Vol. 55 APRIL 1979 No. 2

THE

PAN-PACIFIC ENTOMOLOGIST

MINTZER—Colony Foundation and Pleometrosis in Camponotus (Hymenoptera: For-

micidae)

PARKER—Alfalfa Leafcutter Bee—Reducing Parasitism of Loose Cells During Incu-

bation (Hymenoptera: Megachilidae)

PAPP and JOHNSON—Origins of Psyllid Fallout in the Central Sierra Nevada of

California (Homoptera)

EDMONDS—A New Species of Phanaeus From Mexico (Coleoptera: Scarabaei-

dae)

SKILES—A New Subspecies of Poliaenus nuevoleonis Chemsak and Linsley From

Southern Arizona (Coleoptera: Cerambycidae)

SIMS et al.—Genetic Confirmation of the Specific Status of the Speyeria adiaste Group

in California (Lepidoptera: Nymphalidae)

ROCKWELL—Enmigration Response Behavior: II: The Responses of Drosophila busckii

(Diptera: Drosophilidae)

TILLEY—Some Larvae of Orthocladiinae, Chironomidae, From Brooks Range, Alaska

With Provisional Key (Diptera)

CHANDLER—A New Species of Tanarthrus From California (Coleoptera: Anthici-

dae)

ASHLOCK—A New Eremocoris From California With a Key to North American Gen-

era of Drymini (Hemiptera-Heteroptera: Lygaeidae)

FENDER—A New California Species of Podabrus (Coleoptera: Cantharidae)

SCIENTIFIC NOTES

SAN FRANCISCO, CALIFORNIA «+ 1979

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

106

155

The Pan-Pacific Entomologist

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OFFICERS FOR 1979

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Title of Publication: The Pan-Pacific Entomologist.

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The Pan-Pacific Entomologist (ISSN 0031-0603)

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PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 81-89

COLONY FOUNDATION AND PLEOMETROSIS IN CAMPONOTUS

(HYMENOPTERA: FORMICIDAE)

ALEX MINTZER

Museum of Zool. and Division of Biol. Sci., Univ. of Michigan,

Ann Arbor, 48109

The period of reproductive activity and colony foundation is obviously

crucial in the life cycle of ants. In most ant species, new colonies are es-

tablished by independent females after a mating flight. The reproductives

are subject to predation and the vagaries of weather during and after the

mating flight, and the number of suitable sites for colony foundation avail-

able to individual dispersing females may be a limiting factor. The first

worker brood is usually reared by the isolated female without any additional

assistance. The females might increase the food reserves available for brood

rearing by foraging outside the incipient nest, thereby increasing the indi-

vidual size or number of workers in the first brood, but foraging behavior

exposes the female to additional predators. Foraging by colony foundresses

occurs only in a small number of ant taxa studied, notably in some Ponerinae

and the Australian genus Myrmecia (Wheeler, 1933; Haskins, 1955), in some

attines (Weber, 1972) and in acacia ants belonging to the genus Pseudo-

myrmex (Janzen, 1966). In most ant species, females do not forage and must

rear a worker brood on a limited supply of food reserves. However, con-

specific females may cooperate in jointly rearing a first brood; this is termed

pleometrosis. Holldobler and Wilson (1977) provide a recent review of pleo-

metrosis in ants.

Pricer (1908) and Eidmann (1926) provide accounts of colony foundation

in two species of Camponotus in the eastern United States and Europe.

Their studies show that females of Camponotus ligniperda and C. hercu-

leanus found colonies independently in a manner similar to most other ants.

They do not seek food or capture prey outside their incipient nest chamber

during colony foundation. Camponotus females apparently do not found

colonies through temporary social parasitism involving other ant species,

unlike some ant genera (Wilson, 1971).

This paper describes colony foundation in seven Camponotus species

(representing four subgenera) in the western United States. The effect of an

initial supplementary feeding on colony foundation success was investigated

using two species of Camponotus. Five cases of successful colony foun-

dation by pairs of cooperating females of C. vicinus are described.

82 PAN-PACIFIC ENTOMOLOGIST

Table 1. Collection data for ants used in study of colony foundation.

No. fe-

Species males Locality Date

Camponotus (Tanaemyrmex) 1 Tehachapi Mtn. Park, Los July 8, 1973

vicinus Mayr Angeles Co., California

C. vicinus 4 Iowa Hill, Placer Co., May 1974

California

C. vicinus 7 Iowa Hill May 1975

C. vicinus 14 Iowa Hill May 2, 1976

C. (Tanaemyrmex) 1 Brawley, Imperial Co., July 24, 1974

festinatus (Buckley) California

C. (Camponotus) modoc 1 Iowa Hill, Placer Co., May 1974

Wheeler California

C. (Camponotus) 4 Iowa Hill May 1974

laevigatus F. Smith

C. laevigatus 5 Iowa Hill May 1975

C. laevigatus 1 Flagstaff, Coconino Co., May 10, 1974

Arizona

C. (Myrmentoma) 1 Tustin, Orange Co., April 24, 1972

clarithorax Emery California

C. (Myrmentoma) rasilis 2 Victoria, Victoria Co., May 8, 1974

Wheeler Texas

C. (Myrmentoma) sp. 1 Pinery Valley, Chiricahua Aug. 21, 1973

Mtns., Cochise Co., Arizona

C. (Myrmobrachys) 4 Brownsville, Cameron Co., June 6-7, 1977

planatus Roger Texas

Materials and Methods

The dealate females were collected in California, Arizona, and Texas.

Most were taken in the open, presumably at the conclusion of mating flights.

Several females were found when the incipient nest chamber was broken

open during routing collecting activity. Table 1 lists the collection data for

ants used in this study.

Most of the females were housed in 35 mm plastic petri dishes lined with

filter paper. Water was supplied periodically to all ants by wetting a cotton

ball inserted through the plastic dish lids. The C. planatus females were

kept in glass shell vials with cork stoppers. The dishes or vials containing

founding females were kept together under a thin-wall cardboard box lid or

in incubators to exclude light. Air temperatures ranged from 22 to 29°C

during colony foundation.

Females collected in 1972, 1973, and 1974 were offered honeywater soon

after capture, or as the first larvae neared maturity. In 1975, the effect of

this initial feeding was tested using C. vicinus and C. laevigatus. Three

females of each species were denied access to honeywater until the first

workers eclosed. Two other females of each species received an initial ad

VOLUME 55, NUMBER 2 83

libitum feeding of honeywater shortly after capture. Honeywater was given

to all incipient colonies within two days after eclosure of the first worker,

and the cotton ball was removed from the petri dish lid to allow ants access

to the exterior. Honeywater and insects were supplied on a regular basis

once the worker offspring began foraging. Muscid and Calliphorid flies were

accepted by most species, and Drosophila flies were taken by C. clarithorax

and C. (Myrmentoma) sp. Mealworms (Tenebrio molitor) were accepted by

C. vicinus, which rejected Drosophila. The ants and brood later emigrated

or were transferred to wood or plaster nests with glass tops.

In 1975, two compatible C. vicinus females were placed in one 35 mm

plastic petri dish. In 1976, four pairs of compatible C. vicinus females were

placed in 35 and 60 mm petri dishes (two pairs in each dish size). All dishes

and vials with founding females were examined several times each week to

monitor the development of the first brood.

Results and Observations

I. Colony Foundation and Effects of Initial Feeding

Nearly all of the dealate females used in this study reared workers to

maturity, and required 48 to 74 days to rear the first brood (see Table 2).

During colony foundation most of the ants remained motionless for long

periods of time (e.g., Fig. 1 is a multiple second time exposure). The females

produced a clutch of eggs over a period of several days following installation

in the culture containers. The size of this initial egg clutch varied greatly.

In the C. laevigatus and C. vicinus series, the maximum size of the first

egg clutch ranged from 9 to 16 for single queen replicates. In the C. planatus

series, all females produced six eggs or less. The eggs of species in the

subgenera Camponotus, Myrmentoma, and Myrmobrachys were elongate

and cylindrical in shape. The eggs of the Tanaemyrmex species were broad-

ly oval rather than elongate-cylindrical in shape. The eggs of some species

showed striking changes in appearance during embryonic development, due

to internal cell movement or migration and changes in the chorion, and eggs

of different age could be easily distinguished.

First brood larval growth was usually uninterrupted and rapid. However,

the last larvae to hatch in a brood were often ‘held back’ in the first instar,

and usually did not complete development until the following year. The first

instar larva is a common resting stage during periods of overwintering or

food shortages in Camponotus (Mintzer, unpubl. observation; Holldobler,

1961). In 1974, the C. modoc female and three of the C. laevigatus females

tore loose pieces of filter paper to cover larvae spinning cocoons. Pupal

mortality and abortive adult eclosure were uncommon.

The genus Camponotus is characterized by a polymorphic worker caste.

The workers in the first broods always belonged to the minor subcaste, and

PAN-PACIFIC ENTOMOLOGIST

Table 2. Development of first brood during colony foundation.

Species

. modoc 1974

. laevigatus 1974

. laevigatus 1975

. vicinus 1973

. vicinus 1974

. vicinus 1975

. festinatus 1974

. Clarithorax 1972

. (Myrmentoma) sp.

. planatus 1977

PEVLILP TE LISS PES Be

No. of

repli-

cates

Ree KSB DAK Ne

Duration of immature stages (days)

First First

First egg larva to cocoon to

to first cocoon first

larva spinning worker

21 13 21

Zo 14 26

20 12 20

22 NR NR

24 14 28

21 13 23

NR NR NR

33 NR NR

28 2 14 26

28 12 17

Total

64 (58-69)

55 (48-70)

66 (62-70)

58 (54-63)

68 + 2

57 (54-58)

showed little size variation. The number of workers produced in the first

brood or in the first season varied widely between species, but was often

quite similar within a replicate series (e.g., C. vicinus in 1974). As expected,

the ants aided in tending the brood, but the queen did not abruptly relinquish

Table 3.

Species/female

C. vicinus

BJX

Treatment

fed

unfed

First

egg to

first

larva

20-21

25-26

18-20

17-18

Duration of immature stages (days)

First First

larva to cocoon to

cocoon first

spinning adult

12-14 23-25

13-14 23

12-13 22

11-12 22-24

14-15 22-23

Effects of an initial honeywater feeding on colony foundation.

54-55

57-59

55-56

Colonies founded by four females receiving initial honeywater feeding in 1974

AGX

AHX

AIX

C. laevigatus

fed

unfed

20-21

18-19

20-21

41a

7-8

10-11

17-18

eu)

18-20

15-21

47-50

66-71

Colonies founded by four females receiving initial honeywater feeding in 1974

No. of

workers +

cocoons in

first brood

(August 15)

—_— —

sf ates |

7, 3,9; 9

NOC

2, 4, 6, 6

a Ege consumption by this female probably resulted in the longer egg stage duration ob-

served.

VOLUME 55, NUMBER 2 85

Figs. 1-2. Fig. 1. Colony founding female of Camponotus vicinus with first larval brood.

Fig. 2. Cooperating pair of C. vicinus females with first brood of larvae and cocoons.

brood care after the workers eclosed, and often participated in this activity

throughout the first season. Workers in most colonies began foraging within

25 days following eclosure. The foraging workers were generally timid and

retreated from any disturbance, although some ants tended to investigate

the foraging arena during and after any manipulation, whether or not food

was offered. Worker recruitment was occasionally observed in these small

colonies during emigration or when honey was offered.

86 PAN-PACIFIC ENTOMOLOGIST

Table 4. First brood egg and worker production by single and paired females (1975, 1976).

Maximum No. of workers

egg no. and cocoons

Single female Range 9-16 1-11

Replicates Mean 12.2 6.8

n=9 Std. dev. 2.3 303

Paired female Range 17-30 11-17

Replicates Mean 20.4 13.8

n=5 Std. dev. +5 2.6

Table 3 shows the results of the experiment testing the effect of initial

feeding on colony foundation. One of these ants (female BF) was infertile

and her eggs failed to hatch, even though she had received honeywater. All

other ants reared brood to maturity. Egg eating was not observed among

females which had received honeywater, although unaccounted loss of eggs

was noted in one case (female AE). Egg eating by two unfed ants (AIX and

BJX) was observed, and one egg was consumed in each case. One pupa

disappeared and was presumably consumed by an unfed C. laevigatus fe-

male (AGX). The maximum size of the egg clutch was similar in the two

groups, as was the brood development time and the number of workers in

the first brood.

II. Cooperative Colony Foundation by Two Females

In 1975, a pair of C. vicinus females successfully reared a brood of 11

workers to maturity. The two ants maintained a single egg pile; the maxi-

mum size of this clutch was 21 eggs. In 1976, four pairs of C. vicinius

successfully reared worker broods to maturity. Several pairing attempts

were required to find compatible females for this experiment, as some com-

binations of the ants available led to fighting. One female lost three legs in

such fights after initial pairing. However, paired females that were compat-

ible in the initial stages during egg laying remained compatible for the entire

period of colony foundation. Oral food exchange between females and al-

logrooming was observed, and ‘dominance’ or aversive behavior was very

subtle or absent. Occasionally one female would climb partially on top of

the second female, but biting was never observed (cf. Polistes; West-Eber-

hard, 1969) and the putative dominant-submissive roles were often reversed

when the behavior was observed again. On May 21, an accident resulted in

the loss of all eggs in the dishes, but all of the ants produced a second

clutch. The paired females produced a single egg clutch, which was larger

(p < 0.01, Behrens-Fisher t test) than the egg clutches of single foundress

queens. The time period involved and the details of brood development

were similar in single and paired-female series, but paired females produced

VOLUME 55, NUMBER 2 87

more workers than single females (p < .005, Behrens-Fisher t test). Females

in 60 mm dishes did not rear more workers than did those in 35 mm dishes.

Three of the four pairs of females founding colonies in 1976 remained

compatible through August 1977. One pair was separated after fighting be-

tween the females began in March 1977. The largest colony with two females

had 42 worker offspring in early August 1977.

Discussion

The high proportion of females which successfully founded colonies is

noteworthy. The culture dishes and vials satisfied a requirement for a small

closed cavity during colony foundation, without contamination by patho-

genic bacteria and fungi. The four subgenera of Camponotus surveyed in

the study occupy different habitats in nature. Ants in the subgenus Tanae-

myrmex nest in the soil in the western and southern United States. Campo-

notus laevigatus and C. modoc occur in forested mountain areas in western

North America, and nest in large pieces of wood. The ants in the subgenus

Myrmentoma are largely arboreal and are distributed throughout the United

States and southern Canada. The subgenus Myrmobrachys is the dominant

group of arboreal Camponotus in neotropical habitats, and occurs in the

United States in southern Arizona, Texas, and Florida (Creighton, 1950).

Conditions in the laboratory probably minimized the brood development

time. Water was supplied on a regular basis, and temperature extremes were

avoided during colony foundation. Under these conditions, supplemental

honey feeding had little or no effect on the first brood development time,

and did not appear to increase the size of this brood. Brood cannot be

satisfactorily reared on honeywater alone. However, the ants could use the

sugars present in honey for some of their own metabolic needs, and thereby

free more protein and lipid reserves for use in brood rearing. Such a facil-

itatory effect has been demonstrated in Myrmica (Brian, 1973). Under less

satisfactory culture conditions, beneficial effects of the initial feeding might

become apparent. Brood consumption was uncommon in the laboratory,

but was more frequent among unfed females. In the field, lower average or

fluctuating temperatures would probably prolong or interrupt brood devel-

opment, especially in areas inhabited by C. modoc and C. laevigatus.

The successful foundation of colonies by pairs of cooperating Campo-

notus is significant. Only one other account (Stumper, 1962) of pleometrotic

colony foundation by Camponotus females is available. Stumper’s colony

foundation experiments with C. (Camponotus) vagus Scopoli were plagued

by a high level of brood cannibalism and none of his single-female replicates

reared workers to maturity. However, his pair of compatible females was

successful at rearing workers, and produced more brood than the single

females. Paired or grouped females of Lasius flavus (Fabricius) rear more

88 PAN-PACIFIC ENTOMOLOGIST

brood to maturity in a shorter period of time, and have a higher survivorship

rate than single females (Waloff, 1957). Pairs of compatible C. vicinus found-

ing females also appear to rear larger broods than single females. It is not

clear whether both females are contributing brood during colony foundation.

Even if only one individual in a pair contributes eggs to the clutch, she can

produce a correspondingly larger number of eggs, to be tended by two

founding females. However, the origin of these eggs probably will not in-

fluence the ultimate reproductive success of cooperating females. The re-

productive success of each female is determined by the number of repro-

ductive females and males contributed by each when the colony reaches

maturity, and by colony survivorship functions. The increased number of

workers produced by paired females may have a major positive impact on

survivorship of incipient polygynous colonies. Unrelated females may be

expected to cooperate if their expected reproductive success is increased

above the level for haplometrotic colony foundation. According to kin-se-

lection theory, related females may be expected to cooperate in colony

foundation under some conditions when unrelated females would not, as

long as the inclusive fitnesses of the participating ants are increased (Ham-

ilton, 1964). All of the C. vicinus females involved were collected in a single

locality and it is possible that some are sisters. The behavior of the ants

during pairing attempts suggests that they are discriminating on some basis,

possibly residual odor cues from the parent colony.

Records of mature polygynous colonies of Camponotus in nature are

uncommon. Holldobler (1962) found some in C. ligniperda, where the fe-

males were hostile to each other and maintained separate territories in large

colonies. In a series of experiments, he found that newly fertilized dealate

females were usually incompatible, and those females which did cooperate

failed to rear a first brood to maturity. Holldobler concluded that polygynous

colonies of this ant arise by adoption of females or mating within the colony.

In August 1973, the author excavated a large colony of C. (Tanaemyrmex)

sansabeanus (Buckley) with two dealate females near the Southwest Re-

search Station of the American Museum of Natural History at Portal, Ari-

zona. These ants have shown no aggressive behavior towards each other in

the four years after collection of this colony. Both contribute eggs and egg

eating by the queens has not been observed. It seems likely that polygynous

colonies of this type could occasionally originate through cooperation be-

tween founding females.

Conclusion

Under laboratory conditions, Camponotus females required 48 to 74 days

to rear a worker brood to maturity. The development of this brood was

uninterrupted and supplemental honeywater feeding did not accelerate de-

VOLUME 55, NUMBER 2 89

velopment or increase the brood size. Queens of Camponotus vicinus may

cooperate during colony foundation; females doing so increased the size of

the first worker brood, although the development period was not appreciably

shortened.

Acknowledgments

I thank Randy Oliver and Roy Snelling for collecting some of the ants

used in this study, and George L. Hunt and Peter J. Bryant for providing

space for the ant cultures at the University of California, Irvine. I also thank

Roy Snelling, George Hunt, Michael M. Martin and Richard D. Alexander

for constructive comments and criticism during preparation of this manu-

script.

Literature Cited

Brian, M. V. 1973. Feeding and Growth in the ant Myrmica. J. Anim. Ecol. 42: 37-53.

Creighton, W. S. 1950. The Ants of North America. Bull. Mus. Comp. Zool., Harvard, v.

103.

Eidmann, H. 1926. Die koloniegrundung der einheimischen ameisen. Vergleich. Physio. 3:

776-826.

Hamilton, W. D. 1964. The genetical evolution of social behaviour II. J. Theoret. Bio. 7: 17.

Haskins, C. P., and E. F. Haskins. 1955. The pattern of colony foundation in the archaic ant

Myrmecia regularis H. Insectes Sociaux 2: 115-126.

Holldobler, B. 1961. Temperaturunabhaengige rhythmische erscheinungen bei rossameisen-

kolonien (Camponotus ligniperda Latr. und Camponotus herculeanus L.). (Hym.

Form.). Insectes Sociaux 8: 13-22.

Holldobler, B. 1962. Zur Frage der Oligogynie bei Camponotus ligniperda und Camponotus

herculeanus. Z. Angew. Entomologie 49: 337.

HOolldobler, B., and E. O. Wilson. 1977. The number of queens: an important trait in ant

evolution. Naturwissenschaften 64: 8-15.

Janzen, D. 1966. Coevolution of mutualism beween ants and acacias in Central America.

Evolution 20: 249-275.

Markin, G. P., Collins, H. L., and J. H. Dillier. 1972. Colony founding by queens of the

Red Imported Fire Ant, Solenopsis invicta. Ann. Entomol. Soc. Amer. 65: 1053-1057.

Pricer, J. L. 1908. The life history of the Carpenter Ant. Bio. Bull., Woods Hole, Mass.

14: 177.

Stumper, R. 1962. Sur un effet de groupe chez les femelles des Camponotus vagus (Scopoli).

Insectes Sociaux 9: 329-333.

Waloff, N. 1957. The effect of the number of queens of the ant Lasium flavus (Fab.) (Hym.,

Formicidae) on their survival and on the rate of development of the first brood. Insectes

Sociaux 4: 391-408.

Weber, N. A. 1972. Gardening Ants, the Attines. Mem. Amer. Philosoph. Soc., v. 92.

West Eberhard, M. J. 1969. The social biology of polistine wasps. Misc. Pub. Mus. of Zool.,

Univ. of Michigan 140:1-101. .

Wheeler, W. M. 1933. Colony foundation among ants. Harvard Univ. Press.

