.

Anthocopa (Atoposmia) abjecta (Cresson)

(Figs. 10-13)

Nesting Site : Nests of A. abjecta were found attached to the under

surface of flat stones at 2 locations above 8,000 ft in Cache County,

Fig. 7. Nest of A. c. albomarginata showing composite cells. Some cells were

almost completely lined with macerated plant parts.

Fig. 8. Cocoon of A. c. albomarginata from a wide burrow. The shape of the

cocoon followed the walls of this irregular cell. Pith from the trap stems was

incorporated into the plug at the top of the cell.

Fig. 9. Female and cell of A. c. arefacta.

Fig. 10. Leaf part cells of A. abjecta.

Fig. 11. Leaf part cells of A. abjecta, ventral view.

Fig. 12. Female and cell of A. abjecta.

Fig. 13. Top of cocoon of A. abjecta, showing the coarse silk strands.

Table L Comparative biology of Anthocopa species.

120

The Pan-Pacific Entomologist

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Vol. 51, No. 2, April 1975

121

Utah. One habitat was near the summit of Wellsville Mountain, and

the other was along Beaver Creek near the Idaho-Utah state line.

Nest Construction: Cell walls were constructed entirely from dark

masticated plant material and shaped into domes about 9mm long and

8mm wide (Figs. 10, 11, 12). The rock surface was used as the base.

Cells were capped with the same nest building material that was shaped

into discs — smooth on the outside and roughened on the inside. The

inside of the cell walls was slightly smoothed. When cells were adjacent,

only one wall sometimes separated the cells. The number of cells found

was not recorded because many were old and constructed during pre-

vious seasons. However, one nest contained 16 cells.

Provisions: Hurd and Michener (1955) listed species of Penstenion

as the host plant for A. abjecta. Seven of the cells from which A. abjecta

emerged contained traces of Penstenion pollen. One cell was partially

filled with a white vile smelling liquid.

Feces: In most cells, feces were flattened against the walls, but in

some they were at either end. The links were dark brown, of uniform

length, and without surface markings.

Cocoon: The coarse matlike dark brown strands (Fig. 13) that

formed the cocoon closely lined the cell walls. The strands were loosely

spun on the outside, but inside the cocoon was polished with many

strands visible. The top of the cocoon did not have a nipple.

Sex Ratio: Only females emerged from the cells examined. I did

not find any parasitized cells at the Beaver Creek locality, but at

Wellsville Mountain where only old cells were found, many had been

parasitized by an unknown species of Stelis.

Discussion

Nesting habits of Anthocopa species are indicative of the close rela-

tionship between these bees and other genera such as Hoplitis and

Osmia. Some of the more important nesting patterns among Anthocopa

and related North American genera are: (1) cells made entirely of

masticated plant parts [A. abjecta , Hoplitis biscutellae (Cockerell)],

(2) cells made primarily with fine gravel (A. hypostomalis, Hoplitis

anthocopoides Schenck), (3) cell partition and plug of masticated plant

parts (A. copelandica , Osmia kincaidii Cockerell), (4) cocoon with

nipple (A. hypostomalis , most spp. of Osmia), (5) cocoon without

nipple (most spp. of Osmiini except Osmia), (6) cocoons composed

of 2 distinct layers (A. copelandica, Osmia kincaidii Cockerell), (7)

nests in pre-existing holes (A. hypostomalis, A. copelandica, most spe-

cies of Ashmeadiella, Hoplitis, Heriades, Chelostoma, Proteriades, and

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The Pan-Pacific Entomologist

some Osmia) , and (8) nests under stones [A. abjecta , Osmia spp.

( tanneri Sandhouse, longula Cresson, Integra Cresson)].

Table 1 summarizes the biology of the Anthocopa species considered

in this paper.

Acknowledgments

Thanks are due the persons who helped identify material reared

from the trap stems. C. D. Michener (UK) determined the Apoidea,

B. D. Burks (Systematic Entomology, ARS, USDA) Chalcidoidea, D. S.

Horning (UCD) Chrysididae, and W. Enns (UM) Meloidae. H. Potter

(ARS, Logan) made the pollen identifications. Appreciation is ex-

tended to those persons who reviewed this manuscript — Drs. K. V.

Krombein (USNM), G. C. Eickwort (Cornell), and R. W. Thorp

(UCD).

Literature Cited

Hurd, P. D., Jr. and C. D. Michener. 1955. The Megachiline bees of California.

Bull. Calif. Insect Survey 3: 1-247.

Muesebeck, C. F. W., et al. 1951. Hymenoptera of America North of Mexico,

Synoptic Catalog, USDA Monogr. 2, 1420 p.

Michener, C. D. 1968. Nests of some African megachilid bees, with descrip-

tions of a new Hoplitis. Entomol. Soc. South Africa 31 : 337-359.

Parker, F. D. and R. M. Bohart. 1966. Host-parasite associations in some

twig-nesting Hymenoptera from western North America. Pan-Pac.

Entomol. 42: 91-98.

Parker, F. D. and R. M. Bohart. 1968. Host-parasite associations in some twig-

nesting Hymenoptera from western North America. Part II. Pan-Pac.

Entomol. 44: 1-6.

The Genus Limnia in California

(Diptera: Sciomyzidae)

T. W. Fisher and R. E. Orth

Department of Entomology, University of California, Riverside 92502

The genus Limnia Robineau-Desvoidy (1830: 684) can be distin-

guished from other sciomyzid genera by the following characters:

propleural bristle lacking; vallar (subalar) bristles present; arista with

whitish hairs or pubescence; mid-frontal stripe approximately % width

of frons; wings reticulated; black spot on medifacies lacking; tp (pos-

terior crossvein) arcuate, only slightly sinuate.

As indicated by Steyskal (1965: 691), Limnia is in need of revision.

Such a study is in progress jointly with G. C. Steyskal, L. V. Knutson

(USNM), and the authors. Pursuant to compilation of a California

Insect Survey Bulletin on the Sciomyzidae it has become necessary to

clarify the status of the genus in this State. We recognize four species

in California; L. severa Cresson (Cresson, 1920: 80), L. inopa

(Adams) (Adams, 1904: 448), L. pubescens Day (Day, 1881: 86),

and L. boscii (Robineau-Desvoidy) (R.-D., 1830: 690). The first three

named are western North American in distribution. Limnia boscii is

transcontinental between 35°-47°N Latitude. Both sexes of the species

collected within California are easily separated by external characters.

However, outside of California, examination of male terminalia is

necessary for definitive determination of certain species. Structures

such as the hypandrium, surstylus, and aedeagus are diagnostic for

L. inopa and L. severa (Figs. 6-16).

The objectives of this paper are to (1) resurrect and redescribe

L. inopa (Adams) ; (2) redescribe L. severa Cresson; (3) provide a

key to the Limnia of California; and (4) summarize biological infor-

mation for Limnia in California.

Limnia inopa (Adams)

Tetanocera inopa Adams, 1904: 448. Holotype $ ; Washington Territory;

Repository, University of Kansas.

Limnia costalis var. brevicostalis Melander 1920: 323.

Limnia brevicostalis Melander, Steyskal 1965: 691.

Limnia boscii (Robineau-Desvoidy) 1830: 690, Steyskal 1965: 691.

Examination of the types of L. inopa (Adams) (Adams, 1904: 448) and L.

brevicostalis Melander (Melander, 1920: 323) plus a series of 75 specimens in

The Pan-Pacific Entomologist 51: 123-129. April 1975

124

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the UCR collection and 33 specimens from the collections of CAS, CU, CD A,

KSU, KU, UCB, and USNM lead us to conclude a single taxon is involved.

Distribution: Alta., B.C., Calif., Idaho, Mont., Oreg., Wash.

Head. — Frons yellow, longer than wide, mid-frontal stripe % to % width of frons,

usually widest in females; two frontal-orbital bristles present, anterior socket sur-

rounded by velvety black coloration; ocellars and post-ocellars prominent and

of nearly equal length, longer than fronto-orbitals; velvety black patch present

in anterior corners of frons. Single velvety black patch present on occiput im-

mediately posterad of post-ocellars. Face and gulae whitish; medifacies bare;

parafacies with fine black hairs; central portion of medifacies sometimes with

brown discoloration; face deeply concave. Palpi yellow. Eyes large, oval, dis-

tinctly longer than high. Antennae with arista with dense white short hairs, basal

segments yellow and hirsute; third antennal segment yellowish, usually tinged

with black anteriorly, hirsute; second antennal segment approximately same

length as third, longer than high from lateral aspect, inflated ventrally, com-

pressed dorsally in cross-section, highly polished, ochraceus yellow, usually with

brownish area on dorsal half, usually with two strong bristles on dorso-apical half.

Thorax with notum with two narrow brown vittae centrally, a pair of broader

brown vittae laterally and a pair of smaller vittae appearing posterad, all separated

by dull pruinosity; scutellar bristles prominent; prescutellar bristles variable,

generally vestigial to half length of scutellum. Pleura pruinose; pro- and meso-

pleura brown in upper half. Prostemum without hairs. Legs yellow, tarsal segments

3, 4, and 5 (and sometimes 2) may be dark brown. Wings hyaline, with brown

reticulations; posterior crossvein curved outwardly, may be slightly sinuate; halter

pale yellow. Wing lengths of 20 females 4.3 to 6.0 ,mm (average 5.2 mm), 20

males 3.3 to 5.3 (average 4.6 mm) .

Abdomen pruinose with irregularly defined brown vittae laterally. Post-abdomen

as in Fig. 3.

Limnia severa Cresson

Limnia severa Cresson, 1920: 80. Holotype $ ; Cayton, Shasta County, Cali-

fornia; Repository, California Academy of Science.

Limnia unguicornis var. severa Cresson 1920: 80.

Limnia saratogensis var. severa , Melander 1920: 324.

Limnia saratogensis var. armipes Melander 1920: 324.

Figs.- 1-5. Fig. 1 . Limnia pubescens (Day). Head, male. USA, Calif., Shasta

Co., Cayton; Elev. 3,050 ft.; VII-25-1972; T. W. Fisher and R. E. Orth, collectors.

Arrow indicates strong diagnostic bristles on the anterior dorsal edge on the

second antennal segment. Figs. 2-5. Postabdomen, sinistral view; ae, aedeagus;

ep, epandrium; hy, hypandrium; ss, surstylus; VI, ventral projection of 6th

sternite or protandrium. Fig. 2. Limnia boscii (R.-D.). USA, Calif., Mono Co.,

Fish Slough; Elev. 4,250 ft.; IX-7-1967; T. W. Fisher and R. E. Orth. Fig. 3.

Limnia inopa (Adams). USA, Calif., Modoc Co., East of Cedar Pass; Elev. 5,800

Vol. 51, No. 2, April 1975

125

ft.; VIII-24-1967; T. W. Fisher and R. E. Orth. Fig. 4. Limnia severa Cresson. USA,

Calif., Alpine Co., south of Woodfords; Elev. 6,000 ft.; VII-11-1968; T. W. Fisher

and R. E. Orth. Fig. 5. Limnia pubescens (Day). USA, Calif., Shasta Co., Cayton;

Elev. 3,050 ft.; VII-25-1972; T. W. Fisher and R. E. Orth.

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Limnia armipes, Steyskal 1965: 691.

Limnia severa, Steyskal 1965: 692; Fisher & Orth 1971: 164.

Distribution: B.C., Ariz., Calif., Colo., Oreg., Wash.

Head. — Frons yellow to testaceous, length and width nearly equal, mid-frontal

stripe approximately % width of frons; two fronto-orbital bristles present, anterior

socket surrounded by velvety black coloration, posterior socket to a lesser degree;

ocellars and post-ocellars prominent and of nearly equal length, longer than

fronto-orbitals; velvety black patch present in anterior corners of frons; single

velvety black patch present on occiput immediately posterad of post-ocellars.

Face and gulae whitish; medifacies bare; parafacies with fine back hairs; central

portion of medifacies sometimes with yellow to brownish discoloration; face mod-

erately concave. Palpi yellow. Eyes large, oval, distinctly longer than high. An-

tennae with arista with moderately short and dense hairs, basal segments yellow

and hirsute; third antennal segment yellowish, tinged with black anteriorly,

hirsute; second antennal segment approximately same length as third; longer

than high from lateral aspect; polished, ochraceus yellow, usually with brownish

area on dorsal % to %; usually with two strong bristles on dorso-apical half.

Thorax with notum with two narrow brown vittae centrally, a pair of broader

brown vittae laterad, and a pair of additional smaller vittae appearing posterad,

all separated by dull pruinosity; scutellar bristles prominent; prescutellar bristles

strong, as long as scutellum. Pleura pruinose; pro- and mesopleura brown in

upper % to %. Prosternum usually without hairs. Legs yellow to testaceous,

front tibae and tarsi brownish to black, mid and hind tibae and tarsi less darkly

pigmented, usually only tarsal segments 4 and 5 blackish. Wings hyaline, with

dark brown reticulations; posterior crossvein curved outwardly, usually sinuate;

halter yellow, brownish at apex. Wing lengths of 20 females 4.6 to 6.0 mm

(average 5.4 mm), 20 males 4.0 to 5.4 mm (average 4.9 mm).

Abdomen pruinose with two lateral brown vittae on each side. Postabdomen as

in Fig. 4.

Key to the Species of California Limnia

1. Brown marginal cell of wing with clear spots extending to costal margin;

prosternum with hairs; frons wider than long; second antennal segment

with 2 strong bristles on the anterior dorsal edge; prescutellar bristles ap-

proximately as long as scutellum. (Male terminalia as in Fig. 2)

boscii (Rob.-Desv.)

Brown marginal cell of wing without clear spots extending to costal margin;

prosternum without hairs 2

2. Second antennal segment with 4 or more strong bristles on the anterior

dorsal edge (Fig. 1) frons approximately as wide as long; prescutellar

Figs. 6-16. Figs 6-11. Limnia inopa (Adams). Diagnostic structures of post-

abdomen. USA, Calif.; Modoc Co.; Cedar Pass Campground; Elev. 5,800 ft.;

VIII-8-1968; T. W. Fisher and R. E. Orth, collectors. Legend: aea, aedeagus;

Vol. 51, No. 2, April 1975

127

ap, anterior process; eja, ejaculatory apodeme; vph, ventral process of hypandrium.

Fig. 6. Hypandrium, sinistral view. Fig. 7. Hypandrium. Fig. 8. Surstylus (left) ,

sinistral view. Fig. 9. Aedeagus, sinistral view. Fig. 10. Aedeagus, posterior view.

Fig. 11. Aedeagus, anterior view. Figs. 12-16. Limnia severa Cresson. Diagnostic

structures of post abdomen. USA, Calif.; Inyo Co.; Lake Sabrina; Elev. 9,000 ft.;

IX-6-1967 ; T. W. Fisher and R. E. Orth, collectors. Fig. 12. Hypandrium, sinistral

view. Fig. 13. Surstylus (left) , sinistral view. Fig. 14. Aedeagus, sinistral view.