Wilson, E. O. 1971. The insect societies. Belknap/Harvard Univ. Press.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 90-94

ALFALFA LEAFCUTTER BEE—REDUCING PARASITISM OF LOOSE

CELLS DURING INCUBATION (HYMENOPTERA: MEGACHILIDAE)

FRANK D. PARKER

Bee Biology and Systematics Laboratory, Federal Research,

Science and Education Administration, USDA,

Utah State University, Logan, 84322

The alfalfa leafcutter bee, Megachile pacifica (Panzer), is the most com-

monly used pollinator of western alfalfa seed fields. One method of increas-

ing these bees entails removing their leaf cells from nesting media in the fall

and cold-storing them until spring. Bees are obtained by incubating the cells

for approximately 20 days at 85°F (30°C). Often some cells are parasitized

by chalcid wasps (Monodontomerus—an external parasite, Tetrastichus—

an internal parasite), and since the parasites emerge 11-12 days after begin-

ning incubation of the bees, they have amply opportunity to parasitize many

additional host cells before bee emergence occurs. Various types of light

traps or cell coverings have been used to reduce parasitism (Waters, 1970;

Hobbs, 1973; and Johansen et al., 1973), but none of these methods is

satisfactory. Recently Brindley (1976) recommended dipping cells in car-

baryl insecticide to reduce parasitism. Some growers have sprayed insec-

ticides on loose cells during incubation, but information on bee kill or par-

asite control has not been recorded. Because these insecticide control

methods employ toxic agents and are not commonly used, methods to repel

parasites before parasitism occurred were tested.

Methods

Thirty-three groups (150 cells each) of leafcutter bee cells were placed in

covered 6-o0z plastic cups and incubated at 90°F for 10 days. Before parasite

emergence the cells were removed from the incubator. N, N-diethyl-m-

toluamide (deet) (99.6% stock) was diluted in 20% sugar water to prepare

0, 2, 6, and 18% repellent solutions. Four groups of cells were dipped for

5 sec in each of the four repellent solutions (4 treatments, replicated 4x).

The cells were air-dired at 32°C for 3 hr in the greenhouse. In a second test

vermiculite pieces (1-3 mm) were soaked for 10 sec in each of the 4 repellent

solutions and air-dried as were the leafcutter bee cells. Four groups of cells

were mixed and covered with 3 mm of dried, treated vermiculite (4 treat-

ments, replicated 4x).

The last group of incubated cells was isolated individually in gelatin cap-

sules (1 group, | replication). This treatment was necessary to provide con-

VOLUME 55, NUMBER 2 91

trol bees for the last part of these tests. Then, all cell groups in each treat-

ment were replaced in cups and returned to the incubator.

Four other cell groups (150 cells each) used as a control, were dissected

and their contents were recorded. This group came from the same stock bee

population that was held in cold storage, and it provided data on the inci-

dence of parasitism in the stock bees.

Parasite and bee emergence in the treatments were recorded 3 x/day (8,

11, 16 hr). The parasites were left in the plastic cups, but the bees were

removed during each observation period. One hundred bees (50 ¢6, 50

? °) from each treatment were placed in large covered cages (in the green-

house) and fed a 20% sugar-water solution. Dead bees were recorded and

removed from the cages daily to assess longevity of bees emerged from each

treatment. After all parasites and bees had emerged, cells without exit holes

were dissected and their content were recorded.

Results

Deet-treated cells.—Tetrastichus began to emerge from the cells on the

11th day, and Monodontomerus emerged on the 12th day of incubation. The

Tetrastichus in the 0% test remained active, mating and crawling about the

cells for 13 days, whereas these parasites in the 2, 6, and 18% repellent

treatments concentrated beneath the lids and away from the cells. The Te-

trastichus population in the repellent treatment cups began to decline 4 days

after emergence; no parasites were alive 8 days after emergence. Mono-

dontomerus reacted similarly but often crawled about the cells in all the

cups.

Cells treated with repellents had significantly greater bee emergence than

the 0% treatment, in which no bees emerged. The number of bees exiting

cells was highest in the 6% treatment (Table 1). However, parasite emer-

gence was significantly lower in the repellent-treated cells than in the 0%

treatment. Also, parasite emergence was lower in the 18% treatment (9.1%)

than emergence in the gelatin capsules (21.3%) and emergence in the dis-

sected controls (22.8%). Also, a significantly higher number of dead cells

was found in the 18% deet treatments. Apparently this treatment killed some

of the bees and parasites during incubation.

Vermiculite-covered cells.—Parasite activity in this test was similar to

that observed in the deet treatments; namely, parasites in the cups with

repellent concentrated under the lids and away from the vermiculite. The

emergence and death rate of the parasites were similar.

As with the deet-treated cells, significantly more bees emerged in the

treatments with repellents than in the 0% treatments. Emergence was great-

est in the 6 and 18% treatments and significantly higher than emergence in

the 2% treatment. However, cells in gelatin capsules had some mortality,

92 PAN-PACIFIC ENTOMOLOGIST

Table 1. Effect of vermiculite and deet treatments on emergence of alfalfa leafcutter bees

and parasites, Logan, Utah, 1977.

Live bees Live parasites Dead cells

Avg. Avg. Avg.

Treatment no. % no. % no. %

Vermiculite: 0% 0.00 (0.00)a! 79.00 (52.60)a 70.50 (47.00)a

2% 73.50 (48.70)b 42.00 (28.00)b 29.00 (19.30)b

6% 82.25 (54.60)c ce re (22.50)b 29.25 (19.50)b

18% 87.75 (58.00)c 35.00 (23.30)b 23.00 (15.30)b

Deet: 0% 0.00 (0.00)a 80.00 (53.30)a 67.75 (45.20)a

2% 60.50 (40.30)b 39.50 (26.33)b 48.50? (32.30)b

6% 85.00 (56.70)c 29.50 (19.66)c 30.75 (20.50)c

18% 35.25 (23.50)d 37 (9.10)d 94.25? (62.80)d

1 Numbers followed by the same letter are not significantly different (P < .05).

2 Significant difference between vermiculite and dipped treatments (P < .05).

probably because of desiccation, since 14% of these cells contained dead,

but fully formed adults. If these values are combined, the 47.3% emergence

rate plus the 14% dead adults (61%) in the gelatin capsules approximates

the emergence rate observed in the 18% repellent treatment (58%). How-

ever, significantly lower levels of parasite emergence were found in the

repellent treatment all of which were similar (Table 1). Also, the number of

dead cells was significantly lower in the repellent treatments (15.3-19.5%)

and similar to the number of dead cells found in the gelatin capsules (16.6%).

Caged bees.—The longevity of bees emerging from 2% repellent treat-

ments was similar to that of bees emerging from the gelatin capsule treat-

ment (Fig. 1). Longevity of bees from the 6% dipped cells was also similar,

but the bees from the 6% vermiculite had a slightly higher death rate (Fig.

1). The highest death rate was observed in bees from the 18% dipped and

vermiculite treatments (Fig. 1). All bees died 20 days after emergence be-

cause of uncontrolled, high greenhouse temperatures.

Discussion

The results of these tests indicate that the repellent treatments signifi-

cantly reduced parasitism during incubation of loose cells and thereby in-

creased bee emergence. The 6% deet treatment appeared to cause the high-

est bee emergence, with longevity about equal to that of untreated bees. It

was a surprise to observe a higher death rate for bees emerging from the

vermiculite, since higher death rates were not observed among the immature

stages during the test. Several factors could have contributed to this higher

VOLUME 55, NUMBER 2 93

2% REPELLENT 6% REPELLENT

= deet

BB vermicut ite

a) deet

Bvermiculite

me TTT TT

ig Bvermicul ite

70)

18% REPELLENTS GELATIN CAPSULES

60.

40)

30.

10)

ae 10 15 20 2 10 15 20

DAYS

Fig. 1. Cumulative percent death of adult Megachile pacifica in repellent treatments.

rate. First, the bees were confined in the small cups for several hours after

their emergence. In a commercial operation they could easily escape soon

after emergence. Also, the vermiculite may have absorbed and concentrated

the deet since the repellent odor was very strong when the vermiculite-

covered cells were opened.

In laboratory tests an aerosol repellent works very well in large incubators

where blue-black lights are used over pans of soapy water. The repellent

keeps the parasites away from the cells, and they are more readily attracted

to the light traps. I have sprayed loose cells of our laboratory colony with

repellent for the past 2 seasons and practically eliminated parasitism during

94 PAN-PACIFIC ENTOMOLOGIST

incubation. Bees emerging from our sprayed cells have nested adequately;

their population has increased by over 2-fold each season.

Aerosol repellents are available from several commercial sources, with

concentrations of active ingredients ranging between 6 and 12%. The con-

centration of active ingredients in the aerosol used to spray our laboratory

colony was 71% (Stock No. 6840-00-082-2541, General Services Adminis-

tration, Federal Supply Service, Washington, D.C. 20406). The differences

between the results noted in these tests and those observed in our sprayed

laboratory colony of leafcutter bees may have been caused by a dilution

factor. The amount of applied active ingredient in an aerosol is much less

than in the repellent solutions used to dip cells. Therefore, it may be nec-

essary to use a higher concentration of active ingredients if aerosol sprays

are used.

Acknowledgments

Thanks are due to H. Potter of this laboratory for assistance in the lab-

oratory work. C. Hatley is acknowledged for her analysis of the data and

helpful suggestions on the manuscript. Appreciation is expressed to these

persons who reviewed the manuscript, W. Brindley and D. Davis (Utah

State University).

Literature Cited

Brindley, W. 1976. Carbaryl control of chalcidoid parasites from alfalfa leafcutting bees. J.

Econ. Entomol. 69: 225-228.

Hobbs, G. 1973. Alfalfa leafcutter bees for pollinating alfalfa in western Canada. Can. Dep.

Agric. Publ. 1495. 30 p.

Johansen, C., J. Eves, and C. Baird. 1973. Control of alfalfa leafcutting bee enemies. Wash.

State Univ. Coop. Ext. Serv. EM 2631 (Rev.). 10 p.

Waters, N. 1970. Lights and water traps for alfalfa leafcutter bee incubators. Idaho Agric.

Exp. Stn. Curr. Inf. Ser. 120. 4 p.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 95-98

ORIGINS OF PSYLLID FALLOUT IN THE CENTRAL SIERRA

NEVADA OF CALIFORNIA (HOMOPTERA)

RICHARD P. PApPp! AND JAMES B. JOHNSON

Dept. Entomol. Sci., Univ. of California, Berkeley, 94720

Psyllidae (Insecta: Homoptera) make up a major proportion of the wind-

blown (aeolian) insect fauna deposited on alpine snowfields in the Sierra

Nevada of California (Papp, 1975). During the summers of 1972 to 1974,

1206 psyllid specimens comprising 18 species in six genera were collected

on névé snow at 3353 m on Mt. Conness in the Inyo National Forest. Since

psyllids are an important food for nival aeolian zone predators and scav-

engers (Papp, 1978), knowledge of their origin is basic to understanding the

dynamics of energy flow into the aeolian ecosystem as well as details of the

intrasystem community structure.

Four species of Psyllidae (Psylla hirsuta, P. magna, P. alba and Aphalara

artemisiae) accounted for 93.2% of the total psyllid fallout on Mt. Conness

snowfields (Table 1).

At least 14 other species were taken on snow during 1972-74, including

some species taken only once. Among these were the widespread pear psyl-

la, Psylla pyricola Forster. Hagen (1976, personal communication) has tak-

en this species at high altitudes in the Alps. Another species, Paratrioza

cockerelli (Sulc) has many solanaceous hosts, including potato and tomato.

A complete listing of aeolian psyllids is given in Table 2.

Distribution of the hostplant species for the four common psyllids col-

lected on Mt. Conness snow are given in Table 3. Four species, Cercocarpus

betuloides, Corylus cornuta, Salix melanopsis and Salix Hindsiana, are

found only on the west slope of the Sierra Nevada, and none occur above

Table 1. Common psyllids collected on Mt. Conness snowfields, 1972-74.

Specimens collected

ee % of

Species 1972 1973 1974 Total total

Psylla hirsuta (Tuthill) 0 107 165 eee 22.6

Psylla magna Crawford 0 245 30 275 22.8

Psylla alba Tuthill 1 453 20 474 39.3

Aphalara artemisiae complex 0 21 82 103 8.5

Other spp. 3 15 64 82 6.8

Total 4 841 361 1206 100.0

96 PAN-PACIFIC ENTOMOLOGIST

Table 2. Aeolian Psyllidae collected on Mt. Conness, 1972-74.

Species Host(s)

Aphalara calthae complex Polygonum spp., Artemisia tridentata, Caltha

palustris, others

Aphalara minutissima Crawford Artemisia tridentata

Aphalara veaziei metzaria Crawford Achillea millefolium, other Compositae

Euphalerus vermiculosis Crawford Ceanothus spp., Lupinus sp.

Livia juncorum Latreille Juncus, Carex, Pinus

Paratrioza cockerelli (Sulc) Solanaceae, inc. Solanum tuberosum (potato)

and Lycopersicon esculentum (tomato)

Psylla alba Crawford Salix melanopsis, S. Hindsiana, S. Hindsiana

var. sessilifolia, Cercocarpus ledifolius

Psylla breviata Patch Salix

Psylla hirsuta (Tuthill) Cercocarpus ledifolius, Purshia tridentata,

Corylus cornuta

Psylla insignita Tuthill Cercocarpus ledifolius

Psylla magna Crawford Cercocarpus ledifolius, C. betuloides,

‘*Serviceberry”’ (prob. Amelanchier)

Psylla magnacauda Crawford Shepherdia argentea, Eleagnus sp.

Psylla media Tuthill Cercocarpus ledifolius, C. betuloides

Psylla minor Crawford Salix californica, S. lasiolepis

Psylla pyricola Forster Pyrus communis (pear)

Trioza incerta Tuthill Salix

Trioza sp. (singularis ?) unknown

2500 m in California. The other hostplants, Cercocarpus ledifolius and Pur-

shia tridentata, are distributed along the east slope of the Sierra Nevada at

altitudes ranging up to 3200 m. While climatological data show that pre-

vailing winds across the Sierra are primarily from the west, southwest, or

south (96%, 69%, 88% in 1972, 1973, 1974 respectively) in late spring and

early summer, turbulence resulting from high velocity winds blowing across

the Sierran crest also create diurnal anabatic (upslope winds) from the east-

ern deserts. It is therefore apparent that psyllid fallout along the crest of the

Central Sierra Nevada has at least two origins: a major component from the

Central Valley and the west slope (aeolian transport by geostrophic winds),

and a minor component from the Owens Valley and east slope (aeolian

transport by anabatic winds originating in the deserts leeward of the Sierra

Nevada).

VOLUME 55, NUMBER 2

Table 3. Distribution of common psyllid hostplants. (Data from Munz, 1959, 1968.)

Hostplant (Psyllidae)

Cercocarpus ledifolius Nuttall

(Psylla hirsuta, P. alba)

Cercocarpus betuloides Nuttall

(Psylla magna)

Purshia tridentata (Pursh) DeCandolle

(Psylla hirsuta)

Corylus cornuta Marsh

(Psylla hirsuta)

Salix Hindsiana Benth.

(Psylla alba)

Salix Hindsiana Benth.

var. leucodendroides (Rowlee) Ball.

(Psylla alba)

Salix melanopsis Nutt.

(Psylia alba)

Distribution

Dry, rocky slopes along the east side of the

Sierra Nevada at 1220-3200 m; also in

mts. of W Mojave Desert and N to Modoc

and Siskiyou Cos.; to E Washington,

Montana, Colorado, Arizona, Baja

California.

Chaparall and oak woodlands along the W

slope of the Sierra Nevada below 1830 m;

also cis-montane California to SW Oregon;

N Baja California.

Dry slopes along east side of Sierra Nevada

at 900-3200 m; also in the White Mts.; in

California ranges from Tulare and Inyo

Cos. N to Modoc, Siskiyou and Trinity

Cos.; thence to Lake Co., British

Columbia, Montana, New Mexico.

Damp slopes and banks, below 2200 m, in

many plant communities; in the Coast

Ranges from Santa Cruz Co. N, and in the

Sierra Nevada from Tulare Co. N; to

British Columbia.

Common locally among ditches, on sand

bars, etc., below 3000 ft; many plant

communities; cis-montane California, into

Oregon and Baja California.

Sparingly in Santa Clara and Tulare Cos. to

Kern Co.; more common Ventura to San

Diego Cos. to Baja California.

Stream banks below 8000 ft; many plant

communities; Sierra Nevada N to Modoc

County, coast ranges from Lake and

Sonoma Cos. N; to British Columbia;

Rocky Mountains.

Sierran alpine predators and scavengers which are able to withstand the

rigors of nival foraging are thus able to take advantage of aerial plankton

fallout originating from at least two directions: the forest and riparian biomes

to the west, and the pinyon-sagebrush biomes to the east.

Acknowledgments

The host and distribution data for Psyllidae were taken from California

Insect Survey specimen data and from unpublished notes of the late Dr.

98 PAN-PACIFIC ENTOMOLOGIST

Dilworth Jensen, formerly of the University of California, Berkeley. Ap-

preciation is expressed to the U.S. Forest Service for permission to conduct

research in the Harvey Monroe Hall Natural Area, to the Carnegie Insti-

tution of Washington for use of the Timberline Station facilities, and to the

Division of Biological Control at the University of California, Berkeley for

partial financial support during field research.

Literature Cited

Munz, P. A. 1959. A California Flora. Univ. Calif. Press, Berkeley, 1681 p.

Munz, P. A. 1968. Supplement to A California Flora. Univ. Calif. Press, Berkeley, 224 p.

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

nities in the Central Sierra Nevada of California. Ph.D. thesis, Univ. Calif. Berkeley,

158 p.

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

117-131.

Footnote

1 Current address: Dept. Entomology, Bernice P. Bishop Museum, P.O. Box 6037, Honolulu,

Hawaii 96818.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 99-105

A NEW SPECIES OF PHANAEUS FROM MEXICO

(COLEOPTERA: SCARABAEIDAE)

W. D. EDMONDS

California State Polytechnic University, Pomona, 91768

Phanaeus is a New World genus of dung beetles well known for bright,

metallic colors and striking sexual dimorphism. The purpose of this paper

is to describe a new Mexican species with unusual ecological characteristics.

The primary geographical center of diversity of Phanaeus is tropical Mexico

(Edmonds, 1972). Rather than to Mexican groups, however, the new species

is more closely related to those Central and South American species which

comprise the P. splendidulus-group (Edmonds, 1972). Of these species, it

is most closely related to P. endymion Harold, the only other member which

also occurs in Mexico.

Phanaeus halffterorum, new species

(Figs. 1-3, 6-8)

Holotype.—Male, Mexico, state of Mexico, 8 km W Temascaltepec, 2360

m, VII-11-76, fungus in pine-oak forest, W. D. Edmonds, P. Reyes and B.

Kohlmann.

Paratypes.—3 males, 2 females, same data as holotype; 4 males, 4 females

(1 designated allotype), 5 km E Temascaltepec, Real de Arriba, 2200 m,

VII-10-76, fungus in oak-pine forest, W. D. Edmonds, P. Reyes, B. Kohl-

mann; 1 male, 1 female, 5S km W Temascaltepec, 2200 m, fungus in oak-pine

forest, VII-23-77, W. D. and T. B. Edmonds; 1 male labeled ‘‘Real de

Arriba, VII-1932, 6300 ft, Mexico D. F., Hinton coll., BM 1939-583’"!; 1

male labeled ‘‘Mex. Guerrero, 22 mi S Chilpancingo, 2800 ft, VIII-2-1964,

Richard D. Page, col.’’

Disposition of types.—Holotype and allotype—California Academy of

Sciences, San Francisco (CAS Ent., Type 13184); 1 pair paratypes—British

Museum (Natural History), London; | pair paratypes—Halffter collection,

Mexico City; 1 male paratype—United States National Museum, Washing-

ton, D.C.; 1 male paratype—A. Martinez collection, Buenos Aires; remain-

ing paratypes—temporarily in my collection.

Derivation of epithet.—From the surname Halffter. It is my pleasure to

dedicate this species to my very good friends and colleagues, Gonzalo Halff-

ter and his lovely wife Violeta M. de Halffter, in recognition of their many

contributions to the study of Scarabaeinae.

Major male.—Dorsum dark, shining, iridescent green or dark blue (iri-

100 PAN-PACIFIC ENTOMOLOGIST

Figs. 1-5. Figs. 1-3. Phanaeus halffterorum: Fig. 1—dorsal view female pronotum (arrow

indicates mid-dorsal depression); Fig. 2—dorsal view left side male pronotum; Fig. 3—dorsal

view left elytron of male (arrows indicate 4th stria). Fig. 4—Phanaeus endymion, dorsal view

right side male pronotum. Fig. 5—Phanaeus funereus, dorsal view left elytron of male (arrow

indicates 4th stria).

descence subdued on elytra) except outer margin of head, cephalic process

(‘‘horn’’),? posterior part of head and lower surface of posterior pronotal

angles, which are shining black; venter red-brown to chocolate-brown ex-

cept abdominal sterna, middle and hind femora and pteropleura, which are

tinged with green (or blue). Clypeus distinctly bidentate, teeth rounded;

subclypeal process transverse. Cephalic process long, tapering, curved

evenly posteriorly over pronotum. Prothoracic disk (Fig. 6) triangular, flat-

tened but with distinct undulations laterally and posteriorly; posterior angles

very Salient, directed posterolaterally and slightly upturned apically; middle

of anterior margin with a strong, acute tooth (except Guerrero specimen);

VOLUME 55, NUMBER 2 101

Figs. 6-10. Figs. 6-8. Phanaeus halffterorum, male: Fig. 6—dorsal view pronotum (guide-

line indicates anterior tooth); Fig. 7—right front tibia; Fig. 8—caudal view tips of elytra,

pygidium and 8th abdominal sternum (guideline indicates raised inner margin of elytron). Figs.

9-10. Phanaeus endymion, male: Fig. 9—dorsal view pronotum; Fig. 10—caudal view tips of

elytra, pygidium and 8th abdominal sternum.

pronotum distinctly granulate dorsally (Fig. 2), granulation, while better

appreciated under magnification, visible as fine texturing (like that of sand-

paper) to unaided eye; sculpturing becoming punctate posteriorly and pro-

gressively more clearly punctate and less granulate anterolaterally; disk sha-

greened and highly shining. Front tibia quadridentate (Fig. 7). Elytra (Fig.

3) with very fine, simple striae, interstriae distinctly convex medially such

that striae lie in longitudinal furrows; inner margin a ridge progressively

more raised and keel-like posteriorly which extends beyond apical margin

of elytron as rounded tooth (Fig. 8); lateral margin of elytron distinctly

102 PAN-PACIFIC ENTOMOLOGIST

excised apically adjacent to inner margin. Pygidium (Fig. 8) weakly to mod-

erately punctate, punctures usually coalescent at least medially; each side

with shallow elongate depression.

Minor male.—As above except as follows: Cephalic process shorter,

more upright or reduced to a simple tubercle; flattened, triangular shape of

pronotum much less pronounced, posterior angles reduced to tubercles well

anterior to posterior margin, anterior tooth absent.

Female.—As above except as follows: Cephalic process a trituberculate,

transverse carina extending between ends of postclypeal carinae; most of

pronotal disk shining black, green color not iridescent (blue phase known

in male only). Pronotum evenly covered dorsally with shallow punctures,

appearing very smooth to unaided eye; puncturing sometimes effaced me-

dially and coalescing laterally to produce weak rugosity; surface not sha-

greened, coloring (where present) less brilliant than in male; strongly con-

vex, bearing weakly raised anterior transverse ridge with three isolated

tubercles followed by shallow concavity; distinct mid-longitudinal depres-

sion extending from posterior margin to near middle of disk (Fig. 1, arrow).

Size. —Length 12-19 mm; width (at bases of elytra) 8-12 mm.

Habitat and distribution.—Temperate oak-pine/pine-oak forests ordinar-

ily above 1900 m along southern slopes of the Transverse Volcanic Range

and in the highlands of Guerrero, Mexico; feeding on wild mushrooms;

probably active at dusk and early evening hours.

Discussion

This species is the same mentioned by me in 1972 as incertae sedis (p.

830); females, which were not available then, indicate without doubt that

P. halffterorum is a member of the endymion-complex of the P. splendi-

dulus-group. Pronotal sculpturing of the male, however, requires that the

second alternative of the first couplet of my key to the species groups and

complexes of Phanaeus (p. 829) be modified partially to read as follows:

‘*... or minutely to distinctly granulate or granulorugose (males of endy-

mion complex) ....’’ P. endymion differs from halffterorum by the fol-

lowing combination of characters, the counterparts of which are included

in the above description: the color is weakly shining green or blue and never

highly iridescent on the head and pronotum of the male; most of the disk

of the female is colored, less of it is black; the pronotum of the large males

(Fig. 10) is more broadly triangular, very flat, and the posterior angles are

less salient and directed laterad; even the largest males are without a trace

of an anterior pronotal tooth; the pronotum of the female has at most a fine

mid-dorsal, longitudinal line on the posterior part of the pronotum and is

never distinctly impressed along this line; the disk of the male pronotum

(Fig. 4) is at most only weakly granulorugose, appearing virtually smooth

VOLUME 55, NUMBER 2 103

to the unaided eye and, seen under magnification, often with minute widely

spaced, shining punctures in a field of effaced granules and weak shagreen-

ing; the inner margin of the elytron is not ridge-like, lacks an apical tooth

and is not excised (Fig. 10); the pygidium (Fig. 10) lacks indications of

lateral depressions; the species inhabits lowland evergreen and deciduous

forests of eastern and southeastern Mexico and Guatemala below 800 m,

feeds on excrement of non-herbivores and, occasionally, carrion; active

during early morning hours.

The most useful morphological features for distinguishing halffterorum

from other members of the complex are the shape and sculpturing of the

male pronotum and the strong ridge along the inner margin of each elytron.

Most groups of Phanaeus, including the endymion-complex, are in need of

revision. I have insufficient data to reliably distinguish among the other,

currently recognized members of the complex, blanchardi Olsoufieff, fu-

nereus Balthasar, and pyrois Bates, all of which are from Central or north-

west South America and presumed to be reasonable taxonomic species.

Collectively, however, these three species differ from halffterorum and en-

dymion by the following characters: the elytral interstriae are relatively flat,

the striae are very fine and not lying in pronounced longitudinal depressions

(Fig. 5); the disk of the male pronotum is without granulations, appearing

smooth even under magnification, except for weak roughening anterolat-

erally; the elytra and most or nearly all of the pronotum are dull black, not

shining; or, if a shining red color present (pyrois), it is restricted to the

pronotum.

The Guerrero specimen, which is a large male, differs from those collected

in Temascaltepec by the lack of an anterior pronotal tooth, by being less

distinctly granulate and darker, less shining green. It is interesting that halff-

terorum is evidently more widely distributed geographically than the special

ecological conditions of Temascaltepec might lead one to predict. I do not

know the Guerrero locality, but the elevation (2400 ft, 735 m) suggests a

significantly different ecological setting from that of the environs of Temas-

caltepec. The ecological comments below are based on observations made

in the Temascaltepec area.

Temascaltepec is located on the southern slopes of the Transverse Vol-

canic Range at 19°02’N, 100°02'W. The area supports extensive stands of

oak-pine to pine-oak forests on very uneven terrain. All specimens collected

there have come from inside or near the margins of the forests between 1935

and 2360 m (6350 and 7750 ft). As was originally reported by Hinton (1935)

and later by Halffter and Matthews (1966), who referred to this species as

endymion, P. halffterorum is mycetophagous and attracted only to several

species of wild mushrooms (none yet identified) common during the rainy

season. It has never been found associated with excrement of the many

domestic animals (cattle, horses, burros, swine) which roam the area nor in

104 PAN-PACIFIC ENTOMOLOGIST

human excrement or carrion used to bait pitfall traps. When partially de-

composed, mushrooms are treated by adults as is excrement or carrion by

other Phanaeus: beginning with the stalk, the fungus is packed by pieces

into the blind end of a tunnel dug directly beneath it. Here it is either

consumed by the adults or used to fashion brood balls.

Mycetophagy by scarabaeine dung beetles has been known for many

years. While many species have been collected from decomposing fungi (see

Halffter and Matthews, 1966), very few appear to be as strictly mycetopha-

gous as is halffterorum. The following have also been collected from wild

mushrooms in the Temascaltepec area, although only the former is evidently

strictly mycetophagous: Oniticellus rhinocerulus Bates; Phanaeus daphnis

Harold (Hinton, 1935), otherwise very common in cattle dung; one species

each (not yet identified) of Ateuchus and Onthophagus which are more

commonly collected in cattle and horse dung.

The southern slopes of the Transverse Volcanic Range are interrupted by

a series of valley systems which descend steeply to the valley of the Balsas

River. Temascaltepec is located near the upper (northern) end of one such

valley system. Of zoogeographic interest is the fact that Temascaltepec, like

similar places along the Transverse Volcanic Range, supports a dung beetle

fauna which comprises both nearctic and neotropical elements. Copris,

Onthophagus, Oniticellus and Ceratoptrupes (Geotrupinae) are northern

contributions; Phanaeus, Ateuchus, Canthidium, Deltochilum, Dichotomi-

us and Coprophanaeus are southern representatives. Such faunal mixing in

the Mexican transition zone has been discussed by Halffter (1964, 1976). In

accordance with ideas I presented in 1972, P. halffterorum can be inter-

preted as the product of relatively recent speciation; it undoubtedly repre-

sents the deepest northward eco-geographic penetration of the P. splendi-

dulus-group into North America.

P. halffterorum has been successfully reared in the laboratory. Two pairs

of field-collected adults were introduced into the same vertical terrarium

(95 x 60 X 7 cm) on 12 July, 1976, and provided decomposing fungi from

the type locality. Later, partially decomposed commercial (edible) mush-

rooms were provided to replace the original food supply, which was buried

quickly. Four brood balls were recovered on 14 September; all nesting de-

tails agreed with those of other known Phanaeus (Halffter and Matthews,

1966; Halffter, 1977). A fifth brood ball was recovered on October 6, at

which time surviving adults (1 male, 2 females) were supplied with human

excrement. A sixth brood ball, provisioned with human excrement, was

recovered on 8 November; it yielded an egg in early stages of decomposi-

tion. Although field and laboratory data may suggest strict mycetophagy,

further rearing trials with excrement are necessary before concluding wheth-

er or not halffterorum requires fungi for successful nidification. All five

VOLUME 55, NUMBER 2 105

fungus-provisioned brood balls yielded eggs which were allowed to develop

to 3rd (final) instar larvae. The larva of halffterorum is virtually identical to

those of other known phanaeines (Edmonds and Halffter, 1978).

Acknowledgments

This paper reports partial results of a cooperative project on dung beetle

ecology supported jointly by the National Science Foundation (U.S.—Mex-

ico Cooperative Science Program, Grant No. INT76-06712) and the Consejo

Nacional de Ciencia y Tecnologia (Programa Nacional Indicativo de Eco-

logia—Proyecto ‘‘Interacciones entre ganado y pastizales’’). Mr. Michael

Thompson prepared the photographs; Mr. James McCabe prepared the

drawings.

Literature Cited

Edmonds, W. D. 1972. Comparative skeletal morphology, systematics and evolution of the

phanaeine dung beetles (Coleoptera: Scarabaeidae). Univ. Kansas Sci. Bull. 49: 731-

874.

Edmonds, W. D., and G. Halffter. 1978. Taxonomic review of immature dung beetles of the

subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Syst. Entomol. 3: 307-331.

Halffter, G. 1964. La entomofauna americana; ideas acerca de su origen y distribucion. Folia

Entomol. Mex., No. 6, pp. 1-108. .

Halffter, G. 1976. Distribucion de los insectos en la zona de transicidn mexicana; relaciones

con la entomofauna de Norteamerica. Folia Entomol. Mex. No. 35, pp. 1-64.

Halffter, G. 1977. Evolution of nidification in the Scarabaeinae (Coleoptera: Scarabaeidae).

Quaest. Entomol. 13: 231-253.

Halffter, G., and E.G. Matthews. 1966. The natural history of dung beetles of the subfamily

Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomol. Mex., Nos. 12-14, pp. 1-312.

Hinton, H. E. 1935. Anotaciones acerca de las cosumbres micetofagicas de dos especies de

Phanaeus. An. Instit. Biol. (Mex.) 6: 129-130.

Footnotes

‘ This specimen is evidently one of the series of eleven referred to by Hinton, 1935. I have

been unable to locate the remaining ten specimens.

2 Terminology used here is that established by Edmonds, 1972.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 106-110

A NEW SUBSPECIES OF POLIAENUS NUEVOLEONIS

CHEMSAK AND LINSLEY FROM SOUTHERN ARIZONA

(COLEOPTERA: CERAMBYCIDAE)

DURWARD D. SKILES

Institute of Geophysics and Planetary Physics, Univ. of Calif. Los Angeles,

90024

In a recent review of Mexican Pogonocherini, Chemsak and Linsley

(1975) described the new species Poliaenus nuevoleonis from a male taken

in northeastern Mexico near Monterrey, Nuevo Leon. Subsequently, I have

examined over 70 specimens of that species from the mountains of western

Texas and southeastern Arizona, as well as a single specimen from western

Mexico about 100 km west of the city of Durango. The specimens from

Arizona and western Mexico are readily distinguished from those from Tex-

as and northeastern Mexico and merit subspecific recognition. Below I have

modified the original description of P. nuevoleonis to accommodate the

substantial morphological variation exhibited by the species.

Poliaenus nuevoleonis Chemsak and Linsley

Male.—Form elongate-robust, sides subparallel. Integument shining, red-

brown to blackish, clothed throughout with fine, appressed, moderately

dense pubescence partially obscuring surface, less densely clothed with long

flying hairs which are predominantly dark brown or black on dorsum, white

on venter, intermixed on appendages. Head with pubescence brownish

white, vaguely washed with gold on occiput; frons two-thirds again as broad

as high; eye with lower lobe subquadrate, slightly erect, upper lobe distinct-

ly narrower than interocular space; antenna red-brown, exceeding elytral

apex by three or four segments, scape moderately slender, attaining lateral

prothoracic tubercle, third segment longest, fourth subequal to scape, half

again as long as fifth, fourth to eleventh basally annulate with dense, ap-

pressed, white pubescence. Pronotum with pubescence brownish white,

distinctly washed with golden-orange on basal half; width across lateral

tubercles one and one-fifth to one and one-third times length, dorsal tu-

bercles moderate, distinctly separated by flat pronotal disc, lateral tubercles

moderate to rather prominent, blunt, occasionally slightly recurved. Elytra

two times as long as wide, with a V-shaped antemedian impression extend-

ing from suture to humeri, apices conjointly or separately rounded; lateral

and median costae fine, subdued or moderately prominent, extending from

humeri to apical third or fourth of elytra, thence becoming evanescent and

VOLUME 55, NUMBER 2 107

indicated only by a few minute dark brown penicilli; subsutural costa indis-

tinct, indicated only by a prominently penicillate sub-basal gibbosity and by

an elongate post median swelling containing from one to five prominent,

separate or confluent penicilli; punctures coarse, separate basally and lat-

erally, becoming shallower apically, vanishing abruptly at apical third; pu-

bescence moderate to dense, often largely obscuring basal punctures,

brownish white, washed with gold along subsutural costae, subsutural pen-

icilli dark brown, occasionally margined with orange. Abdominal sternites

densely fringed with white hairs, fifth subequal to fourth, apex broadly

rounded or subtruncate. Length 5.9-10.2 mm.

Female.—Form slightly more robust, antennae slightly shorter than in

male. Upper lobe of eye about half as wide as interocular space. Fifth

abdominal sternite about twice as long as fourth, distinctly impressed me-

dially, apex broadly rounded to subtruncate. Length 6.1-10.9 mm.

Two subspecies, widely separated geographically, can be recognized.

Poliaenus nuevoleonis nuevoleonis Chemsak and Linsley

Lateral and median elytral costae fairly prominent. Pubescence sufficient-

ly dense to largely obscure basal elytral punctures and give dorsum a hoary

appearance. Appressed white pubescence covering at least basal three-fifths

of antennal segments four to ten. Length 7.4-10.2 mim.

Material examined.—MEXICO: NUEVO LEON: Holotype ¢ from Chi-

pinque Mesa, 5400 ft near Monterrey, VII-23-63 (H. Howden); Monterrey,

1°, IW-17-53. COAHUILA: Cuesta La Murrala, 1 ¢ , [X-12-76 (J. A. Chem-

sak, J. Powell, A. and M. Michelbacher). TEXAS: Big Bend National Park,

Chisos Mts., Brewster Co.: 1 36, VII-4 to 6-61 (R. L. Westcott); 4 ¢ ¢,2

2 @, VII-16 to 17-73 (F. T. Hovore, beaten ex Quercus). Davis Mountain

State Park, Jeff Davis Co.: 1 2, VII-18 to 21-73 (F. T. Hovore, at light).

Poliaenus nuevoleonis similnegundo, new subspecies

Elytral costae usually subdued. Pubescence not obscuring basal elytral

punctures, integument sufficiently visible to give dorsum a dark brown ap-

pearance. Appressed white pubescence covering at most basal half of an-

tennal segments four to ten. Length 6.1-10.9 mm.

Known habitat, range, and flight period.—Oak woodland of southern

Arizona (May to September) and Durango, Mexico (July).

Material examined.—Holotype female (California Academy of Sciences

Type #13247) from Santa Rita Lodge, elevation 4960 feet, VII-14-75, Ma-

dera Canyon, Santa Cruz Co., Arizona (D. D. Skiles at light); allotype

(CAS), VII-4-72, Madera Canyon, Santa Cruz Co., Arizona (D. G. Marqua).

Paratypes. ARIZONA. Pima Co.: Kitt Peak, 6000 feet, Baboquivari Mts.,

1 2, VII-9-78 (D. Skiles, at light); Sabino Canyon, 1 ¢, [X-4-61 (J. S.

108 PAN-PACIFIC ENTOMOLOGIST

Buckett). Cochise Co.: Miller Canyon, Huachuca Mts., 1 3d, VII-18-71 (D.

G. Marqua), 1 6, VII-12-75 (E. F. Giesbert); Carr Canyon, Huachuca Mts.,

1 2, [X-4-59 (R. L. Westcott); Cave Canyon, Huachuca Mts., 1 2, VIII-6-

78 (D. Skiles, beaten ex dead Quercus hypoleucoides); Cochise Stronghold,

Dragoon Mts., 1 °, VII-12-77 (D. Skiles at light), Cave Creek Ranch, Chi-

ricahua Mts., 1 6, VIII-19-65 (G. W. Forister, at light); 5 mi W Portal,

Chiricahua Mts., 1 3, VIII-14-58 (G. G. Moore). Santa Cruz Co.: Santa

Rita Mountains, 6000 feet, 1 ¢6, [X-15-33 (Bryant, Lot 238; labeled ‘‘Poli-

aenus negundo (Schaeffer)/Det. Knull °56’’; also labeled ‘‘Pogonocherus

arizonicus Schaeffer’’). Madera Canyon, Santa Rita Mountains, 1 6, [X-4-

66, 146,12, [X-5-66 (M. E. Pendleton): 2 ¢ 6, VIII-16-67 (C. D. Johnson,

at light); 1 ¢d, 1 2, [X-3 to 5-69 (J. Powell, at light); 1 2, VII-23 to 25-58,

1 2, [X-2-59 (R. L. Westcott); 1 9, [X-3-64 (G. H. Nelson, at light); 1 °,

IX-5-59 (L. M. Martin, at light); 1 2°, V-22-63 (J. G. Franclemont, 4880

feet); 1 2, [X-4-67 (A. S. Menke, 4880 feet); 1 ¢, [X-29-63 (V. L. Westerby,

4880 feet); 1 2, VII-1-63, 1 od, VII-2-63, 1 2, VII-5-63 (J. D. Marshall, 4880

feet); 1 2, VIII-6-69, 1 2, VIII-71, 1 2, VIII-28-71 (E. F. Giesbert, at light);

3 6 6, VII-23 to 24-71, 1 6, VIII-28-71, 2 2 2, VIII-8-77,2 6 6,1 2, VII-

12-78 (F. T. Hovore at light); 1 ¢, 1 2, VII-27-76 (F. T. Hovore, beaten

together ex dead Quercus). 1 9, VII-23-74, 1 2, VII-9-75, 1 °, VII-14-75,

1 3, IX-12-75, 1 ¢, [X-20-75, 1 2, VII-10-77, 1 36, VII-12-77 (D. Skiles, at

light and beaten ex dead Quercus); 17 2°, VII-7, 9, 19, 29-71, IX-7-71,

VII-11, 12, 18-72, VIII-4, 8-72, VII-21-73, VII-20-74, VII-15-75, VITI-15-75,

66 6, VII-18, 19, 28-71, [X-7-71, VII-14-75, VIII-15-75 (D. G. Marqua, at

light).

Also assignable to this subspecies but not included as paratypes: 1 6,

Huachuca Mts., Arizona (Van Dyke collection); 1 ¢, 3 mi E El Salto,

Durango, Mexico, VII-3-64 (L. A. Kelton).

Diagnosis.—The fact that P. nuevoleonis, though rather common, re-

mained undescribed until recently is undoubtedly due to its remarkable re-

semblance to P. negundo (Schaeffer) and to the fact that both species are

frequently taken together at light in oak woodlands throughout southeastern

Arizona.

Presumably, these congeners are able to coexist by utilizing different

hosts. Poliaenus negundo is known to attack sumac and box elder (Linsley,

1935), and P. nuevoleonis probably attacks various species of oak, since a

small number of specimens have been beaten from dead, leaved branches

of Quercus sp. in the Davis and Chisos Mountains of Texas (F. T. Hovore,

E. F. Giesbert) and from dead, leaved branches of Q. hypoleucoides A.

Camus in the Santa Rita and Huachuca Mountains of southern Arizona (F.

T. Hovore, D. Skiles). Beyer (1908) reported that four specimens of P.

negundo from Arizona’s Huachuca Mountains emerged in 1907 from oak

twigs girdled in 1905 by Oncideres quercus Skinner. However, as I am

VOLUME 55, NUMBER 2 109

unaware of any other records of P. negundo from oak, I am inclined to

believe that Beyer in fact reared P. nuevoleonis.

Poliaenus nuevoleonis and P. negundo can usually be distinguished with

the unaided eye by the different color patterns of the elytral pubescence. In

the former the pattern is very faint and the elytra appear rather uniformly

brown with a slight hoary wash. In negundo the dark brown base of the

elytra contrasts sharply with the dense, yellowish-brown pubescence of the

antemedian V-shaped impression. Despite this difference, specimens of

P. nuevoleonis are easily mistaken for slightly rubbed specimens of P.

negundo.

Under the microscope the two species are readily distinguished by the

distinct structures of the pronotal disc. In nuevoleonis the discal tubercles

can best be described as circular cones arising from a flat pronotal disc; in

negundo the discal tubercles are little more than the lateral limits of a discal

pronotal gibbosity. In addition, the flying hairs on the dorsal surface of the

posterior tarsi are brown intermixed with white in nuevoleonis, whereas

they are almost invariably entirely white in negundo, though they may ap-

pear dark when illuminated from certain angles.