Fig. 15. Aedeagus (looking posterad) . Fig. 16. Aedeagus (looking anterad) .

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The Pan-Pacific Entomologist

bristles approximately % to % as long as scutellum. (Male terminalia as

in Fig. 5) pubescens (Day)

Second antennal segment with 2 strong bristles on the anterior dorsal edge.

3. Frons approximately as wide as long; pre-scutellar bristles as long as scutel-

lum. (Male terminalia as in Fig. 4) severa Cresson

Frons distinctly longer than wide; prescutellar bristles variable, generally

vestigial to % length of scutellum (Male terminalia as in Fig. 3)

inopa (Adams)

Discussion

Of the 440 Limnia specimens we collected within the State between

VI-10-1965 and VI-26-1974, approximately 80% were L. severa , 11%

pubescens , 8% inopa , and less than 1% boscii. In the 76 collections

made in California which contained Limnia, L. severa occurred alone

at 29 sites, L. inopa alone at 6, L. pubescens alone at 3, and L. boscii

alone at one. Limnia severa and L. inopa co-habited 3 sites ; severa and

pubescens co-habited one site. Three species, inopa, pubescens, severa,

co-habited two sites, Bartle in Siskiyou Co., and Cayton, in Shasta

Co. In 1973-1974 at Bartle, a large diverse habitat, L. severa, L.

pubescens, and L. inopa represented 9%, 4%, and 4%, respectively, of

the 13 sciomyzid species collected. In 1972, 1973, and 1974 at Cayton,

a very restricted habitat, L. pubescens, severa, and inopa represented

55%, 19%, and 1%, respectively, of the 10 sciomyzid species collected.

Limnia boscii was the only species of Limnia we collected at Fish

Slough in Mono Co., 10 miles north of Bishop, and our records indicate

that it occurs only at that locality within California.

Over the nine year period of collecting in California we found Limnia

co-existing with a rather diverse sciomyzid fauna, i.e., 21 species.

Allowing for repeated collections at certain of the sites this summary

does not reflect seasonal population trends but was derived from 76

isolated points in time. On-site separative mechanisms such as time,

space, plant associations, or proximity to water, are discernible. Two

examples are; (1) L. inopa does not appear before mid-summer, and

(2) repeated visits to the Hat Creek and Cayton sites yielded L.

pubescens only at restricted areas of several square meters within the

larger boundaries of those sites. This observation suggests that L.

pubescens requires a rather special set of ecological parameters to exist

at all, but when those requirements are met pubescens can exist in rela-

tively large numbers.

Acknowledgments

Our thanks to G. W. Byers, (Snow Entomological Museum, Univer-

sity of Kansas) who provided the type of L. inopa Adams; to P. H.

Vol. 51, No. 2, April 1975

129

Arnaud, Jr., (California Academy of Sciences) who provided the type

of L. severa Cresson; and to L. V. Knutson (U. S. National Museum of

Natural History) who provided the type of L. costalis brevicostalis

Melander. Review of the manuscript by L. V. Knutson and G. C.

Steyskal is greatly appreciated. The work was partly supported by

U. C. Agric. Expt. Sta. Project 2037.

Literature Cited

Adams, C. F. 1904. Notes on and descriptions of North American Diptera.

Kans. Univ. Sci. Bui., 2: 433-455.

Cresson, E. T., Jr. 1920. A revision of the nearctic Sciomyzidae (Diptera,

Acalyptratae) . Trans. Amer. Entomol. Soc., 46: 27-89.

Day, L. T. 1881. Notes on Sciomyzidae, with descriptions of new species.

Canad. Entomol., 13: 85-89.

Fisher, T. W. and R. E. Orth. 1971. Limnia armipes Melander synonymized

with Limnia severa Cresson (Diptera: Sciomyzidae). Pan-Pac. Entomol.,

47: 164.

Melander, A. L. 1920. Review of the nearctic Tetanoceridae. Ann. Entomol.

Soc. Amer., 13: 305-332.

Robineau-Desvoidy, J. B. 1830. Essai sur les Myodaires. Inst, de France, Sci.

Math, et Phys., Acad. Roy. des Sci., Mem. presentes par divers Savans

(Paris) [ ser. 2], 2: 1-813.

Steyskal, G. C. 1965. Family Sciomyzidae (Tetanoceridae), pp. 685-695. IN:

Stone, A., et al., A catalog of the Diptera of America north of Mexico.

U.S. Dept. Agric., A.R.S., Agric. Handbook No. 276, pp. 1-1696.

Description of Two New Species of Diptera, Cyclorrhapha

from the Ethiopian Zoogeographical Region

Elizabeth Nesbitt

Department of Entomology, South African Institute for Medical Research

Johannesburg, South Africa

From Dr. P. H. Arnaud, Jr., California Academy of Sciences, San

Francisco, the Department of Entomology received a large number of

Calliphoridae and Sarcophagidae. These flies had been collected re-

cently in various parts of Africa and proved very interesting. Two new

species are described here and more will be described in the future.

Phumosia rossi new species (Calliphoridae : Calliphorinae)

In the key to the Phumosia species of the Ethiopian zoogeographical

region (Zumpt, 1956) this species runs down to the P. stabulans- group.

It is well characterized by the male terminalia which show long inwardly

curved paralobi terminating in a distinct knob (fig. 1).

Male — Frons at its narrowest point )4oAi2 as wide as the eye is long. Par-

afacialia and -frontalia black or dark brown with silvery pollinosity, 5 to 8

irregularly arranged cruciate paf; iv and oc strong. First and second antennal

segments brown, third segment darker, twice as long as second. Arista with long

hairs. Bucca about % as high as eye, black with greyish pollinosity, hairs black.

Palpi yellow-brown, almost parallel, slightly curved.

Thorax black with dense olive pollinosity forming three longitudinal ill-defined

and partly indistinct bands on the mesonotum. Chaetotaxy: ac = 0-1 -f- 1-2,

dc — 2 -f- 3, ia — 0-1 -f- 2-3, ph - 1 (outer absent) , h — 2, prs = 1, n — 2,

sa~ 2-3, pa — 2, sc = 2-3 + 0-1, st = 2:1. Pro- and poststigma red-brown to

dark brown, propleuron and prosternum haired, alar- declivity bare. Wings hyaline

or partly tinged especially towards the base. Veins yellow-brown, costal spine

long, a few setae on the upper side of ri l + 5 ' r-m clouded, R s open. Upper squama

brown, lower squama yellowish. Legs with femora dark brown, tibiae and tarsi

yellow-brown. Fore-tibia with several short ad and 1 long submedian pv; mid-

tibia with 1 submedian ad, 0-1 av and 1-2 pd; hind-tibia with 2 ad, 2 pd and 1-2

submedian av; claws and pulvilli as long as last tarsal segment.

Abdomen slender, about 1% times as long as broad. Colour and pollinosity the

same as that of the thorax, with a narrow median stripe, sometimes very weak.

Last three segments with marginal bristles and a few lateral discals in some of

the specimens.

Female — Frons at vertex about % as wide as the eye is long, widening towards

the antennal groove, frontal stripe subparallel, reddish-brown. Bucca about % as

high as the eye is long. Chaetotaxy of head consisting of iv, ev, oc, cruciate paf and

1 thick and 1 shorter and thinner fo.

Length: 5-6 mm.

The Pan-Pacific Entomologist 51: 130-133. April 1975

Vol. 51, No. 2, April 1975

131

Fig. 1. Phumosia rossi n. sp. a) cerci and paralobi, b) phallosome laterally,

c) parameres (Holotype).

There are 8 $ $ and 1 $ before me collected by E. S. Ross and R. E. Leech

at the following localities:

1) Nyasaland (Malawi), 16 miles west of Dedza 24.11.1958 ( holotype $ and

4 paratype $ $).

2) Northern Rhodesia (Zambia), Senga Hill, 40 miles south of Abercorn, 12.11.

1958 (2 paratype S $ and an allotype $ ).

3) Angola, 12 miles south of Villa Teixiera de Silva, 28.V.1958 (1 $ paratype).

Holotype, allotype and 4 paratypes have been returned to the Califor-

nia Academy of Sciences, 3 paratypes have been kindly presented to the

Department of Entomology of the South African Institute for Medical

Research, Johannesburg.

Pterelia kenyae new species (Sarcophagidae : Miltogramminae)

In the key to the species of Pterelia (Zumpt, 1961) the new species

runs down to P. santosdiasi, and on external features it falls within the

“obscurior- complex.” The males are easily distinguished by the

hypopygia.

Male — Eyes bare, facets small. Frons at vertex measuring about % of eye-

length. Frontal stripe yellow, darkening towards vertex, subparallel, at the tip

of the ocellar triangle measuring about DA times as wide as the neighbouring

parafrontalium. Parafrontalia with dense yellow pollinosity, becoming pale yellow

with different light incidence. Ocellar triangle with oc accompanied by several

bristly hairs; iv, ev and f well developed, 2 proclinate fo and several bristly hairs

on the posterior parafrontalium; 10-12 paf. Face pale yellow, antennae yellow,

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The Pan-Pacific Entomologist

Fig. 2. Pterella kenyae n. sp. a) cerci and paralobi, b) phallosome laterally,

c) parameres (Holotype) .

third segment approximately 3 times as long as second, deep yellow; arista bare,

basal half thickened, yellow, tip black. Posterior bucca and occiput black with

short hairs, anterior bucca yellow. Height of bucca about Ho of eye-length. Palpi

yellow, slender, proboscis black.

Thorax black with light grey pruinosity, narrow long undusted stripes visible.

Chaetotaxy: ac — 0 + 1, 2 prescutellar clc and 1 weak presutural dc, ia = 0 + 1;

1 prs, 2 sa, 2 n and 3 h well developed. Pleura white pruinose, pp and pst long

and thick, st = 1:1 or 1:2. Wings hyaline, epaulet black, basicosta yellow, veins

yellow-brown, R 5 broadly open. Thoracic squama white, very broad, halter yellow.

Legs black, fore-tibia with 1 submedial posterior bristle; mid-tibia with 1 ad, 1 pv

and 3 weak pd; hind-tibia with a row of short ad, 3 or 4 av and 1 pd.

Abdomen longer than broad, black dorsally, reddish-brown laterally and

ventrally. Anterior and posterior margins of tergites III, IV and V broadly but

unevenly yellow pollinose. Hypopygium (fig. 2) with long slender cerci, shorter

bifid paralobi with a row of long bristles; the first three anterior bristles are

very well developed. Phallosome with the spinus protruding vertically as in the

other species of the “obscurior- complex”.

Female — not known.

Length: 6-8 mm.

Seven specimens have been received, collected by M. E. Irwin and E. S. Ross

in Kenya, Diani Beach, Kwale District, 4.1.1970 ( holotype and 3 paratypes),

and at Blue Lagoon, Kilifi District, 1.1.1970 (3 paratypes). Three specimens

have been kindly presented to the South African Institute for Medical Research,

Johannesburg.

Acknowledgements

I wish to thank Dr. F. Zumpt, head of the Department of Entomology,

for supervising my studies on Higher Diptera and Professor J. F.

Murray, Director of The South African Institute for Medical Research,

who provided the necessary working facilities.

Vol. 51, No. 2, April 1975

133

References

Zumpt, F. 1956. Calliphoridae (Diptera Cyclorrhapha) Part I: Calliphorini and

Chrysomyiini. Explor. Parc natn. Albert Miss. G.F. de Witte, 87:

1 - 200 .

Zumpt, F. 1961. Calliphoridae (Diptera Cyclorrhapha) Part III: Miltogram-

minae. Explor. Parc natn. Albert Miss. G.F. de Witte , 98: 1-137.

A Description of the First Instar Larva of

Eupompha imperialis and E. edmundsi

(Coleoptera: Meloidae) 1

John D. Pinto

Department of Entomology, University of California, Riverside 92502

The genus Eupompha (Lyttini: Eupomphina) consists of eight dis-

tinctive species, all confined to southwestern North America. The first

instar larva of three of these, E. elegans (LeConte), E. histrionica

(Horn) and E. schwarzi (Wellman) were described by MacSwain

(1956, as Calospasta) . Because background information on larval

anatomy is needed for a forthcoming paper on the courtship behavior

of Eupompha , descriptions of two additional species, E. imperialis and

E. edmundsi are presented at this time.

To facilitate species comparisons, descriptions conform, insofar as

possible, to the terminology employed by MacSwain ( 1956) .

Eupompha imperialis (Wellman)

(Fig. 1)

Body very light brown; heavy, spiniform setae on both dorsum and venter,

some of those on thoracic and abdominal tergites curved upward at apex. Head

wider than long, narrowing slightly to base; six setae between clypeus and frons

(first row behind labrum) ; gula as long as wide, setae inserted on anterior margin.

Eves large, about 15% greater in diameter than mesothoracic spiracles. Antennae

with segment I subrectangular, twice as wide as long; segment II 60% longer

than wide, twice as long as I; segment III twice as long as wide, 30% shorter

than II; sensory organ slightly wider than and subequal in length to III; terminal

seta twice as long as III. Mandibles slender, gradually widened basally, entire

to feebly crenulate; apical seta almost 4X as long as basal seta. Maxillae with

about five weak setae at apex of mala; segment III of palpi 40% longer than wide,

lateral margin 30% longer than medial margin, sensory area with about 20 papil-

lae, two-segmented appendix not evident. Labium with setae of first prementum

long, setae of second prementum minute; segment II of palpi longer than I, two-

segmented appendix at apex of II very small. Thorax with line of dehiscence

confined to pro- and mesonotum, and extreme apex of metanotum; 24 stout,

elongate setae on pronotal disc. Abdomen with posterolateral margin of tergites

abutting against pleurites; spiracles placed in membranous area between pleurites

and anterolateral margin of tergites; first spiracle with diameter about 20% less

than that of mesothoracic spiracle and twice the diameter of second spiracle,

spiracles on segments II-VIII gradually decreasing in diameter; tergites with

posterior marginal row of setae about % tergite length; sternum poorly sclerotized,

1 This study was supported by Grant GB-30907 from the National Science Foundation.

The Pan-Pacific Entomologist 51: 134-139. April 1975

Vol. 51, No. 2, April 1975

135

no evidence of sclerotization on segments I-VI, VII-IX each with a poorly defined

medial sclerite. Legs slender, hind claw long, about % as long as hind tibia;

claws with their two setae separated at base by about Ho claw length, longer seta

not approaching apex of claw (as in Fig. 3). Body length 1.30 mm; caudal

setae .58 mm. 2

Material Studied. Larvae from a mass of 180 eggs laid and hatching on 17 and

25 April, respectively (at 26°C). Adults collected 16 April 1974; Imperial

County, California, ca. 12 mi W Calexico, adjacent to International Boundary;

feeding on flowers of Coldenia palmeri Gray (Boraginaceae) .