From the long series of P. nuevoleonis | have examined, it is apparent

that P. nuevoleonis, P. sparsus Chemsak and Linsley, and P. batesi Linsley

are closely related and may even be conspecific. However, the latter two

species are known only from single specimens so that a definitive analysis

of the specific status of each must await the collection of further material.

I have examined the type of P. sparsus (California Academy of Sciences)

and a 35 mm color slide of the type of P. batesi (courtesy E. G. Linsley

and J. A. Chemsak) and offer the following observations.

The structural similarities between the diminutive type of P. sparsus and

smaller specimens of P. nuevoleonis similnegundo leave me with the

impression that the two species may ultimately prove to be one. Neverthe-

less, when compared side by side with a series of small P. nuevoleonis

similnegundo, the specimen of P. sparsus immediately stands apart as hav-

ing rather testaceous integument rather densely covered with golden pu-

bescence, as opposed to dark brown integument rather sparsely covered

with grey pubescence. In addition, P. sparsus is slightly smaller than the

smallest paratypical P. nuevoleonis similnegundo, and the basal elytral

punctures of the former are distinctly coarser. Hence the two species cannot

be synonymized on the basis of the available material.

On the other hand, I find it difficult to believe that P. nuevoleonis and P.

batesi are specifically distinct. Specimens of nuevoleonis, particularly

densely pubescent ones such as those I have seen from the Chisos Moun-

tains of Texas, key to batesi in Linsley’s (1935) key to the genus. In addi-

tion, although I have seen only a slide of the unique specimen of batesi

from central Guatemala, the slide clearly shows all diagnostic characters,

110 PAN-PACIFIC ENTOMOLOGIST

and the specimen differs from large, densely pubescent specimens of nue-

voleonis only in the more prominent and dramatically orange penicilli of the

subsutural elytral costae. However, the subsutural penicilli of nuevoleonis,

while appearing dark brown or black to the naked eye, are in fact often

margined with orange. This is particularly true of the postmedian penicilli.

Hence the orange penicilli of batesi, which under the microscope are seen

to have dark brown centers at the base of the elytra and a few dark brown

hairs on the apical third of the elytra, merely appear to be extreme versions

of a color tendency apparent in nuevoleonis.

Acknowledgments

I am indebted to E. G. Linsley, without whose generous assistance and

valuable comments this paper would not have been possible, and to J. A.

Chemsak for reviewing the manuscript. I am grateful to F. T. Hovore, E.

F. Giesbert, and D. G. Marqua for important collecting data and the loan

of specimens, and to the following institutions and individuals for the loan

of specimens and permission to study type material: The Essig Museum of

Entomology, University of California, Berkeley (J. A. Chemsak and E. G.

Linsley), the California Academy of Sciences, San Francisco (D. H. Ka-

vanaugh), the Los Angeles County Museum, Los Angeles, California (C. L.

Hogue). F. G. Taylor and R. Doane kindly arranged for me to collect on

the grounds of Kitt Peak National Observatory.

Literature Cited

Beyer, G. 1908. Notes on Oncideres quercus Skinner. J. New York Entomol. Soc. 16: 32.

Chemsak, J. A., and E. G. Linsley. 1975. Mexican Pogonocherini. Pan-Pac. Entomol. 51:

271-286.

Linsley, E. G. 1935. A Revision of the Pogonocherini of North America. Ann. Entomol. Soc.

Amer. 28: 73-103.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 111-116

GENETIC CONFIRMATION OF THE SPECIFIC STATUS OF THE

SPEYERIA ADIASTE GROUP IN CALIFORNIA

(LEPIDOPTERA: NYMPHALIDAE)

STEVEN R. Sims!

Dept. of Entomol., Univ. Calif., Davis, 95616

JOHN G. BRITTNACHER AND FRANCISCO J. AYALA

Dept. of Genetics, Univ. Calif., Davis, 95616

The Speyeria adiaste Edwards group is composed of three closely related

subspecies occurring in suitable habitats in the Coast Ranges and extreme

southern Sierra Nevada of California. These subspecies are well illustrated

on Plate 24 in The Butterflies of North America (Howe, 1975). Speyeria

adiaste was described by W. H. Edwards (1864) and the type locality was

fixed by DosPassos and Grey (1947) as the Santa Cruz Mountains, Califor-

nia, where existing populations of S. adiaste adiaste, the northernmost and

darkest member of the group, occur. Speyeria adiaste clemencei (Com-

stock) has a lighter fulvous ground color and less pronounced dark markings

on the upper wing surface than the nominate species (Comstock, 1925). It

occurs in the Coast Ranges from Monterey Co. to near the town of San

Luis Obispo. The now extinct S. adiaste atossa (Edwards) was recorded

in the Tehachapi and Tejon Mountains and in the Mt. Pinos region. In

overall appearance it resembled its northern counterparts except for a clear

yellow-brown ground color and further reduction of the upper surface mark-

ings.

These three geographically disjunct subspecies form a color cline ranging

from the dark northern S. adiaste adiaste to the pale southern S. adiaste

atossa. Although not yet determined for the adiaste group, several other

subspecies groups in the genus Speyeria (callippe, coronis, zerene) display

extremely close genetic similarity despite evident phenotypic change (Britt-

nacher et al., 1978). It is likely that the adiaste group fits into this pattern

which suggests a rather recent evolutionary divergence of the different color

forms.

The distributional limits of these subspecies seem, in part, to be deter-

mined by the availability of their violet (Viola) food plants and by desic-

cation tolerances of first instar larvae to dry diapause period (summer-fall)

conditions (Sims, unpublished data). Xeric conditions tend to limit distri-

butions of many violet species and prove fatal to species lacking adequate

desiccation resistance. Speyeria adiaste may once have occurred in an un-

broken range throughout the coastal mountains. Division of the range was

112 PAN-PACIFIC ENTOMOLOGIST

possibly influenced by the drier climatic conditions of the Pliocene (Axelrod,

1948) or the pluvial periods of the Pleistocene. The warmer and drier post-

Pleistocene conditions would have supplemented the process of range lim-

itation.

Since the original description, the taxonomic status of S. adiaste adiaste

and the later named subspecies has been in doubt. The most recent system-

atic treatment of the genus Speyeria (DosPassos and Grey, 1947) regards

the adiaste complex as a subspecies of S. egleis (Behr). Although S. egleis

exhibits a much wider distribution than S. adiaste, the groups are (or were)

completely allopatric except in the Tehachapi Mountains. In this latter lo-

cation, the species populations were spatially isolated by elevation, S. egleis

preferring the higher peaks and slopes while S. adiaste atossa frequented

mid-elevation habitats. Other interspecific barriers to reproduction might

well have included differences in species specific pheromones or mating

behavior (Magnus, 1958).

Phenotypically, specimens within populations of the three taxa in the

adiaste group are quite uniform in contrast to many other species in the

genus (Moeck, 1957). Speyeria egleis often exhibits remarkable intrapopu-

lation variation both in coloration of the disc (basal undersurface of hind

wing) and in the silvering or absence of silvering of the hind wing spots.

The taxonomic relationship of S. adiaste has recently undergone another

shuffling in which the group regained specific status (Emmel and Emmel,

1973; Howe, 1975). The purpose of this paper is to present genetic evidence

which ex post facto justifies this latest separation of the S. adiaste group.

Genetic Differentiation

Gel electrophoretic techniques are well known and currently widely used

to study inter- and intraspecific levels of genetic variability in diverse groups

of organisms. Techniques of gel electrophoresis and enzyme assay allow

identification of allelic variation at single gene loci. When data are obtained

from a ‘‘moderate’’ number of enzyme loci, the results, with certain as-

sumptions, may be extrapolated to the genome as a whole. The loci studied

are assumed to represent a random sample of the genome with respect to

allelic variation. Possible sources of bias in such an assumption have been

cited by Lewontin and Hubby (1966) and Ayala et al. (1970). It has been

amply demonstrated that gel electrophoresis is an extremely valuable sys-

tematic tool (see Avise (1974) for review). Ayala (1973) and Ayala and Dob-

zhansky (1974) have used allozyme data as diagnostic characters for sub-

species of Drosophila willistoni and D. pseudoobscura.

We examined genetic variation at 16 loci in all ten species of Speyeria

occurring in California plus several of the subspecies existing in the callippe,

coronis, hydaspe, and zerene groups (Brittnacher et al., 1978). With few

VOLUME 55, NUMBER 2 113

Table 1. Genetic similarity (above diagonal) and genetic distance (below diagonal) for seven

species in the genus Speyeria.

1 2 3 4 5 6 x

1. adiaste tip .866 .798 .852 Be .762

2. atlantis 255 913 954 .950 .801 -922

3. callippe .144 .091 .933 .985 .881 .901

4. coronis LoS .047 .069 .938 .883 .903

5. egleis .161 .0S1 .O15 .064 .872 .938

6. hydaspe .087 22 .126 125 138 .790

7. zerene Bee e: .081 .104 .102 .064 236

exceptions, only males were used in the assays. Females were used in a

companion study of the reproductive biology of the genus.

Populations of S. egleis egleis from the following Sierra Nevada locations

were analyzed: Bowman Lake, Nevada Co., el. 1700 m (n = 8); Donner

Pass, Placer Co., el. 2100 m (n = 5); and Yuba Pass, Sierra Co., el. 2000

m(n = 10). Two populations of S. adiaste clemencei were sampled: Arroyo

Seco Camp, el. 260 m (n = 19) and Chew’s Ridge, el. 1100-1500 m (n =

45), both in the coast range of Monterey Co. California.

Speyeria adiaste clemencei and S. egleis egleis were found to have two

fixed differences at the sixteen loci studied. These were for glyceraldehyde-

3-phosphate dehydrogenase and glucose-6-phosphate dehydrogenase (see

Brittnacher et al., 1978, for details).

The amount of genetic differentiation between S. adiaste and S. egleis

is substantial when compared to the genetic differentiation between other

species of Speyeria. Table 1 summarizes the differentiation found in Spey-

eria using Nei’s (1972) method of calculating genetic distance, D, and ge-

netic identity, 7. It can be seen that the genetic distance between S. egleis

and S. adiaste is greater than the distance between S. egleis and S. atlantis,

S. callippe, S. coronis, and S. zerene. It is thus unlikely, based on this

genetic evidence alone, that the members of the S. adiaste group are sub-

species of S. egleis.

Karyotype Determination

Chromosome numbers of S. adiaste clemencei and S. egleis egleis were

determined in view of the potential for additional substantiation of the allelic

differences. Chromosome counts were made in 19 nuclei in testes of 6 S.

adiaste clemencei males field collected at Chew’s Ridge on June 11, 1974.

Counts were similarly made in 10 nuclei in testes of 3 S. egleis egleis pupae

derived from the population at Loon Lake, El Dorado Co., California.

Cytological techniques followed for S. adiaste clemencei involved fixa-

114 PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Chromosomes of Speyeria, (A) S. adiaste clemencei, N = 29, metaphase; (B) S.

egleis egleis, N = 29, metaphase.

tion for 5 minutes in a 3:1 absolute ethanol:glacial acetic acid solution,

staining with 0.5% lacto-acetoorcein, and squashing on a slide using hand

pressure (Emmel, 1968). A modified squash-air dry technique described by

Goodpasture (1976) was used for chromosome counts of S. egleis egleis.

Preparations were examined under oil using phase contrast illumination at

a magnification of 960. Photographs were taken on Kodak High Contrast

Copy 35 mm film at a film plane magnification of 400.

In all sufficiently clear preparations, the haploid number (N) was found

to be 29 for both S. adiaste and S. egleis (Fig. 1). This count is identical

to the majority of previously determined species in the genus (Maeki and

Remington, 1960). Exceptions occur in the S. callippe and S. coronis groups

where apparently N = 30. Curiously, S. callippe and S. egleis are cytolog-

ically distinct despite having the highest genetic similarity value (.985) of all

Species studied.

Immature Stages

Little other comparative biological data is available on the S. adiaste and

S. egleis groups. Edwards (1897) described and illustrated the life history

of S. egleis from Colorado. Comstock and Dammers (1931) described the

mature larva and pupa of S. adiaste atossa from Lebec, Kern Co., Cali-

fornia. Adequate distinguishing characters cannot be determined from the

descriptions except for the slightly larger mature size of the S. adiaste

atossa larva (35 mm vs. approx. 31 mm for. S. egleis) which may simply be

a sex-related difference. We believe it significant to note that mature larvae

of both are characterized by an irregular yellowish patch on the dorsum of

the head capsule, a trait missing in California S. callippe and S. coronis.

VOLUME 55, NUMBER 2 115

Summary

Speyeria adiaste clemencei, long considered a member of the S. egleis

group, shows a relatively low degree of genetic similarity (I = .852) to this

species. Two fixed differences were present among sixteen loci studied.

From a perspective of genetic relationships in other Speyeria this diver-

gence appears significant at the species level. The marked phenotypic dis-

tinctiveness of the Speyeria adiaste group, uniformity of phenotype within

populations, geographic isolation, and lack of hybridization are additional

factors arguing for recognition of specific distinction. The chromosome num-

ber (N = 29) of both S. adiaste and S. egleis is similar to the majority of

other Speyeria and does not provide an adequate index of relationship. Both

groups have similar immature stages with only minor color differences. In

these two species, it appears that allozyme characters are better differen-

tiated than chromosome number or immature stage morphology.

Acknowledgments

We thank Sterling O. Mattoon for providing pupae of S. egleis egleis and

Dr. A. M. Shapiro for comments on the manuscript.

Literature Cited

» Avise, J. C. 1974. Systematic value of electrophoretic data. Syst. Zool. 23: 465-481.

Axelrod, D. I. 1948. Climate and evolution in western North America during middle Pliocene

time. Evolution 2: 127-144.

Ayala, F. J. 1973. Two new subspecies of the Drosophila willistoni group (Diptera: Drosoph-

ilidae). Pan-Pac. Entomol. 49: 273-279.

Ayala, F. J., and T. Dobzhansky. 1974. A new subspecies of Drosophila pseudoobscura

(Diptera: Drosophilidae). Pan-Pac. Entomol. 50: 211-219.

Ayala, F. J.,C. A. Mourao, S. Pérez-Salias, R. Richmond, and T. Dobzhansky. 1970. Enzyme

variability in the Drosophila willistoni group. I. Genetic differentiation among sibling

species. Proc. Nat. Acad. Sci., USA 67: 225-232.

Brittnacher, J. G., S. R. Sims, and F. J. Ayala. 1978. Genetic differentiation between species

of the genus Speyeria (Lepidoptera: Nymphalidae). Evolution 32: 199-210.

Comstock, J. A. 1925. Studies in Pacific Coast Lepidoptera. Bull. So. Calif. Acad. Sci. 24:

3-4,

Comstock, J. A., and C. M. Dammers. 1931. Notes on the early stages of four California

Argynnids (Lepid.). Bull. So. Calif. Acad. Sci. 30: 40-48.

DosPassos, C. F., and L. P. Grey. 1947. Systematic catalogue of Speyeria (Lepidoptera,

Nymphalidae) with designations of types and fixations of type localities. Amer. Mus.

Nov. 1370. 30 p.

Edwards, W. H. 1864. Notes on the Argynnides of California. Proc. Entomol. Soc. Phil. 3:

434-436.

Edwards, W. H. 1897. Butterflies of North America. Houghton Mifflin and Co., New York.

Ser. 3, pt. IX.

Emmel, T. C. 1968.: Methods for studying the chromosomes of Lepidoptera. J. Res. Lepid.

7: 23-28.

116 PAN-PACIFIC ENTOMOLOGIST

Emmel, T. C., and J. F. Emmel. 1973. The butterflies of Southern California. Nat. Hist. Mus.

of Los Angeles Co., Sci. Series 26. 148 p.

Goodpasture, C. 1976. High-resolution chromosome analysis in Lepidoptera. Ann. Entomol.

Soc. Amer. 69: 764-771.

Howe, W. H. 1975. The butterflies of North America. Doubleday & Co., New York. 633 p.

Lewontin, R. C., and J. L. Hubby. 1966. A molecular approach to the study of genic hetero-

zygosity in natural populations. II. Amount of variation and degree of heterozygosity

in natural populations of Drosophila pseudoobscura. Genetics 54: 595-609.

Maeki, K., and C. L. Remington. 1960. Chromosomes of North American Lepidoptera. Part

4. J. Lepid. Soc. 14: 179-201.

Magnus, D. 1958. Experimental analysis of some ‘‘overoptimal’’ sign-stimuli in the mating

behaviour of the fritillary butterfly Argynnis paphia L. (Lepidoptera: Nymphalidae).

Proc. 10th Int. Congr. Entomol. 2: 405-418.

Moeck, A. H. 1957. Geographic variability in Speyeria. Comments, records and description

of a new subspecies. Privately printed, sponsored by Milwaukee Entomol. Soc. 48 p.

Nei, M. 1972. Genetic distance between populations. Amer. Natur. 106: 283-291.

Footnote

1 Current Address: Dept. of Biol., Univ. Notre Dame, Notre Dame, IN, 46556.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 117-126

EMIGRATION RESPONSE BEHAVIOR: II: THE RESPONSES OF

DROSOPHILA BUSCKI (DIPTERA: DROSOPHILIDAE)'

ROBERT F. ROCKWELL

Department of Biol., The City College, Convent Ave. at 138th St.,

New York, NY, 10031

The movement of members of a population from one geographic locality

to another is an important factor in the evolutionary process. It is clear that

not all members of a given population emigrate. Of those members which

do move, some are compelled by genetic factors, some by environmental

factors and some by the interaction of both. While emigrants must be ca-

pable of surviving and reproducing in the new localities, fine level adapta-

tions to a new area evolve only after arrival. Thus, the possibility of ex-

tending the range of a species depends on the ability of the genetic

architecture of the emigrants to respond to a new environment in addition

to their propensity to move from an old one. It follows that the examination

of genetic and environmental factors influencing movement from one local-

ity to another is of prime importance to evolutionary and behavioral biology.

This is a report from an ongoing study that is examining the interacting

effects of genotypes and environments on the movement of Drosophila from

their place of origin to new locations. The active change in location of

members of a population results from two distinguishable processes, name-

ly, migration and dispersal. While these terms have had various meanings,

Rockwell et al. (1978) encouraged the use of the following definitions: Mi-

gration—the goal oriented movement of a fraction of the population; Dis-

persal—the movement of a fraction of a population as a result of the general

(random) activity of its members. The overall movement of organisms from

their place of origin to a new location is termed emigration response be-

havior.

Emigration response behavior of Drosophila has been examined in the

laboratory by a number of researchers who have used a system devised by

Sakai et al. (1958). That system consists of a set of four interconnected vials

through which individual flies can move (Fig. 1). Emigration response be-

havior is measured as the number (or percentage) of individuals leaving the

central vial. Rockwell et al. (1978) pointed out that this single measurement

of emigration response confounds migration and dispersal. They proposed

that by measuring emigration responses in the Sakai system relative to four

specific environmental configurations, one could obtain a clearer view of

the relative contributions of migration and dispersal to the overall emigration

response behavior of a species (or strain).

118 PAN-PACIFIC ENTOMOLOGIST

Fig. 1. The emigration response behavior apparatus modeled after Sakai et al. (1958). A—

the arrangement of vials; B—low vial; C—high vial.

The specific environmental configurations are the four possible combi-

nations of two levels of each of two factors, namely height of the connecting

tubes and lighting condition. The original Sakai system utilizes connecting

tubes which are about 4 cm above the level of the food surface in the vials

(Fig. 1C). As such, passage from the central vial to the three peripheral vials

can be viewed as three-dimensional; that is, requiring movement through a

volume. The use of vials whose connecting tubes are at the same level as

the food surface changes this to two-dimensional movement involving only

VOLUME 55, NUMBER 2 119

area. If the passage of flies through this system contains a general activity

component, one would expect substantially more emigration in the vials

with the low connecting tubes simply because the flies would have more

chance, in a fixed time period, to encounter the connecting tubes. Thus, the

greater the difference between the emigration responses measured in the

two types of vials (high versus low), the greater is the dispersal component

of the overall emigration response behavior.

The role of light as a stimulus that increases general activity in Drosophila

has been extensively noted (Grossfield, 1971). If light plays this role in the

Sakai system and if the passage of flies through this system contains a

general activity component, one would expect substantially more emigration

in the light than in the dark. Thus, the greater the difference between the

emigration responses measured in the light versus dark, the greater the

dispersal component of the overall emigration response behavior.

The present paper focuses on the emigration response behavior of Dro-

sophila busckii. This species, like D. melanogaster has a global distribution.

However, there is a phylogenetic peculiarity of the genus in that it is the

only member of its subgenus dorsilopha (Throckmorton, 1975). In addition,

D. busckii possesses an inordinately low level of genic variation (and,

hence, heterozygosity) which contrasts it not only with the rest of the genus

but with many other organisms (Prakash, 1973). The inclusion of this species

in the overall study of the interacting effects of genotype and environment

on the movement of Drosophila from one locality to another is especially

important for two reasons. First, because of the taxonomic and genetic

distinctness of this species in the genus, the evaluation of its emigration

response behavior will provide information on the general applicability of

this experimental system, and the conclusions reached using it, to studies

of interlocality movement in the genus.

Second, because of the low heterozygosity of D. busckii, the extent to

which the emigration response changes across the four environmental con-

figurations will provide important information on the general relationship

between behavioral plasticity (i.e., flexibility) and genetic architecture.

While it is widely held that the level of plasticity of a given behavioral trait

is related to underlying genetic architecture such as the level of heterozy-

gosity (Dobzhansky, 1973), most studies on the nature of that relationship

have only used species that are quite heterozygous in nature (Caspari, 1967;

Rockwell et al., 1975).