Eupompha edmundsi (Selander)

(Figs. 2, 3, 5-8) 4

Body very light brown, venter with normal, elongate, spiniform setae through-

out; dorsum with both very short, stout setae, and highly modified, elongate,

clavate, costate setae which are elliptical in cross-section; costae on thoracic and

abdominal setae with apical projections; modified setae somewhat shorter than

unmodified homologues in other species. Modified setae distributed as follows.

Head capsule: widespread except on labium, also, lateral-most seta on first setal

row behind labrum, seta posteromedial to eye, and those on occiput normal;

thorax: on lateral margin of pronotum only; and on lateral and posterior margin

of meso- and metanotum; abdomen: on posterior margin of tergites only.

Unmodified dorsal setae much shorter than homologues in other species.

Head wider than long, narrowing to base; six setae between clypeus and frons

(first row behind labrum) ; gula as long as wide, setae inserted on anterior margin.

Eyes subequal in diameter to mesothoracic spiracles. Antennae with segment I

subrectangular, twice as wide as long; segment II as wide as long, 30-40% longer

than I; segment III twice as long as wide, subequal in length to II; sensory organ

slightly wider and shorter than III; terminal seta short, subequal in length to

III. Mandibles slender apically, abruptly widened at base, entire to feebly

crenulate; apical seta twice as long as basal seta. Maxillae with about 5 weak

setae at apex of mala; segment III of palpi 50% longer than wide, lateral margin

almost twice as long as medial margin, sensory area with about 25 papillae, two-

segmented appendix not evident. Labium with setae of first prementum long,

setae of second prementum minute; segments I and II of palpi subequal in length,

two-segmented appendix at apex of II very small. Thorax with line of dehiscence

confined to pro- and mesonotum, and extreme apex of metanotum; 26 setae on

pronotal disc. Abdomen with posterolateral margin of tergites abutting against

pleurites; spiracles placed in membranous area between pleurites and anterolateral

margin of tergites; first spiracle subequal in size to mesothoracic spiracle and

twice the diameter of second spiracle, spiracles on segments II-VIII gradually

decreasing in diameter; tergites with posterior marginal row of setae less than %

tergite length; sternum poorly sclerotized, no evidence of sclerotization on seg-

ments I-VI, VII-IX each with a poorly defined medial sclerite. Legs slender, hind

claw long, about %o as long as hind tibia; claws with their two setae basally

adjacent, longer seta not approaching apex of claw. Body length 1.22 mm;

caudal setae .60 mm. 2

2 Means based on the measurement of five slide-mounted specimens.

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The Pan-Pacific Entomologist

Figs. 1-2. Head (dorsal view) of first instar larvae of Eupompha. 1. E.

imperialis. 2. E. edmundsi. Illustrations of E. elegans are given by MacSwain

(1956).

Vol. 51, No. 2, April 1975

137

Figs. 3-4. Hind claw of first instar larvae of Eupompha showing relative

length and position of the two basal setae. 3. E. edmundsi. 4. E. elegans.

Material Studied. Larvae from masses of 142, 147, and 187 eggs, respectively.

Respective oviposition and hatching dates as follows: 27 June-12 July; 28 June-

13 July; 3-19 July (at 26°C). Adults collected 27 June 1972; Coconino County,

Arizona, 2 mi W Page; feeding on inflorescences of Heliantlius anomalus S. F.

Blake and Stephanomeria exigua Nutt. (Compositae) .

Discussion

The traits of the two species described here agree with MacSwain’s

(1956) description of Eupompha. The clavate and costate setae on the

dorsum of E. edmundsi are, so far as known, unique within the Meloi-

dae. Their number, position and relative size correspond to their

unmodified homologues in congeners (e.g., compare Figs. 1 and 2) .

The larvae of the five species discovered so far fall into two groups

with E. histrionica and E. schwarzi in one, and E. elegans, E. imper-

ials and E. edmundsi in the other. The latter three can be character-

ized as follows: six setae present between clypeus and frons, antennal

segment II as long as wide or longer, line of dehiscence absent or

incomplete on metanotum, and medial margin of maxillary palpal seg-

ment III distinctly shorter than lateral margin. In E. histrionica and E.

schwarzi there are eight setae between the clypeus and frons, antennal

segment II is wider than long, the line of dehiscence is complete on the

metanotum, and the margins of segment III of the maxillary palpi are

subequal.

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Figs. 5-8. Scanning electron micrographs of the first instar larvae of E.

edmundsi. 5. Modified setae on dorsum of head capsule (5000X). 6. Modified

seta on apical margin of abdominal tergite III (4000X)- 7. Apical view of a

modified seta on head capsule (6000X). 8. Spiracle on abdominal segment I

(6000 X ) •

The larvae of E. elegans , E. imperialis and E. edmundsi are similar,

the highly modified setae of the latter notwithstanding. E. imperialis

is phenetically intermediate. E. elegans is distinguished from both E.

edmundsi and E, imperialis by its darker color and more strongly

Vol. 51, No. 2, April 1975

139

sclerotized abdominal sterna, the presence of the line of dehiscence in

the basal half of the metanotum, the longer apical seta of the claws

(Fig. 4), the placement of the gular setae anterior to the gular margin,

the smaller eyes (smaller than the mesothoracic spiracles) and the

inconspicuous basal mandibular seta. E. elegans and E. imperialis,

on the other hand, are separated from E. edmundsi by the unmodified

dorsal setae, the longer second segment and terminal seta of the

antennae, the longer second segment of the labial palpi, and the less

abrupt basal widening of the mandibles.

The phenetic relationships of larvae and adults correlate fairly well.

Adults of E. schwarzi and E. histrionica are more similar to one another

than to any other Eupompha. E. imperialis and E. elegans adults also

correspond most closely with one another. Adults of E. edmundsi are

most similar to those of E. viridis (Horn), whose larva is unknown.

Males of both species lack certain head, antennal and foreleg modifica-

tions which are found in all other species. These modifications are

epigamic in function (Pinto, MS). Because of their generalized adults,

compelling evidence for including E. edmundsi and E. viridis in

Eupompha was lacking. The close correspondence of the larva of E.

edmundsi to those of other Eupompha, and to E. imperialis in particu-

lar, considerably strengthens its position within the genus.

Acknowledgments

Host plants of E. edmundsi were identified by Mr. Oscar F. Clarke

(U. C. Riverside Herbarium). Figures 1-4 were prepared by Mr.

Carl T. Conley.

Literature Cited

MacSwain, J. W. 1956. A classification of the first instar larvae of the Meloidae

(Coleoptera) . Univ. Calif. Publ. Entomol., 12: 1-182.

The Larva of Cafius sulcicollis LeConte

(Coleoptera: Staphylinidae)

Ian Moore

Division of Biological Control, University of California, Riverside, CA 92502

James, Moore and Legner (1971) described larvae of four common

west coast species of Cafius and presented a key to include two European

species which were already known. Cafius sulcicollis LeConte is not

common in southern California so could not be included in the study.

Vincent D. Roth, Managing Director of the Southwest Research Station

of the American Museum of Natural History at Portal, Arizona has

given us a series of Cafius sulcicollis including larvae from Sonora,

Mexico where it is apparently common. We are taking this opportunity

to describe the larva. Among the west coast species of Cafius , this leaves

only the larvae of C. decipiens LeConte, C. femoralis Maklin and C.

opacus LeConte unknown.

Tabular Key to Some Larvae of Nearctic Cafius

Statement of characters.

Shape and segmentation of urogomphus = SHAPE SEG UROG

SPHER =: Spherical, one-segmented.

CYL = Cylindrical, two-segmented.

Length of urogomphus = LENG UROG

LONG =: longer than pseudopod.

SHORT = shorter than pseudopod.

Length of first segment of maxilla = LENG ONE MAX

LONG = as long as second segment.

SHORT = shorter than second segment.

Length of fourth segment of maxilla = LENG FOUR MAX

LONG = as long as penultimate segment.

SHORT = shorter than penultimate segment.

Source

James = James, Moore and Legner 1971.

SHAPE

SEG

MAX

LENG

UROG

LENG

ONE

MAX

LENG

FOUR

MAX

SOURCE

NAME

CYL

LONG

SHORT

LONG

SPECIMEN

sulcicollis LeConte

CYL

LONG

SHORT

JAMES

luteipennis Horn

CYL

LONG

SHORT

JAMES

lithocharinus LeConte

CYL

SHORT

JAMES

canescens Maklin

SPHER

SHORT

JAMES

seminitens Horn

The Pan-Pacific Entomologist 51 : 140 - 142 . April 1975

Vol. 51, No. 2, April 1975

141

Figs. 1-4. Larva of Cafius sulcicollis LeConte. 1. Clypeus. 2. Pseudopod an

urogomphi. 3. Antenna. 4. Maxilla.

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Larva of Cafius sulcicollis LeConte

Color . — Head pale ferrugineous ; thorax ferrugineous; abdomen, legs, antennae

and mouthparts testaceous.

Head subquadrate, widest near base, very slightly narrowed to apical angles.

Neck about three-fourths as wide as head. Ocelli lacking. Clypeus with nine

teeth, the central tooth and the tooth adjacent to the outer tooth smallest. Antenna

four-segmented; first segment about as long as wide; second segment about as

wide as and about twice as long as first; third segment a little longer and about

as wide at apex as first, tapered to base, with an ovid acorn-type seta at apex

which is almost as long as fourth segment; fourth segment about one-third as wide

and half as long as third, with an acorn-type seta at apex which is slightly smaller

than the acorn-type seta of third segment. Maxilla with stipes as long as palpus;

galea very small, slender; palpus four-segmented, first segment about as long

as wide, second segment about as wide as and twice as long as first, third seg-

ment narrower and a little shorter than second, fourth segment only a little

narrower and a little shorter than third. Ligula not quite as long as first segment

of labial palpus. Labial palpus three-segmented, each segment narrower but about

as long as preceding. Gular sutures united in basal three-fifths, thence diverging

to apex.

Thorax . — Pronotum about one-eighth wider than long; widest at base; sides,

apex and base gently arcuate; basal angles narrowly rounded; apical angles

broadly rounded; surface polished. Mesonotum and metanotum each about as

wide as and half as long as pronotum, with a few scattered fine setae, surface

polished.

Abdomen gradually narrowed from near base to apex, the basal two segments

shorter than the rest; with a few fine scattered setae; surface finely microreticulate.

Pseudopod more than twice as long as wide. Urogomphus a little longer than

pseudopod, two-segmented; first segment almost as long as urogomphus, very

slender; second segment about half as wide and one-fourth as long as first.

Length 8.0 mm.

Material examined, seven specimens from Mexico, Sonora, Punta Cirio, 29.53-

112.40, 20 March 1974, wrack on sandy beach, V. Roth & W. Brown collectors, in

company with 93 adults.

Notes. This species is readily distinguished from all other known

larvae of Cafius by the large apical segment of the maxillary palpus.

The total lack of ocelli is similar to the condition in C. seminitens in

which the ocelli are so pale as to be almost indiscernible.

Literature Cited

James, G., Ian Moore and E. F. Legner. 1971. The larval and pupal stages of

four species of Cafius (Coleoptera: Staphylinidae) with notes on their

biology and ecology. Trans. San Diego Soc. Natur. Hist., 16: 279-289,

8 figs.

A New Species of Apanteles from South America

Being Introduced into California

(Hymenoptera: Braconidae)

Paul M. Marsh

Systematic Entomology Laboratory, IIBIII, Agr. Res. Serv., USDA 20560 1

A new species of Apanteles is described to provide a name for

studies being conducted at the University of California, Riverside.

This species is being released in California against the potato tuber-

worm, Phthorimaea operculella (Zeller) and the tomato pinworm,

Keiferia lycopersicella (Walsingham) . It was collected in Colombia as

a parasite of Scrobipalpula absoluta (Meyrick) . The National Collection

also contains specimens from Chile and Peru. Jack C. Hall, Division

of Biological Control, University of California, Riverside, California,

supplied the material from Colombia and has requested the name.

Apanteles gelechiidivoris, new species

Female. Length of body: 2.5 mm. Color: black except mandibles, palpi, apices

of femora, tibiae and tarsi which are testaceous or dark brown; stigma translucent

and margined on all sides by brown, wing veins brown. Head: shining, finely

punctate, densely covered by short white pubescence; malar space about as long

as clypeus; face only very slightly narrower at clypeus than at antennae, at its

narrowest part about equal to eye height; antennae shorter than body. Thorax:

stout; mesonotum flat, shiny, very finely punctate, punctures separated, densely

and evenly covered with short white pubescence; disc of scutellum flat, shining,

sparsely punctured, polished area on lateral face of scutellum semicircular, small;

propodeum (fig. 1) smooth and shining on dorsal surface, slightly rugose laterally

and strongly rugose medially at apex where rugae form lower part of an areola,

costulae absent; meso- and metapleura smooth and polished. Legs: hind coxa

smooth and polished, with a few scattered punctures dorsally at base; inner spur

of hind tibia a little longer than outer and less than half as long as hind basitarsus.

Wings (fig. 3) : stigma short and broad, broader than length of first segment of

radius, inner and outer sides nearly equal in length; metacarpus as long as

stigma; first segment of radius slightly curved, longer and narrower than inter-

cubitus; nervellus slightly curved at apex; vannal lobe evenly convex and with

fringe of hair. Abdomen: short, often compressed laterally at apex in dead

specimens; median plate of first tergum nearly parallel sided, only slightly

narrower at apex than at base, longer than apical width, smooth at base and

rugose at apex, no indication of median depression at apex (fig. 2) ; median plate

of second tergum nearly four times as wide as long, apical width about two times

basal width, weakly rugose at base (fig. 2) ; hypopygium acute at apex and ex-

tending slightly beyond apex of abdomen; ovipositor about as long as hind tibia,

evenly curved downward.

1 Mail address: c/o U. S. National Museum, Washington, D. C. 20560.

The Pan-Pacific Entomologist 51: 143-146. April 1975

144 .

The Pan-Pacific Entomologist

Vol. 51, No. 2, April 1975

145

Male. Essentially as in female except antennae longer than body, median plate

of first tergnm slightly narrower at apex, and median plate of second tergum

slightly longer.