Methods and Materials

Stocks and Culturing

The strain of Drosophila busckii used in these experiments was collected

at Wyeth Farm, Glenburnie, Ontario, in the fall of 1975. Since that time, it

120 PAN-PACIFIC ENTOMOLOGIST

has been maintained as a mass culture in % pint bottles on a medium com-

posed of equal volumes of instant mashed potatoes and water (Tegosept®

is added to reduce mold contamination). The stock is maintained at 20.0°C,

85% relative humidity and constant illumination.

Equipment

The migration vials are modeled after those of Sakai et al. (1958) and are

depicted in Figure 1. Two types of vials were used:

a) high vials (Fig. 1C) in which the connecting tubes are 4.0 cm above the

surface of the medium;

b) low vials (Fig. 1B) in which the connecting tubes are at the same level

as the surface of the medium.

As depicted in Figure 1A, four high vials or four low vials were connected

with clear tape to form a single migration system. The unused connecting

tubes of the three peripheral vials were plugged with corks. A mixture of

instant mashed potatoes and water (1:1) was used as the medium. The sys-

tems were assembled one hour before the beginning of experimental trials.

Each assembled system was placed in an open plastic tray measuring 28 x

28 x 16 cm high. The inside of each of these trays had been painted white

and the outside had been painted black. The painting of the trays served to

eliminate extraneous visual cues and to provide diffuse uniform illumination.

The trays containing the migration systems were placed in a 20.0°C envi-

ronmental chamber.

Procedure

Flies were collected daily from replicate culture bottles and batches of 20

males and 20 females were placed together in 8 dram food vials. These were

stored in an incubator at 20.0°C, 85% relative humidity and constant illu-

mination. A set of 50 males between 7 and 10 days old was randomly col-

lected from these vials and placed in an empty 8 dram vial for 30 minutes.

Humidified CO, was used as the anesthesia for these procedures.

A given set of 50 males was aspirated into the central vial of either a high

or a low migration system. If the system was to be tested in the dark, the

tray containing the system (and the flies) was immediately covered with

black cloth. Preliminary studies demonstrated that no light entered such

trays. If the system was to be tested in the light, the tray containing the

system was placed beneath a fluorescent light fixture (consisting of two 40

watt tubes suspended 24 inches above the tray).

After inserting the flies and placing the trays in the appropriate lighting

condition, the system was left undisturbed for 24 hours (until 1000 hours on

the following day). At that time, the number of flies in the three peripheral

VOLUME 55, NUMBER 2 121

vials was determined. The total number of flies in the three peripheral vials

was used as the measure of emigration response for a given configuration.

The measure can range from 0 to 50.

Experimental Design and Statistical Procedures

In order to examine the relationship between height and lighting condi-

tion, the experiments were performed in a factorial fashion. Three replicates

of each combination of the levels of the two factors were performed in a

randomized blocks fashion. Preliminary analysis of the data demonstrated

no effects of blocks, so the replicate sources of variation were pooled (Win-

er, 1971). The data were then analyzed with factorial analysis of variance.

The factorial design and analysis were used to assess whether the factors

affect the emigration response behavior and to determine whether the two

factors interact.

The emigration responses of D. busckii were compared to those of D.

melanogaster with factorial analysis of variance. The emigration responses

of D. melanogaster used in that comparison were measured under condi-

tions identical to those just described and formed a part of a study reported

in Rockwell et al. (1978).

Results

The emigration responses of male D. busckii in the four environmental

configurations are given in Table 1 as means with their associated standard

errors. The responses were analyzed with factorial analysis of variance and

the results of that analysis are summarized in Table 2. There is a highly

significant effect of tube height overall; the emigration response is greater

in the low tubes. There is no significant effect of lighting condition on the

emigration response. Importantly, there is no significant interaction between

tube height and lighting condition; tube height modulates the response

equivalently in both the light and the dark.

The emigration responses of D. busckii were compared to those of D.

Table 1. The emigration response behavior of male Drosophila busckii measured in the

four conditions of the Sakai system.

Tube height

Lighting

condition Low High

Constant light 4137 2. 1,76 20,33 +233

Constant dark 36.33 + 4.70 19.33 + 2.84

NB: +Standard errors.

122 PAN-PACIFIC ENTOMOLOGIST

Table 2. Analysis of variance of the emigration response behavior of male Drosophila

busckii.

Degrees of Mean

Source of variation freedom Square

Tube height (H) 1 588.007

Lighting condition (L) 1 12.00

H* L 1 27.00

Error 8 29.08

NB: All sources were tested over the error term.

2 Significant at the 0.01 level of probability.

melanogaster using the three factor analysis of variance summarized in

Table 3. Examining the main effects first, it is clear that there is no overall

difference between the two species. That is, the emigration response be-

haviors of the two species, averaged across the four environmental config-

urations, do not differ significantly. In sharp contrast, the overall emigration

responses for the two tube heights, averaged across lighting conditions and

species, are quite significantly different. The emigration response in the low

tubes is twice that in the high tubes (34.15 versus 16.99). The overall emi-

gration responses for the two lighting conditions, averaged across tube

height and species, are significantly different. The emigration response in

the dark is greater than that in the light (28.57 versus 22.57).

Of crucial importance is the highly significant three-way interaction be-

tween tube height, lighting condition and species. Given the simultaneous

Table 3. Analysis of variance of the emigration response behavior of male Drosophila

busckii and D. melanogaster.

Degrees of Mean

Source of variation freedom square

Tube height (H) 1 1768.17?

Lighting condition (L) 1 216.00°

Species (S) 1 37.50

HxL 1 216.00°

HxS 1 60.16

LxsS 1 96.00

HxLxS& 1 486.00?

Error 16 44.99

NB: All sources were tested over the error term.

2 Significant at the 0.01 level of probability.

3 Significant at the 0.05 level of probability.

VOLUME 55, NUMBER 2 123

occurrence of a significant interaction between tube height and lighting con-

dition and the lack of any significant interaction between tube height and

species, the three-way interaction may reasonably be interpreted as a non-

additive effect between the two species for the tube height by lighting con-

dition interaction. That is, the interrelationship between tube height and

lighting condition, in their joint effect on emigration response, differs be-

tween the two species.

This interpretation is further supported by comparing the separate anal-

yses of tube height and lighting condition effects for the two species. In the

factorial analysis of tube height and lighting condition effects on the emi-

gration response behavior of male D. melanogaster (Rockwell et al., 1978),

it was shown that the tube height by lighting condition interaction is highly

significant. For D. melanogaster, then, the effects of tube height and lighting

condition are not independent; tube height modulates the response differ-

entially with light. Recalling the analysis summarized in Table 2, tube height

modulates the emigration response of D. busckii equivalently in the light

and in the dark. Thus, the interdependence of tube height and lighting con-

dition on the emigration response behavior of these two species is different.

Discussion and Conclusions

It is clear that tube height modulates the emigration response behavior of

Drosophila busckii. The greater emigration response displayed in the low

tubes is consistent with the existence of a general activity component in the

overall emigration response behavior of this species. As explained earlier,

such a component would be expected to result in more two-dimensional

movement than three-dimensional movement.

There is no apparent effect of lighting condition on the emigration re-

sponse behavior of this species; the emigration response is the same in

constant light and constant dark. This result is not in agreement with the

accentuating effect of light on general activity widely noted for Drosophila

(Carpenter, 1905; Manning, 1965). This difference may reflect a peculiarity

of the overall general activity behavior of D. busckii. It may also reflect the

existence of several general activity programs in this species, each of which

may serve as a component of different overall behavior systems. While the

latter explanation is consistent with the results of experiments with D. mel-

anogaster, this entire question is still under investigation.

In general the emigration response behavior of D. busckii measured in

the Sakai system is a composite of both dispersal and migration components.

In that sense, the overall emigration response is like that of D. melanogas-

ter. It is clear, however, that only by assessing the emigration responses

under the four environmental configurations can the underlying nature of

the emigration response behavior be ascertained for either species. As will

124 PAN-PACIFIC ENTOMOLOGIST

be discussed below, it is this underlying nature which is crucial for inter-

specific comparisons.

These two species do not differ significantly in their emigration response

behaviors averaged across the four environmental configurations. While in-

terspecific differences can be shown for the emigration response measured

in specific configurations (e.g., D. melanogaster has a greater response than

D. busckii for the high tubes in the dark), the crucial difference between

these species derives from the comparison of the response spectra of their

emigration response behaviors. The response spectrum of a behavior is the

plasticity of the measured behavioral response with respect to specified

environmental perturbations. Behavioral plasticity is the tendency of a be-

havioral phenotype to change in form or intensity in response to alterations

in the environment. Such plasticity reflects the norm of reaction of the

genetic system underlying the behavior in question (Dobzhansky, 1970).

Rockwell and Seiger (1973) pointed out that the response spectrum of a

given behavior is most important in research directed at the elucidation of

the underlying mechanisms or evolutionary significance of a given behavior.

The emigration response behavior of D. busckii is plastic with respect to

tube height but not with respect to lighting condition. The emigration re-

sponse behavior of D. melanogaster is plastic with respect to both tube

height and lighting condtion. Considering the four environmental configu-

rations, then, it appears that the emigration response behavior of D. busckii

is less plastic than that of D. melanogaster. It follows that with respect to

these environmental perturbations, the norm of reaction of D. busckii is

narrower than that of D. melanogaster for emigration response behavior.

Considering the joint effects of the two factors on the response, it also

appears that the plasticity (and, hence, norm of reaction) of D. busckii is

less complex. It will be recalled that in D. busckii the effects of tube height

were independent of lighting condition. In D. melanogaster, the effect of

tube height was modulated by lighting condition.

Overall the norm of reaction of D. busckii appears to be narrower and

less complex than that of D. melanogaster for this behavior. There is no

doubt that such differences in plasticity and norm of reaction are related to

differences in genetic architecture between these two species. The results

presented here are consistent with the possibility that the reduced plasticity

of this behavior in D. busckii derives directly from the reduced heterozy-

gosity of this species. Formal demonstration of that specific relationship

awaits further investigation.

Since plasticity is, in general, a product of the genetic architecture un-

derlying a given behavioral trait, the level of plasticity must ultimately de-

rive from the evolutionary history and ecological requirements of a species.

Similarly, interspecific differences in plasticity must derive from interspe-

cific differences in these two factors. In fact, a portion of interspecific dif-

VOLUME 55, NUMBER 2 125

ferences in genetic architecture (and plasticity) may reflect the action of

selection directed at the level of behavioral plasticity itself (Emlen, 1973;

Rockwell and Cooke, 1977). It is tempting to relate the differences in plas-

ticity demonstrated in this work to differences in the evolutionary histories

and ecologies of these two species. Such speculation, however, must await

further clarification of the evolutionary role of dispersal and migration in

this genus. It is clear from this work that studies attempting such clarifi-

cation must consider the plasticity and, hence, the norm of reaction of

emigration response behavior to be at least as important as the overall mean

behavior.

Acknowledgments

I wish to thank John Emlen, Joe Griswold, Louis Levine and David Miller

for their useful comments on an earlier version of this paper. This work was

supported in part by a PSC-BHE award (RF11611) and through the good

graces of James Organ, Chairman of Biology, City College.

Literature Cited

Carpenter, F. W. 1905. The response of the pomace fly to light, gravity and mechanical

stimulation. Amer. Nat. 39: 157-171.

Caspari, E. 1967. Gene action as applied to behavior. In Behavior Genetic Analysis, ed. J.

Hirsch. McGraw-Hill, New York.

Dobzhansky, Th. 1973. Genetics of the Evolutionary Process. Columbia University Press,

New York.

Emlen, J. M. 1973. Ecology: an Evolutionary Approach. Addison Wesley, Reading.

Grossfield, J. 1971. Geographic distribution and light dependent behavior in Drosophila. Proc.

Nat. Acad. Sci. (US) 68:.2669-2673.

Manning, A. 1965. Drosophila and the evolution of behavior. Viewpoints in Biology 4: 125—

169.

Prakash, S. 1973. Low gene variation in Drosophila busckii. Genetics 75: 571-576.

Rockwell, R. F., and M. B. Seiger. 1973. Phototaxis in Drosophila: a critical evaluation.

Amer. Sci. 61: 339-345.

Rockwell, R. F., F. Cooke, and R. Harmsen. 1975. Photobehavioral differentiation in natural

populations of D. pseudoobscura and D. persimilis. Behav. Genet. 5: 189-202.

Rockwell, R. F., and F. Cooke. 1977. Gene flow and local adaptation in a colonially nesting

dimorphic bird: the Lesser Snow Goose (Anser caerulescens caerulescens). Amer. Nat.

111: 91-97.

Rockwell, R. F., J. Grossfield, and L. Levine. 1978. Emigration response behavior: I. Effects

of height and light on the Oregon-R and norp-A strains of Drosophila melanogaster.

Egyptian Jour. Genet. Cytol. 7: 123-136.

Sakai, K., T. Narise, Y. Hiraizomi, and S. Iyama. 1958. Studies on competition in plants and

animals. IX. Experimental studies on migration in Drosophila melanogaster. Evolution

12: 93-101.

Throckmorton, L. 1975. The phylogeny, ecology and geography of Drosophila. In Handbook

of Genetics (Vol. 3), ed. R. C. King. Plenum Press, New York.

Winer, B. J. 1971. Statistical Principles in Experimental Design. McGraw-Hill, New York.

126 PAN-PACIFIC ENTOMOLOGIST

Footnote

1 Contribution 2 from the Theoretical Biology Study Group at City College of New York.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, p. 126

SCIENTIFIC NOTE

SYNONYMY OF THE GENUS EUPLUSIA MOURE

UNDER EUFRIESIA COCKERELL

(HYMENOPTERA, APIDAE, EUGLOSSINI)

The genus Eufriesia was originally based on the single species, pulchra.

Subsequently, another species, /ucifera Kimsey, was added. It was distin-

guished from the genus Euplusia by the broad flat scutellum, entire head

and T-IJI-VI or VIII brightly metallic with erect yellow setae, and the rest

of the body black with black setae.

On comparison of male genitalia of Eufriesia pulchra (F. Smith) and lu-

cifera Kimsey with the male genitalia of Euplusia species, I find no differ-

ences to support the separation of these two genera. Unlike other Euglos-

sini, both of these ‘‘genera’’ have strongly bilobed gonostyli and trilobed

gonocoxae.

Many external characteristics are also remarkably similar. Examination

of the entire genus Euplusia reveals a number of species with the same

color pattern as Eufriesia, including formosa (Mocsary) and theresiae (Moc-

sary) and other species with a broad flat scutellum, including violacea (Blan-

chard) and chalybaea (Friese). The genera share the following external char-

acteristics, which distinguish them from all other euglossines: male

hindtibial slit reaching apex; male midtibia with two adjacent felty patches;

face brightly metallic without white maculations, and female with a corbic-

ula.

The differences between these two groups are of species value only. Eu-

friesia pulchra and lucifera actually appear to belong to a species group

containing four species of Euplusia.

The genus Eufriesia described by Cockerell (1908) has priority over Eu-

plusia Moure (1943) which was a replacement for the preoccupied Plusia

Hoffmannsegg (1817).

Lynn Siri Kimsey, Dept. of Entomol., Univ. of Calif., Davis, 95616.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 127-146

SOME LARVAE OF ORTHOCLADIINAE, CHIRONOMIDAE

FROM BROOKS RANGE, ALASKA

WITH PROVISIONAL KEY

(DIPTERA)

LARRY J. TILLEY

Water Resources Division, U.S. Geological Survey,

Menlo Park, CA, 94025

Reconnaissance samples of benthic invertebrates from two arctic-alpine

streams, the Dietrich and Atigun Rivers, Alaska were dominated by Chi-

ronomidae larvae (Slack and others, 1976, 1977, 1979). In both rivers

the headwaters were dominated by the chironomid subfamily Diamesinae

whereas Orthocladiinae predominated further downstream. Although chi-

ronomids are known for their abundance in arctic freshwaters (Downes,

1962, 1964; Hobbie, 1973), little taxonomic information is available for Alas-

kan species. The present report describes and provides a key for the larvae

of eleven taxa in the subfamily Orthocladiinae. A similar report on the

Diamesinae and a single Podonominae from the same area is in preparation.

The Atigun River flows northward and the Dietrich River flows south-

ward, from the Continental Divide in the Brooks Range. The trans-Alaska

pipeline corridor traverses both drainage basins (Fig. 1), but the collections

on which this study is based were made in August 1971 before the start of

pipeline and road construction.

Methodology

Samples were preserved in 40 percent isopropyl alcohol when collected,

and were later separated in the laboratory from detritus by sugar flotation

(Anderson, 1959). The introductory keys for chironomid larvae prepared by

Mason (1973) and Beck (1968) were most useful because they indicated the

morphological characters of greatest value in the separation of species. Oth-

er helpful keys were those of Johannsen (1937), Chernovskii (1949), Roback

(1957) and Pankratova (1970, in Russian).

The chironomid larvae were first sorted into visually distinct groups. A

sample from each group was prepared for microscopic examination by

bleaching in hot 10 percent KOH (potassium hydroxide) solution to dissolve

soft body tissues. Each specimen was then placed ventral side up on a glass

slide in CMC-10! mounting medium and pressed under a 12 mm diameter

coverslip (Greeson and others, 1977). The illustrations for each taxon are

tracings from Polaroid photomicrographs. The heavy backing of the Polaroid

paper was carefully peeled from the prints and the insect parts traced using

128 PAN-PACIFIC ENTOMOLOGIST

ALASKA , —

ATIGUN RIVER

NGE DIETRICH RIVER

xs RM

pRoo ‘

} Location of study area

Trans-Alaska

Pipeline Corridor

170° 166° 162° 158° 164° 150° 146° 142° 138° 134°

io) 100 200 KILOMETERS

Litt =

Fig. 1. Location of the Dietrich and Atigun River basins in Alaska.

a light table. Larval measurements were made to the nearest micrometer

with a calibrated Whipple disc grid in the ocular of a light compound mi-

croscope.

Observations and measurements of the following larval characteristics

were used to separate taxa: 1. Antenna: length of each segment, ratio of

length of first segment to its width (“‘ALAW’’), length of first segment to

that of remaining four segments (‘‘AR’’). 2. Labial plate: relative size,

shape, and length of midtooth or midteeth; bifurcation of midtooth or num-

ber of midteeth; comparison of the width, or length of first pair of lateral

teeth to midtooth or midteeth, and total number of pairs of lateral teeth. 3.

Mandibles: number of teeth and their relative size distribution. 4. Preman-

dibles: number of digits, their relative size and appearance. 5. Preanal pa-

pillae: presence or absence, length versus width. 6. Preanal papillar bristles:

length, number, and location.

Instars were estimated using sizes of various morphological features, in-

cluding body length, head capsule length, and width and length of first an-

tennal segments. To illustrate variability of the averages, standard devia-

VOLUME 55, NUMBER 2 129

STRIATE

RETRACTILE PARALABIAL PLATE

ANTENNAE

MANDIBLE BLADES

B SOum ANNULATE 3rd

ANTENNAL

SEGMEN

LINGUA 1st ANTENNAL

EYESPOT SEGMENT

=tANNULAR

t ORGAN

g PREMANDIBLE

MANDIBLE OQum- 5

men DIAMESA ANTENNAE

fey, ANTENNAE

N LABIAL PLATE

BEARDED

PARALABIAL PLATE

Fig. 2. Head capsule structures (ventral view) used in the identification of larval Chiron-

omidae. (A) Tanypodinae, (B) Chironominae, (C) combined Orthocladiinae and Diamesinae,

A, B and C not drawn to scale, (D) antenna of Diamesa latitarsis (var. 1) to scale.

tions are reported using the symbol ‘‘S.D.’’ Specimens with conspicuously

swollen thorax areas were considered to be fourth (last) instars.

It was not possible to assign specific names to many of these larvae, nor

is it known whether or not a particular taxon has been described. Species

descriptions are based on adults and the immature stages may not be known.

The objective of the field study was to enumerate the taxa and their relative

abundances in samples of the benthic fauna. Hopefully the information in

this paper, which made it possible to distinguish taxa, will be of use to the

taxonomist interested in naming adult chironomids and to the ecologist

studying benthic invertebrates. Specimens are stored at the U.S. Geological

Survey, Western Region Headquarters in Menlo Park, California.

Key to the Common Subfamilies of Chironomidae Larvae and to the Genera

of Some Orthocladiinae From Reconnaissance Samples, Dietrich and Atigun

Rivers, Brooks Range, Alaska, August 1971

1. Head capsule with fork-shaped lingua; antennae long, often 4

length of head or longer, retractile (Fig. 2A)......... Tanypodinae

Head capsule without fork-shaped lingua, labial plate present (Fig.