Holotype female: Colombia, Palmira, V-14-74, A. Sandarriaga, ex. Scrobipal-

pula absoluta on tomato. USNM type no. 73373. Paratypes: Chile: Arica, 7 $ 2,

XI-1965, H. Vargas, ex. Gnorimoschema absoluta on tomato; Arica, 11 2 2,

3 $ $ , no date or collector, ex. Gnorimoschema absoluta. Colombia: 5 2 2,

3 $ $ , same data as holotype; Garagoa, 4 2 2,5 $ $, 4-30-73, E. R. Oatman,

ex. geleehiid on tomato; Mosquera, 5 2 2,5 $ S, 5-4-73, E. R. Oatman, ex.

gelechiid on potato. Peru: LaMolina, 4 2 2,1 $, XI-3-64, A. S. Guido and

Silveira ; Canete, 7 2 2,5 $ $ , XI-3, 6, 26-1964, A. S. Guido and Silveira ; no

locality, 2 2 2, 1966, ex. Gnorimoschema operculella on tomato. Paratypes de-

posited in U. S. National Museum, University of California, Riverside, and in

the Introduced Beneficial Insects Voucher Collection, Beneficial Insects Introduc-

tion Laboratory, Beltsville, Maryland.

Apparently this species is widely distributed along the Andes in

South America. It belongs to the laevigatus group as defined by Nixon

(1965 Bull. Brit. Mus. (Nat. Hist.), Ent., Suppl. 2, p. 181). In South

4 -

Figs. 1~2. Apanteles gelechiidivoris, n. sp. : 1, propodeum, dorsal view, X 160

magnification; 2, abdominal terga 1 and 2, X 185 magnification. Scanning

electron micrographs made at 5kv on a Cambridge II Scanning Electron

Microscope.

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America, it is similar to subcristatus Blanchard but can be distinguished

by the smoother propodeum, sculptured median plate of the second

tergum, and by the median plate of the first tergum being wider at the

apex. In North America, gelechiidivoris is closest to miantonomoi

Viereck but differs in the smoother propodeum and more slender

hind tibia.

Scanning electron microscope time for this project was supported in

part by the Electron Microscope Central Facility, Center of Materials

Research, University of Maryland, College Park.

Host Records for Some Species of Pompilidae From the

Southwestern United States and Mexico

(Hymenoptera)

Frank E. Kurczewski

Department of Entomology , State University of New York

College of Environmental Science and Forestry, Syracuse, New York 13210

The nesting behaviors and host records of many species of Pompilidae

from the southwestern United States and Mexico remain virtually un-

known. This paper attempts to fill in some of the host record gaps.

In addition to presenting first host records for Priocnemioides an-

gusticeps, P. aratus, and Anoplius percitus , new host species of spiders

are listed for Ageniella ( Ageniella ) conflicta Banks, A. (A.) partita

Banks, A. ( Priophanes ) arcuata (Banks), Anoplius ( Lophopompilus )

aethiops (Cresson) , A. ( Notiochares ) lepidus atramentarius (Dahlbom),

A. ( Arachnophroctonus ) relativus (Fox), A. ( Anoplius ) ithaca

(Banks), and A. (A.) toluca (Cameron). The host genus of Dipogon

( Deuteragenia ) thoracicus Townes and a host species of Anoplius

( Arachnophroctonus ) americanus ambiguus (Dahlbom) are verified.

All host records and associations are based upon specimens collected

in Arizona, New Mexico, Texas, and Mexico by R. E. Acciavatti (REA) ,

U. S. Forest Service, Albuquerque, N. M., R. C. Miller (RCM), Cornell

University, Ithaca, N. Y. and V. D. Roth (VDR), The Southwestern

Research Station, Portal, Arizona. H. E. Evans of Colorado State Uni-

versity, Fort Collins, Colo., determined Priocnemioides aratus and con-

firmed the identity of Anoplius percitus. The host spiders were

identified by W. J. Gertsch, Portal, Arizona, Y. D. Roth, and R. A.

Norton, SUNY College of Environmental Science and Forestry, Syra-

cuse, N. Y.

Tribe Pepsini

Priocnemioides angusticeps Townes. — Host: Lycosa antelucana Montgomery

(Lycosidae) , ad. $. TEXAS. Tarrant Co. Colleyville, June 7, 1972 (REA).

Priocnemioides aratus Townes. — Host: Lycosa carol.inensis Walckenaer (Lycosi-

dae) , ad. $ . MEXICO. Sonora. 30 mi. s.e. Agua Prieta on mesa nr. Cajon

Bonito, August 24, 1972 (VDR). “Lycosa spider in burrow of spider.”

Dipogon ( Deuteragenia ) thoracicus Townes. — Host: Xysticus sp. (Thomisidae) ,

imm. ARIZ. Cochise Co. S.W.R.S. 5 mi. w. Portal. 5400', September 18, 1964

(VDR).

The Pan-Pacific Entomologist 51: 147-151. April 1975

148

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Tribe Auplopodini

Ageniella ( Ageniella ) conflicta Banks. — Hosts: (a) Pardosa valens Barnes

(Lycosidae), ad. 2, with all legs amputated at coxal-trochanter joints. ARIZ.,

Cochise County, S.W.R.S. 5 mi. w. Portal. 5400', June 11, 1973 (RCM). (b) P.

valens Barnes (Lycosidae) , imm., with 3rd, 4th legs on both sides amputated.

Collection data : Same as above, (c) Trochosa gosiuta Chamberlain (Lycosidae),

imm., with all legs amputated at coxal-trochanter joints. Collection data: Same

as above except June 14, 1973.

Ageniella ( Ageniella ) partita Banks. — -Host: Pardosa falcifera F. Cambridge

(Lycosidae), imm. $ with ? legs amputated. Collection data: Same as above

except May 27, 1964 (VDR) .

Ageniella ( Priophanes ) arcuata (Banks). — Host: Sassacus papenhoei Peckham

(Salticidae) , ad. 2, with all legs amputated at coxal-trochanter joints. ARIZ.

Cochise Co. Portal, July 31, 1973 (RCM) .

Tribe Pompilini

Anoplius ( Lophopompilus ) aethiops (Cresson). — Hosts: (a) Lycosa santrita

Chamberlain & Ivie (Lycosidae) . ARIZ. Cochise Co. S.W.R.S. 5 mi. w. Portal.

5400', October 14, 1965 (VDR). (b) Lycosa carolinensis Walckenaer (Lycosidae),

ad. $ . N. MEX. Catron Co. 0.7 mi. w. Quemado at Largo Creek. 6900', October

2, 1973 (REA).

Anoplius ( Notiochares ) lepidus atramentarius (Dahlbom). — Host: Lycosa

antelucana Montgomery (Lycosidae), imm. $. TEXAS. Tarrant Co. Colleyville,

June 7, 1972 (REA) .

Anoplius ( Arachnophroctonus ) americanus ambiguus (Dahlbom). — Host: Arc-

tosa littoralis (Hentz) (Lycosidae), imm. 2. N. MEX. Sandoval Co. 1 mi. s. San

Ysidro at Rio Salado. 5500', August 18, 1973 (REA).

Anoplius ( Arachnophroctonus ) relativus (Fox). — Host: Arctosa littoralis

(Hentz) (Lycosidae), ad. 2- (2 records). TEXAS. Cooke Co. 0.7 mi. e.

Rosston at Clear Creek, October 10, 1973 (REA).

Anoplius ( Pompilinus ) percitus Evans. — Host: Trochosa sp., prob. gosiuta

Chamberlain (Lycosidae) , imm. N. MEX. Bernalillo Co. Albuquerque, 2 mi.

n.e. Univ. of Albuquerque. 4980', June 24, 1973 (REA).

Anoplius ( Anoplius ) ithaca (Banks). — Host: Pardosa steva Lowrie and

Gertsch (Lycosidae) , ad. 2 - N. MEX., Grant Co. 0.5 mi. w. Lake Roberts along

Hwy. 35 6000', July 11, 1972 (REA).

Anoplius ( Anoplius ) toluca (Cameron). — Hosts: (a) Schizocosa avida group

(Lycosidae), ad. 2- N. MEX., Hidalgo Co. 20 mi. n. Rodeo, June 13, 1973

(RCM). (b) Trochosa gosiuta Chamberlain (Lycosidae), ad. 2- ARIZ., Cochise

Co. S.W.R.S. 5 mi. w. Portal. 5400', September 17, 1965 (VDR) .

Discussion

In the present paper, Priocnemioides angusticeps and P. aratus are

recorded with prey for the first time. These records bring to five the

number of Nearctic species and subspecies of Priocnemioides for which

host records are now available, and they substantiate the fact that species

of this pompilid genus, regardless of size, prefer wolf-spiders belonging

Vol. 51, No. 2, April 1975

149

to the genus Lycosa. The observation of P. aratus attacking L. caro-

linensis in its burrow and then leaving the spider there suggests that

some species of Priocnemioides may utilize the spider’s burrow as a

nest. On the other hand, Janvier (1930) and Evans and Yoshimoto

(1962) pointed out that other species of Priocnemioides do, in fact,

construct their own burrow and transport the spider backwards on the

ground to the nest.

The record of Dipogon ( Deuteragenia ) thoracicus preying on

Xysticus sp. (Thomisidae) substantiates Evans’ (1974) prey record

for this species and, in addition, underlines the fact that species in this

genus often utilize small thomisids and salticids as prey (see Medler

and Koerber, 1957; Evans and Yoshimoto, 1962; Fye, 1965; Krombein,

1967; Kurczewski and Kurczewski, 1972).

Ageniella ( Ageniella ) conflicta has been reported to prey upon small

lycosids of the genera Arctosa, Lycosa , and Trochosa (Hurd and

Wasbauer, 1956; Townes, 1957; Evans and Yoshimoto, 1962;

Kurczewski and Kurczewski, 1968a). The records of small, adult and

immature Pardosa valens and immature Trochosa gosiuta for A. con-

flicta which are presented herein substantiate this preference.

Ageniella ( Ageniella ) partita has been recorded with prey several

times and these hosts comprise wandering spiders belonging to the

families Gnaphosidae and Lycosidae (summary in Kurczewski and

Kurczewski, 1968b). The record included herein of A. partita preying

upon Pardosa falcifera (Lycosidae) does not alter this host preference.

That of Ageniella ( Priophanes ) arcuata preying upon the salticid

Sassacus papenhoei introduces a new host family for this pompilid

species. Previously, A. arcuata had been reported to provision with

Oxyopidae and Thomisidae (Evans and Yoshimoto, 1962; Evans, 1964).

Although the first three host records for this pompilid have revealed

three host families of spiders, A. arcuata is nevertheless selecting spiders

with rather similar ecologies.

The records presented herein for Anoplius ( Lophopompilus ) aethiops

bring to light two new host species which merely accentuate its pref-

erence for rather large wolf-spiders of the genus Lycosa (summary in

Kurczewski and Kurczewski, 1973). The fact that one of these records

is a male spider is noteworthy in view of the fact that the vast majority

of records for this species of pompilid are for female spiders. Likewise,

the record of Anoplius ( Notiochares ) lepidus atramentarius utilizing

Lycosa antelucana , a new host species, for provisions underlines its

preference for rather large wolf-spiders (see Evans, 1951; Evans and

Yoshimoto, 1955, 1962).

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The Pan-Pacific Entomologist

That of Arctosa littoralis (Lycosidae) for Anoplius ( Arachnophrocto -

nus ) americanus ambiguus substantiates earlier records for this host

species of spider (see Evans, 1951). Additional host records for A.

americanus ambiguus include species of Lycosidae and Oxyopidae

(Evans and Yoshimoto, 1955, 1962; Hurd and Wasbauer, 1956;

Wasbauer and Powell, 1962). The record of Arctosa littoralis as prey

of Anoplius ( Arachnophroctonus ) relativus is, however, new. Previous

records for this species of spider-wasp comprise the lycosid genera

Lycosa and Geolycosa and the agelenid genus Agelenopsis (summary

in Evans and Yoshimoto, 1962; Kurczewski and Kurczewski, 1968a,

b, 1973).

Anoplius ( Pompilinus ) percitus has previously not been collected

with prey and the record presented herein, therefore, indicates an

initial preference for small Lycosidae ( Trochosa sp.). Other species

in this subgenus which utilize small Lycosidae exclusively as prey in-

clude clystera (Banks) (Wasbauer and Powell, 1962), cylindricus

(Cresson) (Kurczewski and Kurczewski, 1968a, 1973), estellina

(Banks) (Evans, 1964), krombeini Evans (Evans and Yoshimoto,

1962; Kurczewski and Kurczewski, 1973), s. stenotus (Banks) (Krom-

bein and Evans, 1955; Kurczewski and Kurczewski, 1973), and stenotus

bequaerti (Dreisbach) (Krombein, 1964). Two of the species, cylin-

dricus and estellina, either utilize the spider’s burrow for a nest or nest

in a pre-existing burrow or depression in the soil.

The record of Pardosa steva as prey of Anoplius ( Anoplius ) ithaca,

albeit new, emphasizes the fact that this species of pompilid prefers

small lycosids of the genera Pardosa and Arctosa (Evans and Yoshimoto,

1962; Kurczewski and Kurczewski, 1968a, 1973). Anoplius ( Anoplius )

toluca has been previously collected with Lycosa sp. (Lycosidae) (Evans,

1964) and, therefore, our two records for the lycosid genera Schizocosa

and Trochosa only narrowly extend the host selection parameters of

this species of pompilid.

Literature Cited

Evans, H. E. 1951. A taxonomic study of the Nearctic spider wasps belonging

to the tribe Pompilini (Hymenoptera : Pompilidae). Part II: Genus

Anoplius Dufour. Trans. Amer. Entomol. Soc., 76: 207-361.

1964. Notes on the prey and nesting behavior of some solitary wasps of

Mexico and southwestern United States (Hymenoptera: Sphecidae

and Pompilidae) . J. Kansas Entomol. Soc., 37 : 302-307.

1974. A review of the species of Dipogon occurring in Central America,

Mexico, and extreme southwestern United States (Hymenoptera,

Pompilidae). Trans. Amer. Entomol. Soc., 100: 29-51.

Vol. 51, No. 2, April 1975

151

Evans, H. E. and C. M. Yoshimoto. 1955. An annotated list of pompilid wasps

taken at Blackjack Creek, Pottawatomie Co., Kansas (Hymenoptera) .

J. Kansas Entomol. Soc., 28: 16-19.

1962. The ecology and nesting behavior of the Pompilidae (Hymenoptera)

of the northeastern United States. Misc. Pub. Entomol. Soc. Amer.,

3: 67-119.

Fye, R. E. 1965. The biology of the Vespidae, Pompilidae, and Sphecidae

(Hymenoptera) from trap nests in northwestern Ontario. Can. Entomol.,

97: 716-744.

Hurd, P. D., Jr. and M. S. Wasbauer. 1956. New host records for North Amer-

ican spider-wasps (Hymenoptera: Pompilidae). J. Kansas Entomol.

Soc., 29: 168-169.

Janvier, H. (M. F. Claude- Joseph) . 1930. Recherches biologiques sur les

predateurs du Chili. Ann. Sci. Nat. Zool., 13: 235-354.

Krombein, K. V. 1964. Results of the Archbold Expeditions. No. 87. Biological

notes on some Floridian wasps (Hymenoptera, Aculeata) . Amer. Mus.

Nov., 2201: 1-27.

1967. Trap-nesting wasps and bees: Life histories, nests, and associates.

Washington, Smithsonian Press, vi -j- 570 p.

Krombein, K. V. and H. E. Evans. 1955. An annotated list of wasps collected

in Florida, March 20 to April 3, 1954 (Hymenoptera, Aculeata). Proc.

Entomol. Soc. Wash., 57: 223-235.

Kurczewski, F. E. and E. J. Kurczewski. 1968a. Host records for some North

American Pompilidae (Hymenoptera) with a discussion of factors in

prey selection. J. Kansas Entomol. Soc., 41: 1-33.

1968b. Host records for some North American Pompilidae (Hymenoptera).

First Supplement. J. Kansas Entomol. Soc., 41 : 367-382.

1972. Same title, Second Supplement. Tribe Pepsini. J. Kansas Entomol.

Soc., 45: 181-193.

1973. Same title. Third Supplement. Tribe Pompilini. J. Kansas Entomol.

Soc., 46: 65-81.

Medler, J. T. and T. W. Koerber. 1957. Biology of Dipogon sayi Banks

(Hymenoptera, Pompilidae) in trap-nests in Wisconsin. Ann. Entomol.

Soc. Amer., 50: 621-625.

Townes, H. 1957. Nearctic wasps of the subfamilies Pepsinae and Ceropalinae.

Bull. U. S. Nat. Mus., 209: 1-286.

Wasbauer, M. S. and J. A. Powell. 1962. Host records for some North Ameri-

can spider wasps, with notes on prey selection (Hymenoptera: Pompili-

dae). J. Kansas Entomol. Soc., 35: 393-401.

A New Species of Chersodromia from Mexico

(Diptera: Empididae)

Paul H. Arnaud, Jr.

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118

In the Americas, the empidid genus Chersodromia Walker is known

from one Neotropical species (from Panama; Smith, 1967) and eight

Nearctic species (from the eastern, southern, and western shores of

the United States; Melander, 1965). In the Palearctic, twenty-two spe-

cies of Chersodromia are presently known (Chvala, 1970). Collin

(1961: 61) considered the genus Chersodromia “an interesting natural

genus of great antiquity in which, while many of the species have be-

come particularly distinct as a result of evolution, practically all the

connecting links between them have continued to exist.” Chersodromia

adults usually occur on sandy beaches (mostly marine) where they

have the ability to run very rapidly over sand. Curran (1931: 11-12)

described the Neotropical species — longicomis — from his collecting at

Patilla Point, Panama. Chersodromia longicornis differed in its selec-

tion of habitat from other known species in that it occurred in holes

and crevices in lava rocks that were several hundred feet from the

shore line, but not wholly submerged at high tide.

The purpose of this paper is to report the discovery and to describe

and compare a new species that is closely related to C. longicornis ,

both in structure and in habitat preference. This new species also oc-

curs on rocks, but in this original collection it was taken from rocks

along the shore line. The type series was swept during mid-day along

with many Dolichopodidae and Canaceidae on the southwestern shore

(fig. 10) of Bahia de Mismaloya, about 8 km southwest of Puerto

Vallarta. The type locality is below the area where the movie “The

Night of the Iguana” was filmed.

Chersodromia madelinae, new species

(Figs. 1, 3-6, 9-10)

Diagnosis. — Elongate, small (length about 1.75 mm. in male; 2.0 mm. in

female), brownish-black species; thorax and abdomen thinly grayish-brown

pollinose; 1 anterodorsal bristle on front tibiae at about basal third; a row of a

dozen short anteroventral bristles on apical half of male mid-tibiae, lacking in

female; hypopygium of male with right cercus broad and not attaining apices of

dististyles and process of left basistyle with long bristles near apex.

Holotype male. — Length, about 1.75 mm., wing length about 2.1 mm. Head

brownish-black, face, frons and back of head grayish pollinose, cheeks about

The Pan-Pacific Entomologist 51: 152-158. April 1975

Vol. 51, No. 2, April 1975

153

Fig. 1. Chersodromia madelinae, new species, right lateral view, paratype

female.

one-fourth eye height, eyes with slight excisions opposite bases of antennae

and concave on more than lower half of hind margin, face carinate, front at

narrowest the width of ocellar protuberance, with a divergent pair of ocellar

bristles and another convergent pair at edges of eyes at side of ocellar

protuberance, two pairs of vertical bristles. Proboscis brown, exposed portion

about half of head height. Palpi blackish, flattened, moderately long haired.

Antennae dark brown, third segment elongate and tapered, with curved pubescence;

aristae slender, about length of first three antennal segments, very short pubescent.

Thorax brownish-black, thinly grayish-brown pollinose; bristles and hairs of

mesonotum black, acrostichals small, 4 pairs of dorsocentrals, scutellum with two

pairs of marginal scutellar bristles, the apical long while the outer very small.

Legs dark brown, front tibiae with 1 anterodorsal bristle at about basal third;

mid-tibiae with 1 anterodorsal bristle at about basal third and with a row of a

dozen short anteroventral bristles on apical half; hind tibiae with six bristles — 2

dorsals, 2 anterodorsals, and 2 anteroventrals. Wings about 2.1 mm. long and

0.75 mm. wide; venation as illustrated (fig. 3), veins light brown, crossveins

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Figs. 2-4. Chersodromia species, wings, upper surfaces. Fig. 2, C. longicornis

(Curran), left wing, paratype female. Fig. 3, C. madelinae, new species, left wing,

holotype male. Fig. 4, C. madelinae, new species, right wing, allotype female.

Illustrations made by projecting wing slide preparations through a B and L Tri-

Simplex Microprojector onto a number 5 Azo single weight paper. Projected at

same magnification.

Vol. 51, No. 2, April 1975

155

Figs. 5-8. Chersodromia species, holotype males, hypopygia. Figs. 5-6, C.

madelinae, new species, ventrolateral and dorsolateral views. Figs. 7-8, C. longi-

cornis (Curran), ventrolateral and dorsolateral views.

indistinct; membrane not clear, with dense microtrichia giving a light yellow-

brown tinge. Halteres with knobs blackish-brown, stalks at bases yellow.

Abdomen brownish-black, minute hairs yellow otherwise black haired, thinly

grayish-brown pollinose, lightly shining. Asymmetrical hypopygium (figs. 5-6)

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Fig. 9. Chersodromia madelinae, new species, allotype female, apical abdominal

segments and ovipositor, lateral view.

rotated, brownish-black, yellow bristled; right cercus broad and not attaining

apex of dististyles; process of left basistyle broader before apex, with long

bristles near apex.

Allotype female. — Length, about 2.0 mm., wing length about 2.5 mm. by 0.9 mm.

in width (fig. 4). Similar in coloration to male; mid-tibia lacks comb of short

bristles on apical half; ovipositor elongate, tapering and as figured (fig. 9).

Type series . — Holotype male, Mexico: Jalisco, Bahia de Mismaloya, 1-1-1971

(P. H. and M. Arnaud), PHA dissection no. 20IV71; mounted on minuten with

postabdomen and hypopygium stored in microvial in glycerine and left wing on

slide in balsam; deposited in the collection of the California Academy of Sciences,

Entomology Type No. 12170. Allotype female, same collection data as holotype,

except PHA dissection no. 7VIII71; mounted on minuten, with portion of abdomen

and ovipositor stored in microvial in glycerine and right wing mounted on slide

in balsam; Entomology Type No. 12170. Paratypes, 4 males and 6 females, same

collection data as holotype; deposited in the following collections: 1 female,

California Academy of Sciences; 1 male, 1 female, National Museum of Natural

History; 1 male, 1 female, American Museum of Natural History (exchange) ; 1

male, 1 female, collection of Dr. Milan Chvala (exchange) ; 1 female, sent to

Director General de la Fauna Silvestre, Mexico, D. F., Mexico; and 1 male, 1

female, collection of author.

The paratypes are rather similar in size. The wing of one male

paratype, which has been mounted in balsam on a slide, is 2 mm. in

length. In working with such small creatures, when they are pinned and

not mounted on microscope slides, measurements are difficult to make.

I take pleasure in naming this species for my wife, Madeline Milliet

Arnaud, who enthusiastically participated in its collection.

Comparison between Chersodromia madelinae and C. longicornis . —

The male of C. madelinae differs from C. longicornis by the following

characters: (1) madelinae has tibial bristling with only 1 anterodorsal

bristle (not 2) on the anterior tibiae and mid-tibiae with a row of

Vol. 51, No. 2, April 1975

157

Fig. 10. Bahia de Mismaloya, Mexico, type locality of Chersodromia madelinae,

new species. Type series swept from rocks in ocean along roadway, right of center.

about a dozen short anteroventral bristles on apical half (not with an

elongate row of about two dozen shorter bristles, on more than apical

two-thirds of tibiae) ; (2) the male hypopygium of madelinae (figs.

5-6) differs from that of longicornis (figs. 7-8) by its shorter, broader

right cercus and the broad long bristled process of the left basistyle.

Conversely longicornis differs from madelinae by its right cercus which

is longer than the processes of the dististyles and the process of the left

basistyle is narrower and without long bristles near apex and with 2

bristles on outer side of mid-region on the apical half. The female of

C. madelinae differs from C. longicornis in tibial bristling of the an-

terior leg as there is only 1 anterodorsal bristle in madelinae while

there are 2 in longicornis.

Chersodromia longicornis (Curran)

(Figs. 2, 7-8)

Coloboneura longicornis Curran, 1931, pp. 11-12.

Chersodromia longicornis : Smith, 1967, p. 39.6.

As a supplement to Curran’s description of longicornis, illustrations of the male

hypopygium and a print of the female wing are included. Figures 7 and 8 illustrate

the hypopygium of the male holotype which is deposited in the collection of the

American Museum of Natural History. See comparison in description of hy-

popygium of C. madelinae, new species. Figure 2 illustrates the wing of the female,

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from the paratype which, through an exchange, is now deposited in the collection

of the California Academy of Sciences. The wing is about 2.5 mm. in length

and 0.9 mm. in width.

Acknowledgments

The author would like to thank the following persons for their aid:

Dr. Lloyd Knutson and Mr. George C. Steyskal, Agriculture Research

Service, c/o National Museum of Natural History, for information

concerning the holotype of Chersodromia longicornis (Curran), and

to Mr. Steyskal also for terminology applied to the male hypopygium

of this genus; Dr. Pedro W. Wygodzinsky, American Museum of Nat-

ural History, for the loan of type material of C. longicornis and for

exchange of material; Miss Carolyn Mullinex, California Academy of

Sciences, who prepared the line drawings of figures 1, 5-9; and Mr.

Maurice C. Giles, California Academy of Sciences, for preparing the

photographic print used in figure 10 which was made from a color

transparency.

Collecting in Mexico was by permit number 45-70 from the Director

General de la Fauna Silvestre, Departamento de Conservacion y Propa-

gacion de la Fauna Silvestre.

Literature Cited

Chvala, M. 1970. Descriptions of nine new species of Palaearctic Chersodromia

Walk. (Diptera, Empididae), with notes on the genus. Acta Entomol.

Bohemoslov., 67(6): 384-407, figs. 1-35.

Collin, J. E. 1961. Empididae. Part I. Tachydrominae. Yol. 6, pp. i-vi, 1-219,

frontispiece, figs. 1-73. In [Verrall, G. H., ed.], British flies. Cam-

bridge, England.

Curran, C. H. 1931. New species of Empididae from Panama. Amer. Mus.

Novit., 467: 1-12.

Melander, A. L. 1965. Family Empididae (Empidae, Hybotidae) . Pp. 446-481.

In Stone, A. et al., A catalog of the Diptera of America north of Mexico.

U. S. Dep. Agric., Agric. Handb. No. 276, pp. I-1V, 1-1696.

Smith, K. G. V. 1967. 39. Family Empididae (Empidae, Hybotidae). Pp. 39.1-

39.67. In, A catalogue of the Diptera of the Americas south of the

United States. Dep. Zool. Secretar. Agric., Sao Paulo.

A New Species of Indialis from India

(Ephemeroptera : Leptophlebiidae)

William L. Peters

Florida A & M University, Tallahassee, Florida 32307

Peters and Edmunds (1970) established Indialis for the species

Indialis badia Peters and Edmunds. The species description was based

on one male subimago and 49 nymphs collected by me and my wife in

Kerala and Andhra Pradesh States, India. Peters and Edmunds (1970)

included the subimaginal description, as the one subimago was the

only subadult or adult known for Indialis. Recently I studied a male

imago from Kerala State which I describe herein as a new species of

Indialis.

The new species is congeneric with Indialis badia ; however, there

are several differences between the species in the wings and eyes. In

the fore wings of Indialis rossi veins R s and MP are forked nearly

equidistant from the base (Fig. 1) ; in Indialis badia vein MP is forked

more basally than the fork of vein R s . In the key to the imagos by Peters

and Edmunds (1970) Indialis keys to the first part of couplet 17 based

on the forks of veins R s and MP. Indialis rossi keys to the

second part of couplet 17, and will eventually key to couplet 25 and

Castanophlebia. Indialis and Castanophlebia can be readily distin-

guished by the wing venation and male genitalia.

The Cu-A area (Fig. 1) of the fore wings of Indialis rossi is broader

and more developed than that of Indialis badia. Also the costal pro-

jection (Figs. 2-3) of the hind wings of Indialis rossi is broader than

that of Indialis badia.

The male eyes of Indialis rossi are separated on the meson of the

head by a length 3% times as long as the maximum width of a lateral

ocellus (Fig. 4). Peters and Edmunds (1970) stated that the male

eyes of Indialis badia meet on the meson of the head; however, the

head of the subimaginal holotype is shriveled and the eyes of Indialis

badia might be separated on the meson of the head. Thus, the generic

limits of the eye character in Indialis have not yet been determined

definitively.

I would like to thank Janice G. Peters for preparation of the illustra-

tions, and Dr. George F. Edmunds, Jr., University of Utah, for bringing

the specimen to my attention. This research was supported by grant

no. 416-15-16 from the Cooperative State Research Service, United

States Department of Agriculture.

The Pan-Pacific Entomologist 51 : 159 - 161 . April 1975

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Fig. 3. Hind wing enlarged. Fig. 4. Dorsal view of head. Fig. 5. Ventral view of

genitalia.

Inclialis rossi, New Species

(Figs. 1-5)

Male imago (in alcohol). — Length: body, 7.1 mm; fore wings, 6.5 mm.