2B and 2C); antennae not vetracttlO... ..sccasee ee wa pein oe des 2

—

PAN-PACIFIC ENTOMOLOGIST

. Premandible absent; preanal papillae at least 3 times longer than

sl POE ae ee, A ye ee UREN ee Eee Podonominae

Premandibles present (Big. 2B and 2€) as sscecs » even ok Shiels ee oe 5

. Paralabial plates present, usually large, conspicuous and striated

RISES EN) ta aroe, cee Oe he he oa Sushi sathmatedia's wale tas BA oe Chironominae

Paralabial plates usually absent, if present paralabial plates without

striations, although sometimes bearded (Fig. 2C) ............. 4

. Third segment of antenna annulate (ringed) (Fig. 2C and 2D); pre-

mandibles usually with more than three digits ........ Diamesinae

Third segment of antenna not annulate; premandibles usually with

one or two and sometimes three digits ....................00. 5

. Generally freshwater, occasionally terrestrial ...................

ee oreo ne Orthocladiinae (exclusive of Clunionini) 6

Seine et ESCs ENS LD <a Oe eek eee Eee ee Oe ogee eee ey Ok ee eee

Ele fs Telmatogetoninae and the Orthocladiinae tribe Clunionini

. Antennae at least one-half as long as head; body less than 5 mm

long; antennae four segmented (Fig. 13A) ..............2.008-

A= Corynoneura (Winnertz) Edwards Alaska sp. I (Fig. 13)

Antennae less than one-half as long as head; body longer than 5

TUOUEY ws seen Sepa tees Sarees a eect aahay eel a ls ate ettecd sega SP ue Ree ange BE a 7

. Labial plate with an even number of teeth; midteeth may appear

TOURS LEC oo Sk ee rary ea Se 5 et el ie te ic ae ee mS Foal as at. 8

Labial plate with an odd number of teeth; midtooth rarely trun-

CEE Solty ipa “a ee tare SPE op EER BOERNE cs ha wiih lol's Re eR OME ary ah. a a PUES 12

. Premandibles with more than one lobe; usually two or three (Figs.

eT Pe 5 al ca Seen csd eck eae Chaetocladius (Kieffer) 9

Premandibles with a single broad, apical lobe (Fig. 3D)..........

ee a Me ra ye Roy ie, eens Eukiefferiella Thienemann 10

. Midteeth of labial plate longer than first pair of lateral teeth (Fig.

TO lee he le tees OR Chaetocladius Alaska sp. I (Fig. 10)

Midteeth of labial plate short, about one-half as long as first pair of

lateral teeth (Fig. 11B)...... Chaetocladius Alaska sp. II (Fig. 11)

. Midteeth of labial plate rounded apically (Fig. 3B); mandibles usu-

ally with one or three long serrations on basal inner margin (Fig.

RT ok a bingy Se «RR a aera Eukiefferiella Alaska sp. I (Fig. 3)

Larva with characters not dS-above .....<06<+ dena dee ese beg ed 11

. Midteeth of labial plate not truncate; preanal papillae present, about

as long as wide; preanal papillary bristles long, about 600 um

long, in fourth instar (Fig. 5B and E) ........................

ee Eukiefferiella bavarica Goetghebuer (Fig. 5)

Midteeth of labial plate usually truncate; preanal papillae absent or

VOLUME 55, NUMBER 2 131

nearly so; preanal papillary bristles short and weak, about 100

ym long in fourth instar (Fig. 4B and E) .....................

fe ee ee ee Eukiefferiella cynaea Thienemann (Fig. 4)

12. Paralabial plates present; premandibles with three lobes (Fig. 12B

72404 Ol Ot Eee ee Parakiefferiella Thienemann, Alaska sp. I (Fig. 12)

Paralabial plates absent; premandible with less than three lobes ..

TEN Re ak = Ee ot re Genus Orthocladius Kieffer 13

13. Midtooth of labial plate about two times as wide as first lateral

tooth; premandible with cleft near apical end; location of Lau-

terborn organs uncertain; labial plate with six pairs of lateral teeth

Ss een ee ee ate inch ekg NaN ox EY Orthocladius s. str. (Fig. 9)

Labial plate with more than six pairs of lateral teeth; or antennae

with sessile Lauterborn organs at third antennal segment (Fig.

BO a, rs ae ade ata WL tse sg dans aE acide taste oi eer a 14

14. Labial plate with six pairs of lateral teeth; sessile Lauterborn

organs at third antennal segment (Fig. 6A) ...............000.

. Orthocladius (Euorthocladius) Thienemann Alaska sp. I (Fig. 6)

Labial plate with more than six pairs of lateral teeth ............ 15

15. Midtooth of labial plate narrow, about as wide as first lateral tooth;

margin of labial plate concave (Fig. 7B); mandibles with all teeth

a4 Ment ap CPUNDE La cect sRel@ ck aegees Be alee RAS eee Aga COR ee cae a mare

.. Orthocladius (Euorthocladius) Thienemann Alaska sp. II (Fig. 7)

Midtooth of labial plate broad, about three times as wide as first

lateral tooth, labial plate convex (Fig. 8B); apical tooth of man-

dible much larger than other mandibular teeth (Fig. 8C) .......

.. Orthocladius (Euorthocladius) Thienemann Alaska sp. III (Fig. 8)

Orthocladiinae

Eukiefferiella Thienemann

Larva, Goetghebuer 1932, in Pankratova 1970.

Eukiefferiella Alaska sp. I

(Fig. 3)

Three instars determined; body length of largest instar (fourth) 2.8—6.0

mm (average 4.1 mm,n = 4,S8.D. = 1.41 mm); of intermediate instar (third)

1.9-4.0 mm (average 2.78 mm, n = 49, S.D. = 0.54 mm); and of smallest

instar (second) 1.6—3.6 mm (average 1.9 mm, n = 22, S.D. = 0.41 mm).

Head capsule of largest instar average 0.33 mm long and 0.21 mm wide (n =

12, S.D. = 0.06 and 0.05 mm). Body color of preserved specimens yellow

during first few weeks of storage, after storage with differing amounts of

leaf and other detritus, many were brown. Head capsules brown, darker

brown with longer storage.

Length of antennal segments of largest instar (fourth) (Fig. 3A), 50: 14:

132 PAN-PACIFIC ENTOMOLOGIST

APICAL

B 1ooum}- e

Oum

Fig. 3. Eukiefferiella Alaska sp. I. (A) antennae, (B) labial plate, (C) left mandible, (D)

right premandible, (E) preanal papillae and papillar bristles.

5: 5: 4 um (n = 4, S.D. = 7.3: 1.7: 0.96: 0.96: 0.82 «m); width of first

segment 19 um (n= 5, S.D. = 2.39 um); AR = 1.86, S.D. = 0.30,

ALAW = 2.64, S.D. = 0.29. Length of antennal segments of intermediate

instar (third) 24: 11: 3: 5: 4 wm (n = 60: 57: 57: 57: 57, S.D. = 3.65: 1.23:

1.22: 0.75: 0.71 um); width of first antennal segment 11 wm (n = 60, S.D. =

1.95 um); AR = 1.11, S.D. = 0.12, ALAW = 2.29, S.D. = 0.29). Length

of antennal segments of smallest instar (second) 13: 9: 2: 3: 3 wm (n = 22,

S.D. = 2.40: 1.22: 0.79: 0.74: 0.79 wm); width of first antennal segment 8

wm (n = 22, S.D. = 1.05 um); AR = 0.76, S.D. = 0.17; ALAW = 1.69,

S.D. = 0.37.

Labial plate (Fig. 3B) with midteeth divided, each midtooth rounded api-

cally, much wider than laterals (division not distinct on worn specimens).

Five pairs of pointed lateral teeth.

Mandibles (Fig. 3C) with five teeth, progressively smaller from apical

tooth to basal tooth. Basal tooth actually a dark area on the apical part of

rounded basal section of mandible. Premandible (Fig. 3D) with apical end

stout and broad, basal part or attached end convoluted.

Preanal papillae short, when present wider than long. Usually six bristles

can be seen at apex of papillae (Fig. 3E); two bristles are shorter than the

other four. The four longer bristles on the largest instar (fourth) about 300

wm long (n = 5); intermediate instar (third) 189 wm (n = 65, S.D. = 72.4

yzm); and for the smallest instar (second) 136 wm (n = 11, S.D. = 37.6 wm).

Microscope slides were prepared for 149 individual specimens. Detailed

measurements were made on 85 specimens. A total of 811 specimens of

Eukiefferiella Alaska sp. I were estimated from 36 samples collected at 12

sampling sites.

VOLUME 55, NUMBER 2 133

APICAL

120um - oo

is \

60um-

OumL B

HM S00umr

OumL

Fig. 4. Eukiefferiella cynaea, Thienemann. (A) antenna, (B) labial plate, (C) left mandible,

(D) right premandible, (E) preanal papillar bristles.

Eukiefferiella cynaea Thienemann

(Fig. 4)

Larva, Thienemann 1936, in Thienemann 1954.

Three instars determined; body length of largest instar (fourth) 3.8—5.7

mm (average 4.73 mm, n= 6, S.D. = 0.66 mm); of intermediate instar

(third) 2.0-4.0 mm (average 3.10 mm, n = 45, S.D. = 0.51 mm); and of

smallest instar (second) 1.4—2.5 mm (average 1.93 mm, n = 11, S.D. = 0.41

mm). Head capsule of largest instar average 0.35 mm long and 0.25 mm

wide (n = 6, S.D. = 0.028 mm and 0.055 mm); of intermediate instar, 0.22

mm long and 0.16 mm wide (n = 41, S.D. = 0.035 and 0.032 mm); and of

smallest instar 0.18 mm long and 0.13 mm wide (n = 11, S.D. = 0.041 mm

and 0.093 mm). Body color yellow to dark yellow or gray, some with banded

appearance (this occurred on many specimens of several taxa where that

part of an overlapping abdominal sclerite was darker than the remainder of

the sclerite). Usually, specimens were darker dorsally. Head capsules light

brown to brown.

Antennae of largest instar (fourth) (Fig. 4A), length of antennal segments

50: 11: 4: 5: 4 um (n = 7: 6: 6: 6: 6, S.D. = 3.36: 1.63: 1.94: 0.41: 1.10 wm);

width of first antennal segment 17 wm (n = 7, S8.D. = 1.25 um); AR = 2.01,

S.D. = 0.20; ALAW 2.97, S.D. = 0.21. Length of antennal segments of

134 PAN-PACIFIC ENTOMOLOGIST

intermediate instar (third) 24: 11: 3: 4: 4 wm (n = 45: 42: 42: 42: 42, S.D. =

2.08: 1.03: 1.29: 0.07: 0.63 um); width of first antennal segment 11 wm (n =

45, S.D. = 0.97 um); AR = 1.13, S.D. = 0.177; ALAW = 2.23, S.D. =

0.26). Length of antennal segments of smallest instar 16: 10: 3: 4: 3.5 wm

(n = 11, S.D. = 2.17: 1.13: 0.87: 1.0: 0.69 um); width of first antennal seg-

ment 9 um (n = 11, S.D. = 1.54 wm); AR = 0.82, S.D. = 0.136; ALAW =

1.83, S.D. = 0.213.

Labial plate (Fig. 4B) of practically all specimens had midtooth truncate

(probably there were two teeth before wear) (Fig. 4B), some specimens with

two teeth were otherwise very similar to those with truncate midtooth, so

are placed in a single taxon. Truncate area 4 to 5 times width of base of

lateral teeth. Five pairs of pointed lateral teeth present with first pair usually

showing apical wear.

Mandibles (Fig. 4C) with 5 teeth, progressively smaller from apical tooth

to proximal tooth; antennae about same length as mandibles. Premandibles

(Fig. 4D) with apical end stout and broad, basal part convoluted.

Preanal papillae absent or nearly so. Four weak bristles present at papillae

sites (Fig. 4E); 105 um long (n = 7, S.D. = 25.3 um) for largest instar

(fourth); 90 um long (n = 66, S.D. = 21.5 um) for intermediate instar

(third); and 85 wm (n = 11, S.D. = 22.8 ~m) for smallest instar (second).

Microscope slides were prepared for 117 individual specimens. Detailed

measurements were made on 84 specimens. A total of 581 specimens of E.

cynaea were estimated from 36 samples at 12 sampling sites.

Eukiefferiella bavarica Goetghebuer

(Fig. 5)

Larva, Thienemann 1935, in Pankratova 1970.

Three instars determined; body length of largest instar (fourth) 3.4—6.0

mm (average 4.69 mm, n = 24, S.D. = 0.68 mm; of intermediate instar

(third) 3.0-3.7 mm (average 3.38 mm, n = 12, S.D. = 0.22 mm); and of

smallest instar 1.6—2.5 mm (average 2.20, n = 3, S.D. = 0.52 mm). Head

capsule of largest instar average 0.36 mm long and 0.25 mm wide (n = 24,

S.D. = 0.03 mm and 0.023 mm); of intermediate instar 0.30 mm long and

0.23 mm wide (n = 12, S.D. = 0.04 mm and 0.035 mm); and of smallest

instar 0.18 mm long and 0.15 mm wide (n = 3, S.D. = 0.052 mm and 0.05

mm). Body color of preserved specimens light yellow during the first few

weeks of storage, after storage with leaf and other detritus, light brown.

Head capsules brown to dark brown.

Antennae (Fig. 5A) with first segment long compared to other Eukieffer-

iella taxa from these samples. Length of antennal segments of largest instar

(fourth) 66: 14: 4: 6: 5 um (n = 15, S.D. = 5.73: 1.79: 0.96: 0.86: 0.72 um);

width of first antennal segment 19 um (n = 15, S.D. = 2.24 wm); AR =

VOLUME 55, NUMBER 2 135

APICAL

120um -

aa im om

Oume

Fig. 5. Eukiefferiella bavarica Goetghebuer. (A) antenna, (B) labial plate, (C) left mandible,

(D) left premandible, (E) preanal papillae and papillar bristles.

2.28, S.D. = 0.22; ALAW = 3.52, S.D. = 0.42. Length of antennal seg-

ments of intermediate instar (third) 44: 16: 4: 6: 4 wm (n = 12, S.D. = 6.06:

1.24: 1.16: 0.57: 0.72 um); width of first antennal segment 16 wm (n = 12,

S.D. = 0.75 um); AR = 2.89, S.D. = 0.38; ALAW = 2.64, S.D. = 0.75.

Length of antennal segments of smallest instar (third) 21: 12: 4: 4: 3 um (n =

3, 8.D. = 5.13: 1.53: 1.15: 1.0: 0.58 wm); width of first antennal segment 11

wm (n= 3, S.D. = 1.53 wm); AR = 0.88, S.D. = 0.26; ALAW = 1.92,

SoD, = O222)5

Labial plate (Fig. 5B) with two large midteeth, 2 to 3 times larger than

lateral teeth. Five pairs of lateral teeth.

Mandibles (Fig. 5C) with five teeth progressively smaller from apical tooth

to proximal tooth. Premandibles (Fig. SD) apically stout and broad, con-

voluted at the base where attached.

Preanal papillae present (Fig. SE) about as wide as long approximately 35

wm long and 37 wm wide. Six bristles attached to end of papillae (in largest

instar, fourth) 600 um long (n = 10, S.D. = 125.0 ym), one bristle attached

to side of papillae 150 wm long (n = 8, S.D. = 37.8 wm). Apical bristles for

intermediate instar (third) 330 wm long (n = 8, S.D. = 61.1 wm); and for

smallest instar (second) apical bristles 230 wm long (n = 1).

136 PAN-PACIFIC ENTOMOLOGIST

APICAL

120um

60um

Fig. 6. Orthocladius (Euorthocladius) Alaska sp. I. (A) antenna, (B) labial plate, (C) left

mandible, (D) right premandible, (E) preanal papillar bristles.

Microscope slides were prepared for 50 individual specimens. Detailed

measurements were made on 39 specimens. A total of 139 specimens of E.

bavarica were estimated from 36 samples collected at 12 sampling sites.

Genus Orthocladius Kieffer

Subgenus Euorthocladius Thienemann

Larva, as per O. A. Saether, written communication, 1973

Orthocladius (Euorthocladius) Alaska sp. I

(Fig. 6)

Three instars determined; body length of largest instar (fourth) 3.5—5.5

mm (average 4.5 mm,n = 4,8.D. = 0.90 mm); of intermediate instar (third)

2.3-3.0 mm (average 2.65 mm, n = 4, S.D. = 0.31 mm); and of smallest

instar 1.5—2.8 mm (average 2.30 mm, n = 27, S.D. = 0.44 mm). Head cap-

sule of largest instar average 0.26 mm long and 0.14 mm wide (n = 2); of

intermediate instar, 0.24 mm long and 0.19 mm wide (n = 4, S.D. = 0.042

mm and 0.013 mm); of smallest instar (second) 0.21 mm long and 0.16 mm

wide (n = 26, S.D. = 0.043 mm and 0.026 mm). Body color of preserved

specimens light green or light yellow, later brown after storage with leaf and

other detritus. Head capsules light brown.

Length of antennal segments of largest instar (fourth) (Fig. 6A), 34: 9: 6:

VOLUME 55, NUMBER 2 137

3: 4 um (n = 4: 3: 3: 3: 3, S.D. = 4.35: 1.15: 2.0: 0.58: 0.58 wm); width of

first antennal segment 18 um (n = 4, S.D. = 4.24 wm). Length of antennal

segments of intermediate instar (third) 23: 11: 3: 4: 4 um (n = 4, S.D. =

1.71: 1.89: 0.96: 0.96: 0.58 «m); width of first antennal segment 12 wm (n =

4,S.D. = 1.41 wm). Length of antennal segments of smallest instar (second)

17: 9: 3: 3: 3 wm (n = 24: 24: 23: 23: 23, S.D. = 2.95: 0.66: 1.58: 0.54: 0.39

ym); width of first antennal segment 12 wm (n = 23,8.D. = 1.57 um); AR =

0.93, S.D. = 0.175; ALAW = 1.41, S.D. = 0.18. Usually sessile Lauter-

born organs are present, attached to apical end of second antennal segment

and are as long as third antennal segment (Fig. 6A).

Labial plate (Fig. 6B) with midtooth ranging from about one and one-half

to twice as wide as the first pair of lateral teeth. The midtooth and first

laterals show wear before remainder of laterals, usually are set apart slightly

and all three truncate. There are six pairs of lateral teeth.

Mandibles (Fig. 6C) with five teeth, progressively smaller from apical

teeth to proximal teeth. Second tooth wider than first tooth, sometimes

mandible teeth appear as two large teeth and three small teeth. Premandibles

(Fig. 6D) apically with a single tapered lobe.

Preanal papillae absent, or nearly so. Six bristles present at papillae sites

(Fig. 6E); 400 wm long (n = 2) on largest instar (fourth); 360 4m long (n =

4, S.D. = 80.1 wm) on intermediate instar (third) and rounded to 315 wm

long (n = 24, S.D. = 67.4 «m) on smallest instar (second).

Microscope slides were prepared for 71 individual specimens. Detailed

measurements were made on 34 specimens. A total of 785 O. (Euortho-

cladius) Alaska sp. I were estimated from 36 samples collected at 12 sam-

pling sites.

Orthocladius (Euorthocladius) Alaska sp. I

(Fig. 7)

Two instars determined; body length of largest instar (fourth) 3.0-6.4 mm

(average 4.15 mm, n = 11, S.D. = 1.19 mm); of smaller instar 3.5 mm long

(n = 1). Head capsule of largest instar 0.33 mm long and 0.29 mm wide (n =

2). Body color of preserved specimens dark yellow-brown or brown, some

with banded appearance.

Antennae (Fig. 7A) with first segment short and wide compared with other

specimens from these samples. Length of antennal segments of largest instar

(fourth) 28: 11: 3: 3: 3 wm (n = 12: 12: 11: 11: 11, S.D. = 2.61: 1.44: 1.19:

0.54: 0.65 4m); width of first antennal segment 21 wm (n = 11, 8.D. = 2.91

wm); AR = 1.39, S.D. = 0.24; ALAW = 1.37, S.D. = 0.14.

Labial plate (Fig. 7B) with midtooth only slightly wider than lateral teeth.

Margin of labial plate on these specimens concave. Nine pairs of lateral

teeth present.

138 PAN-PACIFIC ENTOMOLOGIST

ARIGAL

[FeO

Te ers

Oum

Fig. 7. Orthocladius (Euorthocladius) Alaska sp. II. (A) antenna, (B) labial plate, (C) left

mandible, (D) right premandible, (E) preanal papillar bristles.

Mandibles (Fig. 7C), most showing wear, with teeth progressively smaller

(about same size on worn specimens) from apical teeth to proximal teeth.

Premandibles (Fig. 7D) slightly divided apically.

Preanal papillae absent; 4 pairs of weak bristles at papillae sites (Fig. 7E)

average 90 um (n = 7, S.D. = 17.3 um) on largest instar (fourth); 50 wm

(n = 1) on smaller instar (third).

Microscope slides were prepared for 26 individual specimens. Detailed

measurements were made on 13 specimens. A total of 58 O. Euorthocladius

Alaska sp. II were estimated for 36 samples at 12 sampling sites.

Orthocladius (Euorthocladius) Alaska sp. II

(Fig. 8)

Only seven individuals were estimated from 36 samples collected at 12

sampling sites. Microscope slides were prepared and detailed measurements

made on two specimens.

Both measured specimens were believed to be the same instar. Body

lengths 3.5 and 4.6 mm. Head capsule of larger specimen 0.45 mm long and

0.32 mm wide. Body color of preserved specimens orange-brown and head

capsule brown.

Antennae (Fig. 8A) for larger specimen with first segment was very wide

and stout compared to specimens from other taxa in these samples. Length

of antennal segments for the larger specimen 49: 7: 3: 2: 4 um, width of first

antennal segment 37 um; AR = 3.06; ALAW = 1.32. Width of first antennal

segment about three to four times wider than width of second segment.

Labial plate (Fig. 8B) with midtooth three to four times wider than lateral

VOLUME 55, NUMBER 2 139

150um -

75um/ A

Fig. 8. Orthocladius (Euorthocladius) Alaska sp. III. (A) antenna, (B) labial plate, (C) left

mandible, (D) right premandible, (E) preanal papillar bristles.

teeth. Nine pairs of lateral teeth. Margin of labial plate smooth, slightly

worn with tips of lateral teeth pointed toward the midtooth.