Upper portion of eyes red-brown, lower portion black. Head brown, area around

ocelli washed with black. Antennae brown, flagellum paler. Basal half of

ocelli black, apical half brownish- white. Thorax: brown, carinae darker, sutures

paler, carinae of pleurae and pronotum blackish. Coxae brown, trochanters

pale. (Remainder of legs broken off and missing.) Wings (Figs. 1-3) : longi-

tudinal veins and cross veins of fore and hind wings yellowish-brown, cross veins

in basal half of cells C and Sc of fore wings darker; membrane of fore and hind

wings hyaline, except base of fore and hind wings yellowish-brown, all cross

veins in fore wings surrounded with narrow, yellowish-brown clouds, apical X A

of cells C and Sc of fore wings translucent, white. Abdomen: segments 1-7

translucent, washed with brown, segments 8-10 opaque, washed with brown;

terga 1-9 with a narrow, darker brown, transverse band on posterior margin of

each tergum, band darker and wider on terga 7 and 8, terga 1-9 with a narrow,

lighter brown, longitudinal, median line extended entire length of each tergum;

spiracles dark brown, tracheae washed lightly with dark brown; sterna 1-8 with

a narrow, darker brown, transverse band on posterior margin of each sternum.

Genitalia (Fig. 5) : brown. Caudal filaments pale; wide, dark brown annulations

at articulations.

Female imago and mature nymph.— Unknown.

Holotype male imago, India, Kerala State, Kottayam District, Kittikanam,

nr. Peermade, 1000 M, 22 March 1962, E. S. Ross and D. Q. Cavagnaro. Holotype

Vol. 51, No. 2, April 1975

161

is preserved in alcohol and deposited in the collections of California Academy

of Sciences.

Etymology. — Species is named for Dr. E. S. Ross, California Academy of

Sciences.

Discussion . — Indialis rossi can be distinguished from Indialis badia

by the following combination of characters in the male imago, (1) all

cross veins in fore wings are surrounded with narrow, yellowish-brown

clouds (Fig. 1), (2) abdominal segments 1-7 are translucent and

washed with brown, (3) caudal filaments are pale, with wide, dark

brown annulations at articulations, and (4) the costal projection of

the hind wings is well developed and broadly rounded at the apex

(Figs. 2-3).

Literature Cited

Peters, W. L. and G. F. Edmunds, Jr. 1970. Revision of the generic classifica-

tion of the Eastern Hemisphere Leptophlebiidae (Ephemeroptera) .

Pac. Insects, 12: 157-240.

A New Acanthinus from Panama

(Coleoptera: Anthicidae)

Donald S. Chandler

Department of Entomology

The Ohio State University, Columbus, Ohio 43210

In a recent series of ultraviolet light trap samples taken in the

Canal Zone of Panama, there were present three species of Acanthinus

which could be placed in the aequinoctialis-G roup of Werner (1967).

Two of the species were named, aequinoctialis (LaFerte) and zeteki

Werner, while the third was undescribed. In several instances all three

species were collected during the same night and so present a problem

in species separation. The new species is more similar to aequinoctialis

and zeteki than to any of the others in the group and actually combines

certain of the characters which Werner used for the separation of these

species in his key.

The shape of sternite 8 of the males is the best character for species

separation. The emargination of this sternite in the new species is

moderately deep and wide (Fig. 2). In zeteki the emargination is very

deep and narrow (Fig. 3). Sternite 8 in aequinoctialis may be shallowly

emarginate (Fig. 1). Werner notes in his key to the group that the

sternite of aequinoctialis varies from truncate to gently convex. This

description varies slightly from Figure 1, but examination of specimens

from Brazil has convinced me that the Panama population is conspecific.

The new species may be separated by rearrangement of Werner’s key

to the aequinoctialis group.

Key to Acanthinus of the Aequinoctialis Group

1. Dorsal portion of elytra subopaque due to extremely fine granulation on the

surface between the punctures 2

Dorsal portion of elytra smooth and shiny 3

2. Base of head shallowly, almost evenly curved between the distinct temporal

angles. Elytra usually with an obscure, often interrupted, dark band across

the middle rugosus (LaFerte)

Base of head slightly produced at middle, the temporal angles feeble;

therefore nearly evenly semicircular behind the eyes. Elytra not banded

tucumanensis (Pic)

3. Male sternite 8 with deep notch at apex (Fig. 3) ; rugules on head ap-

parently not oriented in a particular direction; antennae appearing heavy

due to thickening of intermediate segments; usually with a dark midband

zeteki Werner

Male sternite with notch shallower (Fig. 2) or gently concave to convex

The Pan-Pacific Entomologist 51: 162-164. April 1975

Vol. 51, No. 2, April 1975

163

Figs. 1-5. Terminal sternites and genitalia of Acanthinus. Fig. 1. Stemite

8, aequinoctialis. Fig. 2. Sternite 8, umbilicatus. Fig. 3. Sternite 8, zeteki. Fig.

4. Internal sac, showing gonopore armature; a, ventral view; b, lateral view.

Fig. 5. Aedeagus.

(Fig. 1) ; rugules on head appearing to be at least slightly oriented longi-

tudinally; elytra without midband 4

4. Male sternite 8 with a distinct notch; head often with longitudinal area at

center of head without rugules and with sparse umblicate punctures,

punctures may be obscured by definite longitudinal rugules; antennae ap-

pearing heavy due to thickening of intermediate segments umbilicatus n. sp.

Male sternite 8 gently concave to convex; head always with distinct rugules

at center, no umbilicate punctures visible; antennae without thickening of

intermediate segments aequinoctialis (LaFerte)

The format of Werner is used in presenting the species description.

Acanthinus umbilicatus, New Species

1.98-2.43 mm., rufescent with paler elytra, varying to dark brown with elytra

brownish, coloration of elytra uniform. Similar in general appearance to

aequinoctialis, but with “heavy” antennae, strigules distinctly longitudinally

oriented and frequently with a longitudinal smooth central area on the head.

Holotype male: length 2.43 mm. Head 0.50 mm. long, 0.52 wide across eyes,

0.40 behind, tempora flat and nearly parallel, 0.07 long, temporal angles quite

sharp and at ca. 120°, base evenly curved between them. Surface shiny, punctures

small, umbilicate, smoothly strigose longitudinally, strigulations not dense, longi-

tudinal smooth area at center. Setae inconspicuous, tactile setae erect and fairly

stout, 0.08 mm. Eyes prominent, 0.25 X 0.18 mm., separated by 0.31, a line across

their hind margins 0.15 from base. Last segment of maxillary palpi securiform,

0.17 X 0.09 mm. Antennae fairly stout, gradually thicker toward apex, with tactile

setae 0.04 mm. long. Measurements in 0.01 mm.: 10/7, 7/5, 9/5, 7/5, 8/5, 7/5,

6/5, 7/7, 8/7, 9/8, 12/7. Segment 1 eccentrically pedunculate, 2-10 thickest near

apex, 7-10 subtruncate at base, truncate at apex, 11 subtruncate at base, convexly

tapered to a blunt point near base. Pronotum 0.50 mm. long, 0.34 wide at base,

0.31 at constriction, 0.43 maximum (0.46 including tubercles), 0.21 at collar,

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tubercles short, bluntly pointed. Disc slightly convex, shiny, covered with large,

flat-bottomed punctures, sharply defined by a rugule between them, rugules

with slight longitudinal orientation, no microreticulation in disc. Elytra 1.42

mm. long, 0.65 wide at the distinct humeri, 0.75 maximum, with a feeble, postbasal

transverse depression. Punctures sharply defined, moderately deep in basal half,

gradually smaller but distinct behind, 0.04 mm. from center to center. Setae

appressed, 0.20 mm., barely clearing punctures; tactile setae erect, 0.07 mm.,

moderately stout. Mesosternum strigulose, longitudinal medially to oblique

laterally, the strigules not anastamosing ; 0.10 wide laterally, leaving 0.07 of

mesepisterma exposed laterally. Fringe setae appressed, 0.07 mm. long anteriorly.

Mesepisterna with deep punctures at anterior margin, smooth and shiny behind,

fringe setae denser posterolaterally. Metasternum with longitudinal strigules,

smoother medianly; metepisternum smooth next to sternum, longitudinally strigu-

late laterally. Abdominal sternum 1 with few longitudinal strigules, micro-

reticulate, the rest with only microreticulation, setae appressed, sparse, 0.04 mm.

Sternum 7 essentially as in aequinoctialis. Sternum 8 not down-curved, concave

across base, with moderately deep U-shaped median notch, 0.28 mm. wide, 0.18

from one tip to the other, notch ca. 0.07 mm. deep. Ventral surface microreticulate,

edges of notch with some long setae. Apex of tergum 8 with punctulate and

pubescent zone, 0.26 mm. wide, 0.12 long, set off anteriorly by a sharp, transverse

ridge. Front femur 0.48 X 0.15 mm., front tibia 0.37 X 0.07, hind femur 0.52 X 0.17.

Aedeagus with tegmen gradually tapered (Fig. 5) ; gonopore armature essentially

as in other species, with spines in the internal sac near the gonopore armature

(Fig. 4) .

Female. — Similar to male, but with sternite 8 unmodified.

Holotype male: Albrook Forest Site, Fort Clayton, Canal Zone, Panama,

23/24 May 1968, black light trap, R. Hutton, Lot No. 200. 45 paratypes: 18 males,

25 females, same locality as holotype, 25 May, 1967 to 10 May, 1968, Hutton and

Llaurado. Individuals were collected every month except July and October.

52 same locality; 2 males, 5 females, 25/26 May 1967, Hutton & Llaurado. 1 male,

1/2 June 1967, Hutton & Llaurado. 1 female, 3/4 August 1967, Hutton &

Llaurado. 2 females, 7/8 September 1967, Hutton & Llaurado. 1 female, 28/29

September 1967, Hutton & Llaurado. 1 male, 2 females, 30 November/1 December

1967, Hutton & Llaurado. 1 female, 28/29 December 1967. 1 female, 19/20

January, Hutton & Llaurado. 2 males, 30/31 January 1968, R. Hutton. 1 male,

3 females, 23/24 February, R. Hutton. 1 female, 7/8 March 1968, R. Hutton. 2

males, 3 females, 21/22 March 1968, R. Hutton. 2 males, 1 female, 28/29 March

1968, R. Hutton. 3 males, 1 female, 4/5 April 1968, R. Hutton. 2 males, 10/11

April 1968, R. Hutton. 1 male, 2 females, 18/19 April 1968, R. Hutton. 1 male, 1

female, 9/10 May 1968, R. Hutton. 2 males, Las Cumbres, Canal Zone, 10

January/1 April 1974, H. Wolda. The holotype will be placed at the University

of Arkansas with half the paratypes. Two paratypes will be placed at the United

States National Museum and the remainder in the author’s collection.

My thanks are given to C. A. Triplehom for reviewing the paper.

Literature Cited

Werner, F. G. 1967. A revision of Acanthinus (Coleoptera: Anthicidae). IV.

Ann. Amer. Entomol. Soc., 60: 255-273.

Vol. 51, No. 2, April 1975

165

SCIENTIFIC NOTES

Additional Distribution Records for Nearctic Species of Ptenus Kirby

(Hymenoptera: Argidae: Sterictiphorinae) 7 — Smith (1970. Trans. Amer.

Entomol. Soc. 96: 79-101) revised the Nearctic species of Ptenus Kirby, recog-

nizing 2 previously named species and describing 10 new species. The collection

of the Department of Entomology at Texas A & M University (College Station)

contains several specimens of Ptenus spp. collected in Texas and Mexico, some

which extend the known distributions of several species into east-central and

southeastern Mexico. The purpose of this paper is to redefine the distributions

previously described by Smith, and to list additional collection records of some

of the Nearctic species.

Ptenus bicolor Smith was described from eastern and southeastern Texas, and

extreme northeastern Mexico (Matamoros). The known distribution of this

species is extended to include eastern Mexico south to southeastern Vera Cruz.

Additional records include: TEXAS: Cameron Co., Brownsville; San Patricio

Co., Welder Wildlife Refuge. MEXICO: Nuevo Leon, 9 mi. s. Monterrey; San

Luis Potosi, El Salto; Tamaulipas, 11.3 mi. n. Ciudad Victoria, and 8.5 mi. s.

Soto La Marina; Vera Cruz, 7 mi. n. e. Mata Espino. Collection dates range

from early June to mid-August.

Ptenus imus Smith was described from southern Texas (Laredo and Browns-

ville) and northeastern Mexico (Tampico). The known distribution of this

species is extended north along the Gulf of Mexico coast to Corpus Christi.

Additional records include: TEXAS: San Patricio Co., Corpus Christi St. Park,

25-VIII-1962.

Ptenus magnus Smith was described from western Texas, New Mexico and

Arizona. Additional records include: TEXAS: Jeff Davis Co., Madera Canyon,

w. of Ft. Davis, 9-VIII-1969.

Ptenus modestius Smith was described from Arizona, California, and Texas as

far east as Brownsville and Austin. The known distribution of this species is

extended north in Texas to Bosque Co., north along the Gulf of Mexico coast to

San Patricio Co., and south into Mexico to southwestern Tamaulipas (16 mi. n. e.

Jaumave). Additional records include: TEXAS: Bosque Co., 2 mi. w. Iredell;

Cameron Co., 3 mi. w. Brownsville; Gonzales Co., Palmetto St. Park; Presidio Co.,

2 mi. s. Shafter; Refugio Co., 8 mi. s. Woodsboro; San Patricio Co., 7 mi. n.

Sinton, and Mathis. MEXICO: Tamaulipas, 11.3 mi. n. Ciudad Victoria and 16.3

mi. n. e. Jaumave. Collection dates range from mid-March to mid-July.

Ptenus nigropectus (Norton) was known previously from “eastern and northern

Texas, east through Louisiana to Mississippi” (Smith, ibid.). Additional records

extend the known distribution of this species to include Texas, from the pan-

handle south to San Antonio, southeast to Brownsville, east through Louisiana

into Mississippi, and south into Mexico to northern Vera Cruz (18.6 mi. s.

Tampico). Additional records include: TEXAS: Bosque Co., 3 mi. w. Laguna

Park; Brazos Co., Minter Springs, and College Station; Burnet Co., Longhorn

Cavern St. Park; Kerr Co., Kerrville; Refugio Co., 8 mi. s. Woodsboro. MEXICO:

Nuevo Leon, 9 mi. s. Monterrey; Vera Cruz, 18.6 mi. s. Tampico. Collection dates

in Texas range from mid-April to early June. The Mexican specimens were

collected in mid-August.

1 Technical Article 11412, Texas Agricultural Experiment Station, College Station.

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The Pan-Pacific Entomologist

Ptenus parvus Smith was described from western Texas and Arizona. Additional

records include: TEXAS: Brewster Co., 43 mi. s. Alpine, 7-VI-1972; Presidio Co.,

3 mi. n. Presidio, l-IX-1966.