Mandibles (Fig. 8C) with a very large, scythe-shaped apical tooth and

four, much smaller, proximal teeth. Premandibles (Fig. D) a large tapered

lobe apically.

Preanal papillae absent; 4 or 5 bristles (Fig. 8E) arising from each papillae

site about 500 um long.

Subgenus Orthocladius s.str.

(Fig. 9)

Information provided by Ole A. Saether, 1973

Two instars determined; body length of largest instar (fourth) 2.6—6.0 mm

(average 4.67 mm, n = 31, S.D. = 0.77 mm); for smaller instar (third) 1.9-

3.0 mm (average 2.46 mm, n = 9, S.D. = 0.37 mm). Head capsule of largest

instar 0.41 mm long and 0.31 mm wide (n = 30, S.D. = 0.067 mm and 0.048

mm); and of smaller instar 0.24 mm long and 0.18 mm wide (n = 8, S.D. =

0.025 mm and 0.016 mm). Body color of preserved specimens yellow, yel-

low-brown, some with green tinge to other colors. Head capsules yellow,

light brown or yellow-brown.

140 PAN-PACIFIC ENTOMOLOGIST

APICAL

120um

Pd OR

60um+-

200um- j

100um-

Oum+

Fig. 9. Orthocladius s.str. (A) antenna, (B) labial plate, (C) left mandible, (D) right pre-

mandible, (E) preanal papillar bristles.

Antennae of largest instar (Fig. 9A) with segment lengths of 49: 14: 6: 5:

4 um (n = 29: 28: 28: 28: 28, S.D. = 4.18: 2.25: 1.33: 0.99: 0.90 um); width

of first antennal segment 22 um (n = 27, S.D. = 3.95 wm); AR = 1.77,

S.D. = 0.258; ALAW = 2.30, S.D. = 0.387. Length of antennal segments

of smaller instar 20: 9: 4: 3: 3 wm (n = 9, S.D. = 4.5: 1.50: 0.53: 0.33: 0.73

yum); width of first antennal segment 12 wm (n = 9, S.D. = 0.93 wm); AR =

1.07, S.D. = 0.103; ALAW = 1.73, S.D. = 0.438.

Labial plate (Fig. 9B) with a wide, rounded midtooth, two to two and

one-half times as wide as lateral teeth. There are 6 pairs of lateral teeth,

usually the first pair is slightly worn or rounded.

Mandibles (Fig. 9C) with 5 teeth progressively smaller from apical teeth

to proximal teeth. Crenulations on the outer margin of the mandibles are

prominent on some specimens, weak or absent on others. Premandibles

(Fig. 9D), a tapered lobe with a small notch near apex.

Preanal papillae slight (Fig. 9E), always wider than long. Six bristles pres-

ent at apex of each papilla; 485 wm long (n = 29, S.D. = 123.1 wm) on the

largest instar (fourth); 320 wm long (n = 9, S.D. = 14.9 wm) on the smaller

instar (third).

Microscope slides were prepared for 41 individual specimens. Detailed

VOLUME 55, NUMBER 2 141

120um : jute

)

60um-

Oum- B

200um -

1 O0Oum ie

Oum*

Fig. 10. Chaetocladius Alaska sp. I. (A) antenna, (B) labial plate, (C) left mandible, (D)

right premandible, (E) preanal papillae and papillar bristles.

measurements were made on 39 specimens. A total of 103 specimens of

Orthocladius s.str. were estimated from 36 samples collected at 12 sampling

sites.

Genus Chaetocladius (Kieffer)

Larva, Thienemann 1921, in Pankratova 1970

Chaetocladius Alaska sp. I

(Fig. 10)

One instar determined (third or fourth) body length 3.0—5.4 mm (average

3.97 mm, n = 11, S.D. = 0.70 mm). Head capsule, 0.28 mm long and 0.20

mm wide (n = 11, S.D. = 0.057 mm and 0.041 mm). Body color of pre-

served specimens yellow-brown, and head capsule brown.

Antennae (Fig. 10A), each segment 36: 10: 5: 4: 3 wm long (n = 13: 13:

11: 11: 11, $.D. = 5.72: 1.55: 1.49: 1.21: 0.92 wm); width of first antennal

segment 16 um (n = 13, S.D. = 2.91 um); AR = 1.56, S.D. = 0.32;

ALAW = 2.23, S.D. = 0.45.

Labial plate (Fig. 10B) with 2 large, dark midteeth and S pairs of lateral

teeth. Last pair of teeth nearly trilobed single tooth.

142 PAN-PACIFIC ENTOMOLOGIST

APICAL

120um

ian

60um

oun 200.m

100um

Oum

Fig. 11. Chaetocladius Alaska sp. II. (A) antenna, (B) labial plate, (C) left mandible, (D) left

premandible, (E) preanal papillae and papillar bristles.

Mandibles (Fig. 10C) with 5 teeth, apical tooth large, second tooth one-

half size of apical tooth and last three teeth much smaller. Premandible (Fig.

10D) with three lobe-like digits.

Preanal papillae (Fig. 10E) present, about as long as wide (20 um long

and 20 um wide). Five bristles at apices of papillae, 325 wm long (n = 9,

S.D. = 56.6).

Microscope slides were prepared for 15 individual specimens. Detailed

measurements were made on 13 specimens. A total of 17 specimens of

Chaetocladius Alaska sp. I were estimated from 36 samples collected at 12

sampling sites.

Chaetocladius Alaska sp. II

(Fig. 11)

Only a single specimen was taken from 36 samples collected at 12 sam-

pling sites.

Body length of specimen 3.6 mm, head capsule not measured. Body color

of preserved specimen banded, brown bands and brown-white between

bands, head capsule dark brown.

Antennae (Fig. 11A) length of segments 45: 13: 3: 3: 3 um; width of first

antennal segment 25 wm; AR = 2.05, ALAW = 1.8.

Labial plate as in Fig. 11B. Two midteeth each only one-half as long as

VOLUME 55, NUMBER 2 143

150um/- APICAL

[EO >

75um_ ° A

Oum

Fig. 12. Parakiefferiella Alaska sp. I. (A) antenna, (B) labial plate with paralabials, (C)

left mandible, (D) right premandible, (E) preanal papillae and papillar bristles.

first or second pairs of lateral teeth. Five pairs of lateral teeth, last three

pairs smaller than first two pairs.

Mandibles (Fig. 11C) with five teeth, apical tooth large and remaining four

teeth small and progressively smaller from second tooth to proximal teeth.

Premandibles (Fig. 11D) with two large tapered lobe-like digits with a lateral

(mesad) bulge.

Preanal papillae (Fig. 11E) present, about as long as wide (32 um long

and 35 wm wide) with a robust appearance. Six pairs of bristles about 300

wm long.

Genus Parakiefferiella (Thienemann)

Larva, Thienemann 1936, as in Pankratova 1970

Parakiefferiella Alaska sp. I

(Fig. 12)

Only 11 individuals were estimated from 36 samples collected at 12 sam-

144 PAN-PACIFIC ENTOMOLOGIST

pling sites. Microscope slides were prepared for five specimens and detailed

measurements made on two specimens.

Body length of five specimens 3.5—4.5 mm (average 4.0mm, n = 5,S.D. =

0.5 mm). Head capsule 0.38 mm long and 0.28 mm wide (n = 2). Body color

of preserved specimens yellow, head capsule yellow-brown.

Antennae as in Fig. 12A. Length of antennal segments 50: 16: 5.5: 3: 2.5

ym (n = 2); width of first antennal segment 29 wm (n = 2); AR = 1.85;

ALAW = 1.72.

Labial plate (Fig. 12B) with a single midtooth one and one-half to two

times wider than lateral teeth. Six pairs of lateral teeth present with para-

labial plates beginning apically at outer base of second pair of lateral teeth

to posterior of the base of the last pair of lateral teeth (paralabial plates not

easily detected on these specimens).

Mandibles (Fig. 12C) with five teeth, progressively smaller from apical

tooth to proximal tooth. Premandibles (Fig. 12D) with two tapered lobes.

Preanal papillae (Fig. 12E) wider than long (about 20 wm long and 32 wm

wide). Four terminal bristles about 425 um long.

Genus Corynoneura (Winnertz) Edwards, in Pankratova 1970

Larva, Pankratova 1970

Corynoneura Alaska sp. I

(Fig. 13)

Only 6 specimens of C. Alaska sp. I were estimated from 36 samples

collected at 12 sampling sites, all specimens came from one site. One instar

determined (third or fourth), body length 1.2 mm long (n = 2). Head capsule

of a single specimen 0.14 mm long and 0.11 mm wide. Body color of pre-

served specimens yellow, head capsule light brown.

Antennae (Fig. 13A) with two annular organs, the first at midpart of first

antennal segment and the second about three-fourths the length of first an-

tennal segment from the base. There is a small spur-like blade about 4 wm

long at apex of second antennal segment. Antennae with four segments,

_ length of antennal segments 63: 47: 45: 2 um (n = 2); width of first segment

17 um; AR = 0.66; ALAW = 3.71.

Labial plate (Fig. 13B) with midtooth small, first pair of lateral teeth much

larger than midtooth. Total of 6 pairs of lateral teeth, second pair smaller

than third pair, and third pair almost as large as first pair.

Mandibles (Fig. 13C) with five teeth, first tooth smaller than second tooth.

Third through fifth tooth progressively smaller. Premandibles (not illustrat-

ed) poor on both specimens, possibly divided into two lobes.

Preanal papillae short (Fig. 13E), 4 bristles at apex about 160 um long.

Spines, one group each at dorsal base of each proleg. Each spine group

VOLUME 55, NUMBER 2 145

21 ny

105um- LO

q .

Oum*

Fig. 13. Corynoneura unnamed Alaska sp. I. (A) antenna, (B) labial plate, (C) left mandible,

(D) spines, of posterior prolegs, (E) preanal papillar bristles.

(Fig. 13D) with one long spine 48 u~m long and a second spine (at base of

first spine) 11 wm long.

Acknowledgment

Dr. Ole A. Saether of The Freshwater Institute in Winnipeg, Canada,

identified representative samples of each taxon. Most of the drawings were

prepared from specimens examined by him.?

Literature Cited

Anderson, R. O. 1959. A modified flotation technique for sorting bottom fauna samples.

Limnol. and Oceanog. 4: 223-225.

Beck, W. M., Jr. 1968. Chironomidae. In Keys to water quality indicative organisms (south-

eastern United States). Edited by F. K. Parrish, Fed. Water Pollut. Control Admin.,

vl—v22.

Chernovskii, A. A. 1949. Identification of larvae of the midge family Tendipedidae. Izdat.

Akad. Nauk S.S.S.R., Transl. by Lees, E., (1961) (ed. K. E. Marshall), Natl. Lending

Library for Sci. and Tech., Boston Spa, Yorkshire, England, 300 p.

Downes, J. A. 1962. What is an Arctic insect? Canadian Entomol. 94: 143-162.

. 1964. Arctic insects and their environment. Canadian Entomol. 96: 279-307.

Greeson, P. E., T. A. Ehlke, G. A. Irwin, B. W. Lium, and K. V. Slack, editors. 1977.

Methods for collection and analysis of aquatic biological and microbiological samples.

U.S. Geol. Survey Techniques Water-Resources Inv., book 5, chap. A4, 332 p.

146 PAN-PACIFIC ENTOMOLOGIST

Hobbie, J. E. 1973. Arctic limnology—A review. In Britton, M. E., ed., Alaskan arctic tundra.

Arctic Inst. North America Tech. Paper 25: 127-168.

Johannsen, O. A. 1937. Aquatic Diptera. Part III. Chironomidae: subfamilies Tanypodinae,

Diamesinae, and Orthocladiinae. Mem. Cornell Univ. Agr. Exp. Sta. 205: 1-84 [reprint-

ed 1970 by Entomological Repr. Specialists, Los Angeles].

Mason, W. T., Jr. 1973. An introduction to the identification of chironomid larvae. Cincinnati,

Ohio, Natl. Environmental Research Center. 1-90.

Pankratova, V. Y. 1970. [Larvae and pupae of midges of the subfamily Orthocladiinae of the

fauna of the USSR.] Leningrad ‘Nauka.’ 1-343. In Russian.

Roback, S. S. 1957. The immature Tendipedids of the Philadelphia area. Monog. Acad. Natl.

Sci., Philadelphia 9: 1-152.

Slack, K. V., J. W. Nauman, and L. J. Tilley. 1976. Evaluation of three collecting methods

for a reconnaissance of stream benthic invertebrates. U.S. Geol. Survey J. Research 4:

491-495.

Slack, K. V., J. W. Nauman, and L. J. Tilley. 1977. Benthic invertebrates in an arctic

mountain stream, Brooks Range, Alaska. U.S. Geol. Survey J. Research 4: 419-519.

Slack, K. V., J. W. Nauman, and L. J. Tilley. 1979. Benthic invertebrates in a north-flowing

stream and a south-flowing stream, Brooks Range, Alaska. Water Resources Bulletin,

15: 108-135.

Thienemann, A. 1935. Chironomiden-Metamorphosen. XII. Deutsche Ent. Zeitschr. 4: 86-96.

Thienemann, A. 1936. Alpine Chironomiden (Ergebnisse von Untersuchungen in der Gegend

von Garmisch-Partenkirchen), [Alpine Chironomidae Report on investigations in the

vicinity of Garmisch-Partenkirchen]. Arch. Hydrobiol. 30: 167-262.

Thienemann, A. 1954. Chironomus. Die Binnengewasser, Stuttgart 20: 1-834.

Footnote

1 Mention of trade names or commercial products does not constitute endorsement by the

U.S. Geological Survey nor recommendation for use.

2 Dr. Saether is now Professor of Systematic Zoology at the University of Bergen, Norway.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 147-148

A NEW SPECIES OF TANARTHRUS FROM CALIFORNIA

(COLEOPTERA: ANTHICIDAE)

DONALD S. CHANDLER

641 Cherry #5, Chico, CA, 95926

Recently a large series of an undescribed species of Tanarthrus was col-

lected along the Leslie salt pans in San Francisco Bay. The distribution of

the new species is of interest since the species most similar to it, T. iselini

Chandler, is found only in central New Mexico. Following the distributional

mechanisms proposed in my revision (Chandler, 1975), this distribution sug-

gests a pre-Pleistocene separation of populations.

The species description follows that of Chandler (1975). All measurements

are in millimeters. I would like to thank Christine A. Janus-Chandler for

checking the manuscript.

Tanarthrus occidentalis, new species

(Figs. 1-2)

General description.—Elytra usually piceous, fuscous areas may be pres-

ent to cover basal third and apical fourth; head and pronotum fulvous; elytra

feebly ridged and punctate, pubescence directed posteriorly, microreticu-

lation present; pronotal angles rounded, roughly sculptured by punctation,

microreticulate between punctures; head subtruncate, base with medial

depression, shallow punctures distinct, moderately dense, microreticulation

distinct, eleventh antennal segment distinctly constricted, portion before

constriction as long as to slightly shorter than tenth, portion after constric-

tion as long as to slightly longer than tenth; length 2.61—3.55.

Male holotype.—3.2 km (2 mi) W Newark, California. Head 0.83 long,

width behind eyes 0.79, length from base to eyes 0.35, width at antennal

bases 0.43, length from base to antennal bases 0.59; basal portion of eleventh

antennal segment 0.07 long, after constriction 0.10 long, tenth segment 0.10

long. Pronotum 0.76 long, width at base 0.49, widest point 0.68 at 0.50 from

base; collar 0.04 thick, 0.30 wide. Elytra 1.94 long, width at humeri 0.90,

scattered erect setae at 40—60 degree angle. Profemur 0.66 long, 0.19 wide,

protibia 0.67 long; mesofemur 0.65 long, 0.20 wide, mesotibia 0.65 long;

metafemur 0.79 long, 0.21 wide, metatibia 0.72 long.

Genitalia with tegmen as long as phallobase; internal sac with numerous

spines; primary gonopore with sclerotized ring.

Female without erect elytral setae.

Relationship.—most similar to T. iselini in the medial basal depression of

148 PAN-PACIFIC ENTOMOLOGIST

Figs. 1-2. Fig. 1. Tenth and eleventh antennal segments. Fig. 2. Ventral view of male

genitalia.

the head and the erect elytral setae of the male. Separated from iselini by

the entire antenna being more slender in appearance and both portions of

the eleventh antennal segment being longer than wide. In iselini the divi-

sions of the eleventh segment appear almost moniliform and the tenth seg-

ment and basal portion of the antennal constriction are as long as wide.

Distribution.—Known only from the Leslie salt pans near Newark along

San Francisco Bay. HOLOTYPE male, 3.2 km (2 mi) W Newark, off Dum-

barton Bridge, Alameda County, California, V-27-1976, C. Y. Kitayama. 33

PARATYPES: 15 males, 5 females, eutopotypical; 1 male, 12 females, same

locality, V-15-1978, D. S. Chandler, under debris along salt pans.

The holotype will be deposited in the California Academy of Sciences,

with paratypes to be placed in the United States National Museum and the

Floyd G. Werner collection, Tucson, Arizona.

Literature Cited

Chandler, Donald S. 1975. A revision of Tanarthrus LeConte with a presentation of its Mid-

Cenozoic speciation (Coleoptera: Anthicidae). Trans. American Entomol. Soc. 101: 319-

354.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 149-154

A NEW EREMOCORIS FROM CALIFORNIA WITH A KEY TO

NORTH AMERICAN GENERA OF DRYMINI

(HEMIPTERA-HETEROPTERA: LYGAEIDAE)

PETER D. ASHLOCK!

Dept. of Entomol., Univ. of Kansas, Lawrence, 66045

Peculiar species often present problems of placement, and such is the

case with a short-winged drymine (Rhyparochrominae) described herein.

Specimens of the species, which were collected in northern California on

Sargent’s Cypress, have been in my collection for several years, and addi-

tional specimens were found in the collection of the California Academy of

Sciences. The species looks like an Eremocoris, but is an unusual member

of that genus. It has shorter wings than any other species and is quite

uniformly dark reddish brown, lacking the patterns on the pronotum and

hemelytra common in the genus. The degree of brachyptery is similar to

that of Togodolentus wrighti (Van Duzee), and in both species the anterior

lobe of the pronotum is much longer than the hind lobe and somewhat

swollen. These pronotal characters are associated with the wing modifica-

tion. There were no macropterous specimens available of either species.

Study was begun to determine if the new cypress bug is an odd Eremo-

coris, or a second species of the monotypic genus Togodolentus, or whether

Togodolentus wrighti itself is merely another peculiar species of Eremo-

coris.

The hind tibia of the cypress bug has along its entire length numerous

long, erect hairs that are about three times as long as the measurement

across the tibia (see Fig. 1). Sweet (1977) described this condition for E.

ferus (Say) from eastern North America and used it to distinguish E. ferus

from E. borealis (Dallas). Other Eremocoris with long tibial hairs are E.

setosus Blatchley (eastern U.S., Canada), E. plebejus Fallen (Europe, Si-

beria), and E. semicinctus Van Duzee (California, Idaho). These long tibial

hairs may have confused E. P. Van Duzee (1921) when he described E.

semicinctus, for the allotype of his species is a female of the cypress bug

described below. Eremocoris species with short, appressed tibial hairs, in

addition to E. borealis cited by Sweet (1977), are E. depressus Barber

(southeastern U.S.), E. dimidiatus Van Duzee (Colorado), E. obscurus Van

Duzee (western U.S., Canada), and E. opacus Van Duzee and E. inquilinis

Van Duzee (both California). All other North American drymines, and To-

godolentus wrighti, have short, appressed hairs on the hind tibia.

The long tibial hairs found only in some species of Eremocoris may be

a synapomorphous character delimiting a holophyletic group within the ge-

150 PAN-PACIFIC ENTOMOLOGIST

nus. If so, placing the cypress bug in Togodolentus would make Togodo-

lentus polyphyletic, which is contrary to good systematic practice.

Whether Jogodolentus wrighti is enough like Eremocoris species to be

placed in the genus is another question. The buccula of most North Amer-

ican drymines appears from the side as a prominent lobe obscuring the base

of the labium. Viewed ventrally, the bucculae extend posteriorly as carinae

that join to enclose a gular region. In all Eremocoris, including the cypress

bug, these carinae extend to or nearly to the base of the head, enclosing a

posteriorly tapering gular region. In Togodolentus, the carinae extend only

to the level of the anterior margin of the eye, and the enclosed gular region

is parallel-sided.

Further, most North American drymines have the lateral margins of the

pronotum to some degree explanate. (Thylochromus is an exception.) In

Togodolentus, the explanate margin is wider than in any other North Amer-

ican drymine (at the middle, wider than the middle part of the second an-

tennal segment). In species of Eremocoris, these margins are not as wide

as the second antennal segment at its midpoint. Given this information, I

conclude that the cypress bug is a true Eremocoris, and that the genus

Togodolentus should be retained.

Eremocoris cupressicola, new species?

(Fig. 1)

Head.—Vertex obscurely roughened, elevated between eyes, obscurely

covered with short, sparse hairs, trichobothrial hairs very long, prominent;

length 2.10, width including eyes 1.95, anteocular length 1.26, antenniferous

tubercle length 0.42, eye length 0.51, eye width 0.36, interocular space 1.17,

bucculae most prominent as anteriorly projecting lobes, continuing as low

carinae to near base of head, enclosing a tapering gular region; labium just

exceeding posterior coxae, reaching base of abdomen, first segment just

exceeding anterior margin of prosternum, segment lengths from base 1.95,

2.70, 1.92, 0.69; antennae clothed with short, appressed hairs, segments I,

II, and III with a few longer hairs apically and segment I with three setae

basally on medial surface, clypeus not reaching midpoint of segment I, seg-

ment lengths from base 1.26, 2.40, 1.98, 1.92.