Ptenus texanus (Norton) is known from Texas and Mexico. Additional records

include: MEXICO: Coahuila, Puerta de la Goriona, el. 4900 ft., Sierra del Carmen,

13-VII-1938.

Ptenus vanus Smith was described from New Mexico, Arizona, and the bordering

Sierra del Carmen mountains of Mexico (Coahuila). The known distribution of

this species is extended south in Mexico to central Vera Cruz. Additional records

include: MEXICO: Vera Cruz, 4 mi. w. Conejos, 29-VI-1971.

Ptenus vargus Smith was described from 2 females taken at “port of entry” from

Mexico at Brownsville, Texas. The first specific records from Mexico are Nuevo

Leon, 7.5 mi. s. Monterrey and 9 mi. s. Monterrey, and San Luis Potosi, 13.4 mi.

w. El Naranjo. Collection dates range from mid-March to mid- August. The

proximity of these localities to the Mexican-United States border suggests that

P. vargus probably occurs also in southern Texas, as suggested by Smith (ibid.).

I wish to thank D. R. Smith (ARS, USDA, Washington, D. C.) for his sugges-

tions and for his determinations of many of the specimens reported in this paper. —

Harold N. Greenbaum 2 , Department of Entomology, Texas A & M University,

College Station, Texas 77843.

A Mountain-top Swarm of the Hemipteran Nysius raphanus in New

Mexico, with Notes on Other Insects. — Capulin Mountain National Monu-

ment, situated in extreme northeastern New Mexico, has as its central attraction

an extinct volcanic cinder cone. This is thinly forested with pinyon pine and

rises a thousand feet (summit is 8,215') above the surrounding treeless plain.

When Ray G. Martinez, Jr., Park Superintendent, kindly gave me a permit for

collecting insects there (May 9, 1974), he said that visitors had complained for

the past two weeks of vast swarms of small “flies” on the summit trail around the

crater rim. Near midday, at the summit parking lot, hikers descending from the

trail did indeed seem harassed. A short climb up the trail brought me into an

area where the air was filled with flying, small, hard-bodied insects that pelted

one’s face like grains in a sandstorm. This situation extended for about a

half-mile of the trail. The insects proved to be a species of lygaeid, identified as

Nysius raphanus Howard by Jon L. Herring of the USDA Systematic Laboratory

at Beltsville. Both females and males were present in about equal numbers. This

species has long been regarded as a synonym of N. ericae (Schilling), the False

chinch bug.

As the day progressed, the wind became brisk, and the Nysius sought shelter

in the small pinyons. A blow with a net handle on a tree produced a thick gray

cloud of insects that quickly drifted down wind; each pass of the net through

such a cloud captured a few hundred of the insects. Nysius was not seen near the

base of the mountain.

Near the highest point of the mountain, just down wind from the tip of a tall

2 Present Address : Department of Entomology and Nematology, University of Florida, Gainesville,

Florida 32611.

Vol. 51, No. 2, April 1975

167

pine, was an aggregation of 10 to 20 oestrid flies, which easily kept its position

in the wind. A specimen sent to R. J. Gagne (also of the Systematics Laboratory)

proved to be Cephenomyia pratti Hunter.

At mid-morning of the same day the long-faced black bee Andrena porterae

Cockerell was common on flowering bushes of Ribes at the base of the peak.

This is, as far as I know, the easternmost record for this Rocky Mountain species.

Probably Mt. Capulin’s chief claim to entomological fame is the satyrid but-

terfly Oeneis alberta capulinensis Brown 1970: 137 (J. New York Entomol.

Soc.), which at the time of its description had status as an isolated eastern

population of alberta. However, Brown predicted it would eventually be found in

the highlands to the north and west. In order to reduce possible collecting pres-

sure on the small Mt. Capulin colony, it would be well to point out here that I

observed alberta on May 10 in numbers on Johnson’s Mesa, about 10 mi. NNW of

Mt. Capulin, at an altitude of 7,600'. — LJ. N. Lanham, University of Colorado

Museum, Boulder.

Additional Notes on Chagas’ Trypanosome in California and Arizona. — -

Wood (1975, Exptl. Parasitol. In Press) indicated 25 locations in California for

recovery of Triatoma protracta protracta (Uhler) naturally infected with Try-

panosoma cruzi Chagas. Further unreported California localities for recovery of

Chagas’ trypanosome in T. p. protracta include the following: from Neotoma

houses in Carden Valley, El Dorado Co., 26 & 28 November 1963: 24 (4 $, 7

2, 8-5th and 5-4th instar nymphs) with 3 $ and 5 2 positive for T. cruzi, A. L.

Gladwill; at black light in Lytle Creek, San Bernardino Co., 12 September 1964:

4 (2 $ , 2 2 ), 1 2 positive, J. A. Robertson; 1 positive $ from a home in Yreka,

Siskiyou Co., 21 July 1967, forwarded from the State Department of Agriculture

by R. Hawthorne; 1 positive 2 from the living room couch in a home in the

Hollywood Hills, Hollywood, Los Angeles Co., 20 August 1967, R. DeRover, Jr.;

and 8 (2 $ , 3 2 and 3-5th instar nymphs), 2 2 and 3-5th positive 10 August

and 2 September 1974 from Neotoma houses in Wildwood Park, Thousand Oaks,

Ventura Co., S. F. Wood.

From 1964 through 1970, 28 T. p. protracta were reported to me from homes in

Beverly Hills (Benedict Canyon), Los Angeles County. Microscopic examination

of the feces of 3 ^ and 11 2 revealed 1 2 positive for T. cruzi from inside a

home 7 September 1968 and another 2 positive from an outside screen 8 July

1969. Thus, there are numerous reservoir mammal carriers of Chagas’ zoonosis

in the southern end of the Santa Monica Mountains.

During the summer of 1965, A. L. Gladwill, caretaker at the Griffith Park

Boys’ Camp in Los Angeles, California, collected 13 T. p. protracta (6 $ , 7

2 ) from the vicinity of his living quarters (Wood & Wood, 1967, Pacific Insects

9: 544, Fig. 5) from 26 August to 31 October and placed them, dead or alive,

in his home refrigerator (1-2°C). These conenose bugs were transferred 6 June

1966 to the author’s refrigerator and removed 6 July 1966 for examination of the

rectal contents for trypanosomes. One 2 with feeble leg movements when isolated

by Gladwill 4 September 1965 revealed 3 trypo- and 16 epi-mastigotes of T. cruzi

when examined 305 days after capture! One $ collected alive 30 August 1965

168

The Pan-Pacific Entomologist

from inside the home (Fig. 5, above the garage, loc. cit.) revealed 1 trypo- and 8

epi-mastigotes when examined 310 days after capture. Another 2 collected

on 28 August 1965 showed 12 trypo- and 8 epi-mastigotes 312 days after capture.

These are remarkably long survival times and show that freshly captured Triatoma

can be stored in a household refrigerator to preserve Trypanosoma cruzi in the

insect vector for at least 312 days.

Infection with T. cruzi for all California triatomes obtained by me through 1974

was 25.1% or 945 of 3,761 bugs. Most were alive when rectal contents were

examined.

From Arizona, between 1966 and 1971, 24 (12 $,12 2 ) Triatoma rubida uhleri

Neiva were received. Two $ were positive for T. cruzi of 22 collected in Tucson,

Pima Co., 14 June 1969 by G. Harwood. One of 2 2 was positive from Phoenix,

Maricopa Co., collected 12 June 1970 by F. Plettenberg.

The infection rate for all Arizona triatomes examined through 1974 was 6.6%

or 88 positive of 1,323 examined mostly from dead, dried specimens. — Sherwin F.

Wood, 614 W . Shenandoah St., Thousand Oaks, CA 91360.

Foraging Behavior of Agapostemon on Oenathera caespitosa in south-

eastern Wyoming (Hymenoptera : Halictidae). — Linsley, MacSwain and

Raven (1963, Univ. Calif. Pubis. Entomol. 33: 25-58) have provided information

on the Apoideaen visitors to eleven species of Oenothera in and around the Great

Basin. This note reports on a species previously unrecorded on Oenothera caes-

pitosa caespitosa Munz and describes its foraging behavior on the blossom.

Observations were made on two small populations of Oenothera caespitosa on

two consecutive mornings, May 26 and 27, 1974, from 5:30 to 9:30 a.m. The

weather was clear, warm and without wind. The area, a slightly rocky sandstone

patch surrounded by typical open short-grass prairie, is located about seven miles

S. S. E. of Laramie, Wyoming. As reported previously, (ibid.) the flowers at

this time of morning were devoid of nectar but did possess significant amounts

of pollen.

The first activity was recorded at 8:30 on May 26 and 8:20 on May 27 when

females of Agapostemon texanus began to appear on the flowers. A total of four

bees were recorded on May 26 and ten on May 27. Typically, a bee would

alight upon the proximal half of a petal and walk to the base of a filament. It

would then ascend the filament to the anthers where pollen collected by the fore-

legs was transferred to the scopa. The pollen grains, which are large, triangular

and inter-connected by viscin threads formed a rather loose and easily detachable

mass on the scopa. When finished with one anther the bee flew to the next fila-

ment and repeated the process. Usually, two or three anthers were collected

from on each flower. In no case did a bee forage on all four anthers of the same

flower. Analysis of the pollen taken from the scopa of three bees revealed only

Oenothera pollen. This flower constancy occurred despite the fact that several

other species were in bloom and contained available pollen. Females of Aga-

postemon texanus carrying full pollen loads have been collected on these other

species at other times. Of the fourteen recorded visitors none were seen con-

tacting the stigma at any time. Thus, Agapostemon texanus does not seem to be

Vol. 51, No. 2, April 1975

169

an important pollinator of Oenothera caespitosa and may be regarded as a pollen

thief.

Although evening observations were not conducted it is probable that this

flower is pollinated by nocturnal hawkmoths as previously reported (ibid.).

The stigmas of all plants examined contained noticeable amounts of pollen. The

early-morning oligoleges of the genus Andrena ( Onagrandrena ) found by Linsley,

et al. (ibid.) were not present on the study site. — V. J. Tepedino, Department of

Zoology and Physiology, University of Wyoming, Laramie, Wyoming 82071.

Insect Associates (Diptera: Chironomidae, Sphaeroceridae) of Darling-

tonia calif ornica (Sarraceniaceae) in California. — In June, 1974, I had the

opportunity to study specimens of Darlingtonia californica Torrey, the California

pitcher-plant, in Nevada County, California, near Willow Springs. The plants

were growing along the stream margins at 6500 feet elevation in a sphagnum bog

surrounded by pine-fir forest. In view of the presumably digestive nature of the

fluids contained in the pitchers (Street, H. E. and H. Opik. 1970. The Physiology

of Flowering Plants: Their Growth and Development. American Elsevier Pub-

lishing Co.: New York, N. Y. 263 pp.), it was of interest to find two species of

dipterous larvae living and developing in the upper layer of reddish fluid above

the insect debris in the pitchers. One species was a chironomid midge, apparently

Metriocnemas edwardsi Jones, the other a sphaerocerid in the genus Leptocera.

I am indebted to Dr. E. I. Schlinger of the University of California, Berkeley, for

his identification of the flies.

The chironomid larvae were abundant in 15-20 pitchers examined in the field.

They appear to be detritus feeders, but may be herbivores. The larvae stayed near

the top of the detritus. Jones found M. edwardsi larvae eating insect remains in

plants in Siskiyou County, California, and reported that in captivity specimens

pupated on the outside of the pitcher, on the basal portion of the leaf or on

adjacent moss (Jones, F. M. 1916. Entomol. News 27: 385-392). Jones was

convinced, by the presence of larvae in almost every suitable Darlingtonia leaf,

that the larvae were habitually and possibly exclusively associated with it. To my

knowledge, M. edwardsi has never been reported from any host other than D.

californica, nor from outside California, although the plant occurs in Oregon.

Because of the uniqueness of the habitat utilized, it seems quite unlikely that the

same species of chironomid would be found in other aquatic situations in the same

area. Although there are 17 described species of Metriocnemus in North America,

M. edwardsi is the only one as yet described from California. The rest are northern

in distribution.

The Leptocera larvae were more numerous than the chironomids. These crawl

up the walls of the pitchers, and pupate among the retrorse hairs found in

the lower portion of the lube. Adult Leptocera were also found in the pitchers.

Some are probably caught by spiders, such as the eriogonids found in some of

the atria at the tops of the plants, while others escape and colonize new pitchers.

Various species of Leptocera are commonly found among low plants in this type

of boggy area. This particular species has apparently been able to adapt completely

to the conditions in the pitchers, and is probably limited to Darlingtonia.

170

The Pan-Pacific Entomologist

It is interesting that the eastern genus of pitcher-plant, Sarracenia, has many

recorded dipterous associates, while Darlingtonia has but three. It is also interesting

that no mosquito larvae were found in any of the plants, since they are recorded

from Sarracenia. This study was supported in part by the NIH Training Grant,

Department of Entomological Sciences, University of California, Berkeley. — S. L.

Szerlip, Division of Entomology and Parasitology, University of California,

Berkeley, 94720.

Identification of the damsel bugs, Nabis alternatus Parshley and N.

americoferus Carayon (Heteroptera : Nabidae).— Harris (1928, Entomol. Am.

9: 1-97) records 6 species of Nabidae from California; 3 of these are extremely

rare, and known from only a few specimens. Harris stated that of the 3 common

species, N. ferus (L.) was European in origin and had been introduced and be-

came widely distributed in the U.S. while Pagasa fusca (Stein) and Nabis alter-

natus Parshley were native. However, extensive genitalic studies of North Ameri-

can and European specimens of “/V. ferus ” (Carayon, 1961, Bull. Mus. Nat’l.

Hist. Nat. Paris (2) 33: 183-96) have shown the American form, now N. americo-

ferus Carayon, to be distinct from N. ferus , which is now thought not to occur in

California.

Development of a couplet to separate the two species of Nabis resulted from

difficulties encountered in rapidly identifying California Nabidae during studies

on predatory Hemiptera in Sacramento Valley alfalfa hay fields. Identification

using Harris’s monograph or the keys of Werner and Butler (1957, Ariz. Agr.

Exp. Sta. Tech. Bull. 133: 1-12) proved impractical due to the very similar

overall appearance and variable color patterns in our 2 common species of Nabis.

Also, we would point out that keys and descriptions of il N. ferus ” in both taxo-

nomic works apply to N. americoferus and not to the true European N. ferus.

The following couplet was constructed using specimens of Nabidae subsampled

from the predation study and identified by experienced Hemipterists. Its ac-

curacy was then confirmed by testing it on identified museum specimens from a

wide geographic range. The color pattern of the scutellum is best observed when

the specimen is in alcohol. All characters used in the couplet refer to both sexes

unless otherwise indicated.

Couplet to separate Nabis alternatus and Nabis americoferus.

— Scutellum dark brown with a pale yellow “V” shaped area extending to the

posterior margin (Fig. 2) or scutellum yellow except for brownish areas cen-

trally and at the anterior lateral margins (Fig. 1) ; male clasper (Fig. 5) wide

(0.20-0.28 mm) and long (0.52-0.60 mm) ; body length 7.0-9 mm

N. americoferus Carayon

— Scutellum dark brown with two oval yellowish areas, each located mid-

laterally (Fig. 3) ; male clasper (Fig. 4) narrow (0.11-0.16 mm) and

short (0.40-0.44 mm) ; body length 6.4^8. 1 mm Nabis alternatus Parshley

Vol. 51, No. 2, April 1975

171

Figs. 1-5. Structures used in separating species of Nabis; 1-3, scutellum;

4-5, lateral view of male genitalia, showing claspers. Figs. 1, 2, 5, N. americoferus.

Figs. 3, 4, N. alternatus.

We are grateful to Drs. Peter Ashlock, Richard Bohart, Jon Herring, John

Lattin and Mr. Tim Vargas for their assistance in various aspects of this project.—

J. H. Benedict and W. R. Cothran, Department of Entomology, University of

California, Davis, California 95616.

172

The Pan-Pacific Entomologist

BOOK REVIEWS

The Insects of the Los Angeles Basin. Charles L. Hogue, x -f- 173 pp. incl. 2

color plates, 250 figs. ; Los Angeles County Museum, Sci. Ser. 27, 1974. $6.00,

8 X 10“ paperback.

This fine little book is a pictoral guide to the common and conspicuous insects

and related arthropods occurring in the greater Los Angeles area. Because these

bugs are those most commonly seen in man’s most heavily populated region

of western North America, their numbers include many homodynamic kinds with

a disproportionate representation of “weedy,” introduced species and house and

garden insects. Therefore, though the number of species treated is not large

(about 200 are discussed and another 75 mentioned), most are widespread

species, so that the book will prove useful throughout the Pacific States. Virtually

all species discussed are illustrated, either by excellent photographs or shaded

drawings, many taken from technical literature.

Taxonomically, the coverage is uneven, geared to a level of information that

should answer questions of a layman. Thus orders or families of small insects

are characterized by group only, while a few to many species are detailed for

larger kinds like dragonflies and butterflies as well as for insects affecting man,

such as lice and mosquitoes. The Lepidoptera are given top billing, with more

than 60 species shown. Emphasis is also directed to household problems, with

sections on pantry moths and beetles and to housing insects alive, with instruc-

tions for keeping mantids and ant colonies in captivity and raising caterpillars.

A representative sample of scorpions, spiders, mites, ticks and other terrestrial

arthropods is also discussed, again with emphasis on those of potential medical

importance.

Introductory sections include discussions of structure and function, making a

collection, insect classification, and habitats around Los Angeles. These parts and

the text on individual orders and families are accompanied by literature references,

and there is a general section referring the reader to other sources of information,

both special topics and general ones, such as children’s books.

Common names of each species are accompanied by parenthetical mention of

the scientific names (without author), and the text is remarkably free from errors,

both typographical and technical. The color plates add aesthetic value but are

not essential, being habitat photographs of 16 species, most of which have been

reproduced in black and white in the text.

Although the price seems high considering the audience, this is the best popular

manual available to the public that deals with west coast insects, especially those

of urban areas. — J. A. Powell, Division of Entomology and Parasitology, Uni-

versity of California, Berkeley 94720.

The Moths of America North of Mexico: Pyraloidea and Gelechioidea. This

ambitious project which proposes to describe and illustrate all of the 10,000 T

species in our fauna has now published the first two fascicles dealing with the

so-called microlepidoptera. These follow a somewhat more formal format with

less anecdotal digression of popular manual style that was characteristic of

the several earlier issues treating sphingids, saturniids, and other hombycoids.

Vol. 51, No. 2, April 1975

173

Fascicle 13.1: Pyraloidea, Pyralidae (Part), by Eugene Munroe; published

in 3 parts, A (Scopariinae and Nymphulinae) (1972), B (Odontiinae and

Glaphyriinae) (1972), and C (Evergestiinae and plates to all 3 parts) (1973) ;

304 -)- xx pp., 13 color and 10 bl. & wh. plates [price about $82.00, non-subscrip-

tion].

This is half of the first of 4 fascicles planned to cover the pyraloids, probably

the second largest superfamily of Lepidoptera, with more than 1,500 species in

North America. The introduction contains the first comprehensive review

of higher classification in the superfamily and in the Pyralidae to have appeared

in many years.

These 3 parts enumerate some 227 species of which nearly 20% are previously

undescribed, despite the fact that the Odontiinae was monographed by Munroe in

1960 and various other taxa have been recently described. Virtually all species

are illustrated in the superb color photographs, and genitalia of new species are

shown by microphotographs. Most of the color plates are reproduced at twice

natural size, while the somewhat larger moths of the Evergestiinae and some

Scopariinae are shown at natural size.

The text is very thorough, more resembling a taxonomic revision than a general

manual. It includes lengthy synonymies and rather detailed descriptions and full

data citations for the new taxa, which in addition to the new species include 7 new

genera and more than 20 new subspecies, encouraging nomenclatural recognition

of geographical variation that American microlepidopterists have been relatively

free from in the past.

This fascicle suffers from the paradox that the earlier macro-moth volumes

instigated: a decision to treat the fauna as though these were taxonomic re-

visions, incorporating undescribed material, yet using mainly a few large eastern

collections as sources of raw data. The former concept results in uneven treatment

between taxa, greatly expanded text length, delays in completion and encourages

high costs; while the latter attitude results in failure to incorporate all up to

date knowledge, particularly that concerning western species. Munroe has used

some western collections, in contrast to earlier fascicles where perhaps a con-

vincing argument could be mustered supported by the much better fieldwork

coverage by the industrious collectors of sphinges and bombyces. However, the

use of western material in pyralids is sporadic both in taxonomic spectrum and

in recency of solicitation, so that there are a number of statements concerning

geographical distributions and lack of biological information that were out of date

when written.

Fascicle 6.2: Gelechioidea, Oecophoridae, by Ronald W. Hodges; 142 -f- x pp.,

7 color and 1 bl. & wh. plates (1974) [price about $57.00, non-subscription].

This volume is of general interest in two ways. First it proposes a higher

classification for Oecophoridae, based on review of the world fauna, and this

is one of the largest families of microlepidoptera with more than 4,000 described

species. Secondly, this is the first family of microlepidoptera to have been re-

viewed following an earlier taxonomic revision based on genital characters, namely

Clarke’s treatment in 1941. Thus this is our first chance to see whether an as-

ymptote in faunal size is being approached in Nearctic microlepidoptera, and it

appears that it is. Hodges defines some 147 species, and only about 10% are

174

The Pan-Pacific Entomologist

previously undescribed, while a like proportion of names are considered new

synonyms. Therefore the size of this family is only slightly enlarged, and con-

temporary microlepidopterists can be encouraged to know that their present

treatments are near the crest, at least in non leafminer taxa.

With the exception of new taxa, the oecophorid text is quite terse, with concise

morphological and distributional summaries and abbreviated synonymies that often

do not include previous generic assignments so that new combinations appear to

revise the original combinations only. On the other hand, some superfluous items

are incorporated, such as citation of plant common names, a lengthy quote from

previous literature on page 44, and proposal of unique tribes (pp. 93, 136) in

subfamilies that contain no other tribes. In general the coverage of biological,

phenological, and geographical knowledge is excellent, although there are a few

errors and omissions in biological information.

This is the first group in the series to include quite small moths, and the color

plates are unbelievably good, showing the adults at twice natural size for all

genera except a few like Decantha which are magnified 4 times. In addition

there are excellent drawings by Elaine Hodges illustrating genitalia of new

species and some previously described ones that were evidently deemed necessary

for comparison purposes.

An innovation towards economy has been made without loss of clarity, with

the use of smaller type organized in double column format, so that each page

potentially carries about 17% more print than in predecessor volumes. However,

the text incorporates 32 generously spaced, large figures which are more detailed

than necessary so that greater reduction was not warranted in favor of details

in non-taxonomic features such as shading and setae. The figures comprise about

18% of the text, much more than would have been necessary had they been less

detailed drawings originally and were they aggregated as full page groups.

These two fine treatments will be musts for serious students of the pyralids

and oecophorids, but considering the prices they are hardly manuals for the general

lepidopterist. It is apparent that the dual goal of a hybrid monograph that will

be useful and available to laymen, yet be comprehensive and prepared by meticulous

professionals is impossible in the face of today’s production costs. — J. A. Powell,

Division of Entomology and Parasitology, University of California, Berkeley,

94720.

Vol. 51, No. 2, April 1975

175

RECENT LITERATURE

A Revision of the Scarabaeine Dung Beetles of Australia II. Tribe Scarabaeini.

E. G. Matthews, Australian lournal of Zoology, Suppl. Ser. No. 24, pp. 1-211.

The highly specialized dung beetle fauna of Australia has received admirable

monographic treatment in the first two parts of this work. Part II, dealing with the

Canthonina, describes and keys 94 species in 16 genera, of which three genera and

52 species are considered new. Discussions of food, behavior, and biogeographic

relationships, including the significance of wing reduction in many Australian

species, supplement the keys and descriptions of genera and species. One species

from each genus is portrayed in excellent half tone illustrations, and details of

important taxonomic features are provided by over 300 line drawings. — Editor.

Los Tipos de Insectos de la Coleccion Luis E. Pena g. Luis E. Pena. Bol.

Soc. Biol. Conception, 47:259-282. 1974.

The author lists 114 species for which he holds the holotypes. Of these, 84 species

are Tenebrionidae, and 23 species are from other families of Coleoptera, especially

Cerambycidae. The Pena collection also contains a few holotypes of Lepidoptera,

Hemiptera and Orthoptera. Paratype material represents an additional 151 species,

mostly Tenebrionidae, with moderate representation of other families of beetles and

of Lepidoptera. Although small compared to the holdings of major museums, the

Pena collection probably comprises one of the most important assemblages of in-

sects, particularly Coleoptera, from southern South America. — Editor.

A Systematic Monograph of New World Ethmiid Moths (Lepidoptera:

Gelechioidea) . J. A. Powell. Smithson. Contr. Zool., no. 120, iv -f- 302 pp.

1973. ($3.85, U. S. Government Printing Office, Washington, D. C. 20402.)

This exhaustive treatment of a previously confusing family of moths is truly

monographic in scope. Both sexes, when available, of each of the 133 new world

species are described. Clear line drawings of genitalia are provided for nearly all

species, and 192 photographs arranged in 16 plates, illustrate the adult moths.

Although genitalia, especially in males, provide excellent differentiating characters

which were important in defining species groups, the keys to genera and species are

based on external characteristics, making this work potentially useful to a much

larger audience. Immatures are not described, but analysis of larval and pupal

characters appears in tables and discussions. A large amount of biological in-

formation is integrated with the systematics portion of the text, with separate

discussions on host plant relationships, phenological patterns, and biogeographic

distribution. Taxonomic conclusions are derived in part from numerical analysis

of 42 characters, with results expressed as phenograms and Primnets. The text is

indexed by general subject, as well as host plant name. — Editor.

176

The Pan-Pacific Entomologist

Butterflies of the World. H. L. Lewis, with a Foreword by J. M. Chalmess-

Hunt. Follett Publishing Company, Chicago, Illinois, 1973. XVI -f- 312 pp.,

208 color plates.

Illustrates about 70 percent of the species of butterflies, representing nearly all

genera. Organized by taxa within biogeographic regions.

The Ecology of Insect Populations in Theory and Practice. L. R. Clark, R.

Hughes and R. F. Morris. Halstead Press, John Wiley and Sons, Inc., New

York, New York, 1974. 248 pp., paper, $6.75.

Insects and Diseases. Keith Snow. Halstead Press, John Wiley and Sons, Inc.,

New York, New York, 1974. 208 pp., $9.50.

Vol. 51, No. 2, April 1975

177

OBITUARY

Donald D. Penny was born September 22, 1894 at Los Gatos, California and

died February 8, 1975 in Aptos, California at the age of 81. He entered the

University of California in 1914 and received a B.S. degree in Entomology in

1917. While at the University, he was greatly influenced by Professors C. W.

Woodworth and E. 0. Essig. As a student, he conducted extensive investigations

on Aleyrodidae, an interest which continued after graduation. His types were

presented to the California Academy of Sciences.

On completing college, Penny went to Sacramento to work in the state insec-

tary under the supervision of Harry S. Smith. His primary responsibility was

the production of ladybird beetles. In 1919, at the urging of Smith, Penny took

and passed the examination for Horticultural Commissioner for Santa Cruz County.

While there he became very much interested in oil sprays. In cooperation with

W. H. Volck he played an important role in the development and use of the new

phytonomic summer oil sprays. These relatively safe oil sprays were rapidly

accepted throughout the developed agricultural world and marked an important

advancement in pest control.

In September 1923, Don, as he was known to friends and associates, joined

Cal Spray Chemical (Ortho Division of Chevron Chemical Company) where he

remained until he retired on October 1, 1959. He served the company in a number

of capacities which included marketing, research and development, and formula-

tion of products. He introduced Chevron products into Australia and New Zealand

in 1927 and 1928 and into Hawaii in 1927. At the time of his retirement, he was

Assistant to the Manager for Research of the company.

Penny served a term as President of the Entomological Society of Southern

California and was an Emeritus member of the Entomological Society of America.

Don was a most friendly and cooperative person. His principal hobby during

his active career revolved around applied entomology. After retirement he devoted

much time to his garden. He is survived by his wife, Dorothy, who lives in Aptos,

California, and a son, William, who resides in Oakland, California. — Leo R.

Gardner, retired, Chevron Chemical Co., Ortho Division and A. E. Michelbacher,

Emeritus Professor of Entomology, University of California, Berkeley, 94720.

178

The Pan-Pacific Entomologist

ZOOLOGICAL NOMENCLATURE

Announcement A. (n.s.) 95

Required six months’ notice is given of the possible use of plenary powers by

the International Commission of Zoological Nomenclature in connection with the

following names listed by case number: (see Bull. zool. Nom. 32, part 1, 27th

March 1975).

1079. Forcipomyia Meigen, 1818 (Insecta, Diptera) : designation of type-species.

1892. Ruling on the stem of the family-group name based on Sphaerius Waltl,

1838 (Insecta, Coleoptera).

Comments should be sent in duplicate, citing case number, to the Secretary,

International Commission on Zoological Nomenclature, c/o British Museum (Nat-

ural History) , Cromwell Road, London SW7 5BD, England. Those received early

enough will be published in the Bulletin of Zoological Nomenclature. — Margaret

Green, Scientific Assistant.