Pronotum.—Sparsely clothed with long, erect setae, anterior lobe ob-

scurely roughened and punctate, collar region and lateral explanate margins

delimited by row of punctures, posterior lobe moderately punctate, distance

between punctures from diameter of a puncture to three times this distance;

medial length 2.70, greatest length 3.09, anterior and posterior margins deep-

ly emarginate, anterior lobe prominent, swollen, lateral explanate carina of

even width, about as wide as diameter of second antennal segment at mid-

dle, becoming wider between lobes, posterior lobe poorly differentiated

VOLUME 55, NUMBER 2 151

Fig. 1. Eremocoris cupressicola Ashlock, new species, dorsal view, holotype.

152 PAN-PACIFIC ENTOMOLOGIST

from anterior lobe, median length of anterior lobe 1.86, median length of

posterior lobe 0.84, scutellum with surface curved down laterally, clothed

with erect hairs and punctures like those on posterior lobe of pronotum,

length 1.86, width 1.80.

Hemelytra.—Brachypterous, reaching abdominal segment V, clavus and

corium with erect setae similar to those on pronotum, veins not evident,

length of corium 4.35, length of claval commissure 1.20, membrane without

evident veins, greatest width 1.86, greatest length 0.72.

Legs.—Fore femora greatly incrassate, width 1.35, length 3.39, armed

beneath with two ranks of spines, with one large subapical spine on inner

rank, accompanied by five small spines basally and four small spines api-

cally, outer rank with three small apical spines. Fore tibia curved, with

small tubercles on inner surface. Hind tibia with entire surface covered with

long, erect setae three times as long as diameter of tibia.

Color.—Rather uniform dark reddish brown; acetabulae, coxae, lateral

margins of hemelytra a little paler; tarsi yellow.

Length.—Female holotype 7.50; females, range 6.60 to 7.95, mean 7.52;

males, range 6.90 to 7.05, mean 6.95.

Holotype female (California Academy of Sciences).—California, Marin

Co., Carson Ridge, under bark of Cupressus sargenti, I-22-1957 (J. A.

Chemsak).

Paratypes (California Academy of Sciences and the author’s collection).—

All California. 1 male, same data as holotype; 1 female, same data (P. D.

Ashlock); 1 female, same data but I-9-1957 (J. A. Powell); 1 female, same

locality but beating Cupressus sargenti, II-1-1958 (P. D. Ashlock); 2 fe-

males, same locality, V-6-1962 (C. W. O’Brien); 1 female, same locality,

XIJ-15-1962 (J. A. Chemsak); 2 females, same locality, found dead under

bark of dead Cupressus sargenti, VIII-9-1978 (P. D. Ashlock & E. Rogers);

1 male, same locality, habitat, date, collectors, but collected as nymph,

emerged VIII-15-1978; 2 females, same locality, under board, [X-24-1963

(P. D. Ashlock & N. T. Davis); 1 female, Marin Co., Cypress Ridge, V-29-

1920 (E. C. Van Dyke) [allotype of Eremocoris semicinctus Van Duzee]; 1

female, Marin Co., Fairfax, V-11-1919 (E. P. Van Duzee); 1 male, Alameda

Co., Cedar Ridge, IIJ-22-1931 (E. C. Van Dyke); 1 female, Sonoma Co., 1

mi NE Occidental, V-17-1964 (C. W. O’Brien); 1 male, Sonoma Co., 2 mi

E Camp Meeker, XI-24-1962 (P. D. Ashlock); 1 female, same data (C. W.

O’Brien); 1 female, Lake Co., St. Helena Creek, IIJ-11-1951 (J. Helfer); 2

females, Lake Co., Highland Springs, V-10-1932 (R. L. Usinger); 1 female,

Lake Co., Middletown, Putah Creek, V-11-1928 (E. P. Van Duzee).

In addition to the form of the brachyptery and the uniform color, distinc-

tive features of E. cupressicola include the longest head of any North

American member of the genus: the anteocular distance is greater than the

interocular distance. The males have strongly curved fore tibiae, with sev-

VOLUME 55, NUMBER 2 153

eral short spines on the inner surface. The first of my specimens were col-

lected under the bark of cypress trees in January, where the insects were

probably overwintering. Beating the trees themselves has produced other

specimens, and the best results have come from beating branches that have

open cones with seeds. Presumably the bugs feed on the seeds, competing

with Kleidocerys obovatus (Van Duzee) (Lygaeidae, Ischnorhynchinae),

which is common in the same habitat.

In the twenties and thirties, such collectors as E. P. Van Duzee and E.

C. Van Dyke referred to a specific locality in Marin County, California, as

‘‘Cypress Ridge,’’ and some specimens of E. cupressicola bear this label.

The correct name for this locality is Carson Ridge. The cypress forest lies

past a locked gate at the end of Carson Road, which leaves the main road

in the town of Woodacre.

Earlier keys (Barber, 1918; Torre-Bueno, 1946) to North American dry-

mine genera combine this tribe with the Lethaeini and place the genus Thy-

lochromus in the tribe Rhyparochromini. Slater and Baranowski (1978) do

not distinguish tribes in their key, and omitted Togodolentus and Thylo-

chromus because of the rarity of their species. Since no complete and correct

key to genera has ever been provided, and since a better separation of

Eremocoris and Togodolentus has been achieved, a new key to the six

genera of Drymini found north of Mexico follows. The only other Western

Hemisphere genus of Drymini listed in the Slater catalogue (1964) is Scy-

thinus Distant, whose only species, S. splendens Distant, is not available

for study. However, the key should be useful for Mexico as well. The most

recent characterization of the Drymini is that of Sweet (1967), and the group

can be recognized by the two trichobothria (not three) that are placed an-

teriorly on abdominal segment five.

Key to the Genera of Drymini of America North of Mexico

1. Lateral margin of pronotum angulate, not foliaceously expanded;

brachypterous forms without a trace of membrane; Pacific Coast

SE a tas Gate a pe ashe SE ses OURS Mara diel Thylochromus Barber

Lateral margins of pronotum foliaceously expanded at least between

anterior and posterior lobes; brachypterous forms with obvious

eel rele E ANS Pe ics Peed Gee & o oe ens Soe ae ea eee p.

2. Ventral abdominal sutures straight and reaching lateral margins; flat-

tened, cone-living bugs ...............000- Gastrodes Westwood

Ventral abdominal suture [V—V curving anteriorly and not reaching

lateral abdominal margins; robust, mostly ground-living bugs.... 3

3. First antennal segment shorter than distance between eyes; apex of

clypeus reaching at least to middle of antennal segment! ....... 4

First antennal segment longer than distance between eyes; apex of

clypeus not reaching middle of antennal segment I............. 3

154 PAN-PACIFIC ENTOMOLOGIST

4. Antennae densely covered with semierect hairs that are longer than

diameter of segments; east of Rocky Mts........... Drymus Fieber

Antennae with only an occasional erect hair, most hairs appressed

and shorter than diameter of segments ....... Scolopostethus Fieber

5. Bucculae extending posteriorly as carinae only as far as level of

anterior margin of an eye, enclosing a parallel-sided gular region;

width of lateral pronotal expansions at middle of anterior lobe

greater than diameter of antennal segment II measured at its mid-

Le Ct PIAA epee tc et Dae Eas, ele Ag 4 oes ae Togodolentus Barber

Bucculae extending posteriorly as carinae to base of head, enclosing

a gular region that narrows posteriorly; width of lateral pronotal

expansions at middle of anterior lobe not wider than diameter of

antennal segment IJ measured at its middle ..... Eremocoris Fieber

Acknowledgments

I am grateful to Dr. Paul H. Arnaud, California Academy of Sciences, for

the loan of specimens of E. cupressicola, and for permitting me to study

Van Duzee’s types of the genus Eremocoris. The excellent illustration was

executed by Mr. S. Thurston, University of Connecticut, Storrs.

Literature Cited

Barber, H. G. 1918. Synoptic key to the Lygaeidae (Hemiptera) of the United States, Part II:

Rhyparochrominae. Psyche 25: 71-88.

Slater, J. A. 1964. A Catalogue of the Lygaeidae of the World. 2 vols. University of Con-

necticut, Storrs.

Slater, J. A., and R. M. Baranowski. 1978. How to Know the True Bugs (Hemiptera-Heterop-

tera). Wm. C. Brown Co., Dubuque, Iowa.

Sweet, M. H. 1967. The tribal classification of the Rhyparochrominae (Heteroptera: Lygaei-

dae). Ann. Entomol. Soc. Amer. 60: 208-226.

Sweet, M.H. 1977. Elevation of the seedbug Eremocoris borealis (Dallas) from synonymy with

Eremocoris ferus (Say) (Hemiptera: Lygaeidae). Entomol. News 88(7-8): 169-176.

Torre-Bueno, J. R. de la. 1946. A synopsis of the Hemiptera-Heteroptera of America north

of Mexico, Part III: Family XI—Lygaeidae. Entomol. Amer. 26: 1-140.

Van Duzee, E. P. 1921. Characters of some new species of North American hemipterous

insects, with one new genus. Proc. Calif. Acad. Sci. (Ser. 4), 11(10): 111-134.

Footnotes

1 Contribution no. 1627 from the Department of Entomology, University of Kansas, Law-

rence, Kansas 66045.

2 All measurements in millimeters.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 155-158

A NEW CALIFORNIA SPECIES OF PODABRUS

(COLEOPTERA: CANTHARIDAE)

KENNETH M. FENDER

835 Ashwood Avenue, McMinnville, OR, 97128

The following new (to science) species is named for its principal collector:

Daniel K. Young, Michigan State University, an avid student of the Pyro-

chroidae and of the pedilid genus Pedilus.

Podabrus youngi, new species

Head pale flavous, becoming black behind on the dorsal surface, black

area commencing at anterior margin of eyes and deeply arcuately receding

medially to posterior margin of eyes, behind eyes extending along lateral

median line of neck, not attaining base of neck; apical margin of last segment

of maxillary palpi narrowly infuscate; labial palpi piceous; antennae becom-

ing infuscate beyond middle of fourth segment. Pronotum and scutellum

flavous. Elytra black. Head, prothorax beneath and legs flavous. Meso-

thorax and metathorax black. Abdomen piceous. Pubescence fine, short and

Sparse, aureous on pale portions, black on black portions.

Male.—Head shining, as wide as pronotum, moderately rapidly narrowed

behind eyes, eyes moderately large and prominent. Clypeus impunctate save

for fine close punctures along anterior margin. Head finely sparsely puntured

between eyes, punctures separated by two to three times their diameters,

a little more coarsely punctured behind, becoming rather coarsely rugose

punctate on neck; an evident arcuate interocular ridge extending from eye

to eye; apical segment of maxillary palpi elongate triangular, twice as long

as wide, widest medially, inner side straight, apical side arcuate, outer side

shallowly concave; antennae slender, extending to about apical third of

elytra, third segment shorter and thicker than second, fourth segment half

again as long as third, intermediate segments about four times as long as

wide. Pronotum shining, slightly transverse, widest medially, lateral margins

arcuate from obliquely angulate hind angles to rounded anterior angles, disc

broadly explanate at anterior angles, broadly and rather deeply reflexed at

posterior angles, anterior margin shallowly concave, posterior margin more

deeply concave and deeply guttered, convexities moderately high and un-

evenly reniform in shape, median longitudinal impressed line feebly indi-

cated and not eroded, discal punctures fine and sparse. Scutellum shining,

finely sparsely punctured, subtriangular, sides straight, apex sharply round-

ed. Elytra shining, parallel sided, about three times as long as their width

156 PAN-PACIFIC ENTOMOLOGIST

Figs. 1-3. Podabrus sierrae. Fig. 1. aedeagus of male, dorsal view. Fig. 2. same, lateral

view. Fig. 3. same, ventral view. Figs. 4-6. Podabrus youngi. Fig. 4. aedeagus of male,

dorsal view. Fig. 5. same, lateral view. Fig. 6. same, ventral view.

at the humeri, finely rugose punctate basally, becoming more coarsely so

apically. Thorax shining beneath, pubescence of mesothorax and metatho-

rax cinereous, rather long and somewhat depressed. Abdomen dull, pubes-

cence shorter and more dense than on mesothorax and metathorax. Legs

slender and long, protibiae and metacoxae not sexually modified, all claws

broadly cleft. Aedeagus as in Figs. 4-6. Length (head extended) 14 mm.

VOLUME 55, NUMBER 2 157

Female.—Similar to male. Head narrower than pronotum, eyes smaller

and less prominent. Pronotum more transverse, sides more rounded. Length

(head extended) 15 mm.

Holotype male and allotype female.—California, Mariposa County, Sierra

National Forest, Summerdale Campground, 11-15 June 1973, D. K. & D.

C. Young, in the collection of the California Academy of Sciences at Mr.

Young’s request. 208 paratypes with the same data. In a letter, Mr. Young

noted that they were on the foliage of willows. Paratypes also from the

following California localities: Tuolumne Co.: Strawberry (11 miles north),

24-VI-51, R. W. Morgan; Strawberry, 12-VII-52, M. Cazier, W. Gertsch &

R. Schrammel; Dardanelle, 13-VII-52, M. Cazier & W. Gertsch; Crocker

Station, 6 mi s. of Mather, 12-VI-61, C. D. McNeill; Mather, 9-VI-61, D.

R. Miller; Long Barn, 16-VI-61, R. R. Snelling; Miguel Meadows, Swamp

Lake, 24-VI-76, black light, R. P. Allen. Alpine Co.: 4 mi w. of Woodfords,

25-VI-61, A. S. Menke; Woodfords, 28-VI-62, W. E. Simonds; Hope Valley,

18-VII-48, R. C. Bynum; Markleeville, 2-VII-50, collected at light, W. C.

Day. Eldorado Co.: Fallen Leaf Lake, 10-VII-35, 6500 feet, F. E. Blaisdell;

Tallac, VII-99, Van Dyke; Tahoe, Tallac, 5-VIH-15, E. P. Van Dyke. Madera

Co.: Oakhurst, 5-VI-42, on Ceanothus, Arthur J. Waltz; Bass Lake, 14-VI-

34. Mono Co.: 4 mie. of Monitor Pass, 24-VI-62, C. D. McNeill. Mariposa

Co.: Yosemite Valley, 23-VI-21, VanDyke coll.; Yosemite Valley, Mariposa

Grove, 30-VI-37, 6500 feet, E. Herald. Nevada Co.: Cisco, VI. San Ber-

nardino Co.: San Bernardino Mts., 2-VII-17, 7500 feet.

Some of the paratypes are in the collection of the author and the others

are to be deposited in various collections, some of which include: Daniel

K. Young; Michigan State University, East Lansing; California Department

of Food and Agriculture, Sacramento; California Academy of Sciences, San

Francisco; Oregon State University, Corvallis; Museum of Comparative

Zoology, Harvard University; National Museum of Natural History, Wash-

ington, D.C.

The following California localities are also represented but could not be

satisfactorily placed as to county: Sugar Pine, 21-VII-33, 25-VII-33, E.

Schiffel. There is more than one Sugar Pine in California. Myers, 28-VI-30,

A. T. McClay. This may be a misspelling of Meyers, Eldorado County.

Deerpark Inn, July. I was finally able to find a Deerpark, Placer County in

a 1938 Shell Oil Company road map of California.

In Fall’s (1928) review of Podabrus this species would key to P. sierrae

Fall. In all of the specimens of P. sierrae examined, the black of the back

of the head always extends to and includes the base of the neck. The scu-

tellum is black. The male aedeagus is less elongate and the ventral lobes are

less strongly produced (Figs. 1-3).

In P. youngi the coloration of the head is quite variable. The black area

may be reduced to a rather narrow arcuate interocular fascia. It may be

158 PAN-PACIFIC ENTOMOLOGIST

expanded beneath to nearly attain the gular sutures. However, in the more

than 200 specimens examined, the black never does attain the base of the

neck. The scutellum is flavous.

Literature Cited

Fall, H. C. 1928. A review of the North American species of Podabrus. Entomol. Amer.

8(new series)2: 65-103.

PAN-PACIFIC ENTOMOLOGIST

Vol. 55, No. 2, pp. 159-160

SCIENTIFIC NOTE

NOTES ON THREE OREGON LEPTURINE

CERAMBYCIDAE (COLEOPTERA)

Cortodera militaris constans L. & C.—Linsley and Chemsak (1972, Univ.

Calif. Publ. Entomol. 69: 109-111) segregated C. militaris into three sub-

species, listing only the nominate form as occurring in the Pacific North-

west. Concurrently, they described the subspecies C. m. constans from

northeastern California, which might suggest that this taxon also inhabits

adjacent regions of Oregon. Collections verifying this assumption have re-

cently been made in Lake Co.: (6 6d, 3 2 ) 3.3 and 5.4 mi SE Quartz

Mt., VI-16-1977 on Ranunculus occidentalis Nutt. (R. L. Penrose); (3

636,15 22 ) 14.8 mi N. Lakeview, VI-14-1977 on R. occidentalis and

Potentilla gracilis Dougl. ex Hook. (R. L. Penrose); (13 64, 30 2 2) VI-8-

1978 on R. occidentalis (R. L. Penrose and R. L. Westcott). Although these

specimens are clearly best placed with constans (on the basis of anatomical

similarities and geographic proximity), they exhibit more variation in elytral

coloration than is evident in Modoc County material. The name constans

was given on the basis of the uniformity of the type series, i.e., ‘‘all indi-

viduals black with red humeri.’’ In the Lake County material cited above,

60 specimens (86%) are typical constans, 9 (13%) have elytra brown/vittate

and 1 (1%) is wholly black. Inclusion of Oregon populations would therefore

require a redefinition of constans to encompass predominately black pop-

ulations in which the red humeral condition is usually expressed. It should

also be noted that four of the six specimens from Quartz Mt., in marked

contrast to other Lake County specimens, have the reddish mark vaguely

defined and restricted to the humeral angle. This reduction in maculation

size, combined with the occurrence of a wholly black individual could in-

dicate that transitional populations between constans and typical militaris

remain to be discovered in the Klamath Basin. Although much remains to

be done to unravel the population structure of this species, specimens at

hand suggest a divergence in melanistic tendencies on opposite sides of the

Cascade Range. Whereas constans is typically melanic, black individuals

comprise only 20% of the beetles examined from western Oregon. Another

interesting phenomenon is found in the distribution of local populations with

red and black individuals within the range of the nominate subspecies. All

bicolored specimens seen are from Washington (Tacoma, Olympia and Che-

halis), and Linsley and Chemsak’s statement that black militaris tend to

have reddish humeri would seem to be valid only as regards specimens from

the northern portions of the subspecies range.

Typocerus serraticornis Linsley and Chemsak.—Recently described from

specimens collected in Nevada, Utah, New Mexico and Idaho (1976, Univ.

160 PAN-PACIFIC ENTOMOLOGIST

Calif. Publ. Entomol. 80: 69), this species also occurs in southeastern Or-

egon. A series was collected in Harney Co., Little Cottonwood Creek, VI-

6-1978 (R. L. Penrose and R. L. Westcott). Beetles were sporadically abun-

dant in the flowers of Sphaeralcea grossulariaefolia (Hook. and Arn.) Rydb.

(currant-leaved desert mallow) which was growing intermixed with the lar-

val host, Sand Dropseed (Oryzopsis hymenoides (Roem. and Schult.) Rick-

er). Occasional Typocerus were also taken on flowers of Eriogonum oval-

ifolium Nutt. An additional collection was made 22 mi NW of Denio

Junction, Humboldt Co., Nevada, on June 7. At this site, beetles were quite

abundant but very localized, nearly all having been encountered on a single

165 square meter area of sand, covered almost exclusively with O. hyme-

noides. They were observed flying slowly about, and sitting and mating on

this grass. Occasionally, females were noted crawling on the surface of the

sand around the plant bases in search of oviposition sites. Larvae were

found at both localities boring in culm bases, below the soil surface, indi-

cating at least a two-year life cycle.

Ortholeptura obscura (Swaine and Hopping).—There have been but three

prior references to this species, all of which are descriptive (Swaine and

Hopping, 1928, Nat. Mus. Canada Bull. 52: 56; Hatch, 1971, Univ. Wash.

Publ. Biol. 16: 132; Linsley and Chemsak, 1976, Univ. Calif. Publ. Entomol.

80: 133). These references are apparently based upon a single male and

female specimen. The literature has been reflective of specimen availability

and I have seen only two specimens of obscura (both from northeastern

Oregon) in Pacific Northwest collections during the past decade. Due to the

rarity of this species, and the absence of any published biological informa-

tion, the following observations are presented. On July 25, 1978, five spec-

imens were collected at Wallowa Lake State Park, Wallowa County, Ore-

gon, as follows: 1 6 was attracted to white light; 1 2 was found clinging to

the underside of a freshly fallen branch of Pseudotsuga menziesii (Mirb.)

Franco (Douglas Fir); 1 teneral adult 6 and 2 pupae, 1 d, 1 2, were col-

lected from pupal cells in the dry, hard outer sapwood near the top of an

old Douglas Fir stump. A single ¢ was swept from foliage of Snowberry,

Symphoricarpos sp. at Field Spring State Park, Asotin County, Washington,

VIII-30-1966 by R. L. Westcott.

Special thanks is extended to Dr. J. D. Lattin, Curator, Oregon State

University Entomological Collection, Corvallis, for permission to study

specimens of C. militaris present in the recently acquired M. Hatch Col-

lection.

Richard L. Penrose, Oregon Department of Agriculture, Salem, Oregon

97310.

January issue mailed 21 June 1979

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