Vol. 50

JANUARY 1974

No. 1

THE

Pan-Pacific Entomologist

PINTO — Courtship behavior in Linsleya compressicornis and its taxonomic

significance (Coleoptera: Meloidae)

SCOTT — Adult behavior and population biology of Poladryas minuta, and

the relationship of the Texas and Colorado populations (Lepidoptera:

Nymphalidae)

HARADON — Vaejovis spicatus : A new scorpion from California (Scor-

pionida: Vaejovida)

LANGSTON — The maritime earwig in California (Dermaptera: Carcinophor-

idae)

CHEMSAK AND LINSLEY — Notes and descriptions of some lycid-like Neo-

tropical lepturine Cerambycidae (Coleoptera) _

KOMPFNER — Larvae and pupae of some wrack dipterans on a California

beach (Diptera: Coelopidae, Anthomyiidae, Sphaeroceridae)

HORN — Observations on primary and secondary parasitoids of California oak

worm, Phryganidia calif ornica , pupae (Lepidoptera: Dioptidae)

DAILEY, PERRY, AND SPRENGER — Biology of three Callirhytis gall wasps

from Pacific slope Erythrobalanus oaks (Hymenoptera: Cynipidae)

ALCOCK — Observations on the behavior of Mallophora fautrix Osten Sacken

(Diptera: Asilidae)

STEINER — Unusual caterpillar-prey records and hunting behavior for a

Podalonia digger wasp: Podalonia valida (Cresson) (Hymenoptera:

Sphecidae)

McDONALD, HEED, AND MIRANDA — The larval nutrition of Minettia

flaveola and Phaonia parviceps and its significance to the Hawaiian

leaf -breeding Drosophila (Diptera: Lauxaniidae, Muscidae, Dro-

sophilidae)

SCIENTIFIC NOTES 22, 52, 67, 72, 84, 85, 86,

BOOK REVIEW

BOOK NOTICE

RECENT LITERATURE 27,

PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY

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in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

THE PAN-PACIFIC ENTOMOLOGIST

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The Pan-Pacific Entomologist

Vol. 50

January 1974

No. 1

Courtship Behavior in Linsleya compressicornis

and its Taxonomic Significance 1

(Coleoptera: Meloidae)

John D. Pinto

Department of Entomology, University of California, Riverside 92502

The systematic position of the small, western North American genus

Linsleya MacSwain has been the subject of recent controversy. Although

members of this group were originally placed in Lytta on the basis of

superficial adult resemblance (Horn, 1873), larval anatomy has now

convinced most workers that they belong near Epicauta and Pleuro-

pompha in the Epicautina (MacSwain, 1951, 1956; Selander, 1955,

1964a; Arnett, 1960; Werner, Enns and Parker, 1966). Yet, other

authors (Kaszab, 1959; Gupta, 1965) continue to treat Linsleya as al-

lied to Lytta. This study of courtship behavior was undertaken to pro-

vide additional information bearing on this question. Courtship in

Linsleya ( Linsleya ) convexa (LeConte) was studied previously but

failed to indicate subtribal affinity (Selander and Pinto, 1967). The

behavior of a second species, L. ( Linsleyina ) compressicornis (Horn),

is described below. This species is of considerable interest since its

courtship is remarkably similar to that of several species of Epicautina

and unlike any behavior known in the Lyttina 2 .

Adults utilized in this study consisted of 30 males and 28 females

collected from their host plant, Menodora spinescens Gray (Oleaceae),

2 miles E of Big Pine, Inyo County, California, on 9 June 1973. Sexual

isolation was effected 12 hrs after collection and was maintained when-

ever observations were not in progress. All observations were made in

the laboratory from 12-20 June 1973, at 26-30 °C. Specimens were

maintained and observed in plastic cages IIV 2 cm long and 8 cm wide.

1 This study was supported by Grant GB-30907 from the National Science Foundation.

2 This paper follows Selander’s (1964b) classification in treating the Lyttina and Epicautina as

subtribes within the Lyttini. Other authors (e.g. MacSwain, 1956) give these two taxa tribal status.

The Pan-Pacific Entomologist 50: 1-8. January 1974

The Pan-Pacific Entomologist

Ethological data are based on six hours of observation utilizing nu-

merous randomly selected pairs, and include 200 feet of motion picture

films.

L. compressicornis has been divided into two subspecies by Selander

(1955). This study is based on members of the nominate form.

Courtship Behavior

All components of courtship in L. compressicornis are performed by

the male while positioned behind the female (posterior position). At

no time does he mount her, or overtly contact her with his legs in any

other way. For descriptive purposes courtship is divided into three

phases, a preliminary phase, a display phase, and a genital phase.

At the onset of the preliminary phase the male orients directly behind

a quiescent female. His maxillary palpi are often extended and ir-

regularly touch the female’s pygidium. The position and state of the

male’s antennae vary. They may be held motionless and out to the side

or, more frequently, with segments II-XI directed downward at an ap-

proximate right angle to segment I and the anterior surface of each

antenna facing the side of the female’s last abdominal segment. While

in the latter position the antennae are often rapidly vibrated antero-

posteriorly. Periods of vibration vary in duration from one to 20 sec

and comprise, on the average, 50-60% of total courtship time. Antennal

vibration is usually repeated several times before the start of the display

phase. Due to decamping by the female, however, some bouts never

progress beyond preliminary phase activity.

Although antennal vibration is the dominant component of courtship

in terms of time, its significance is not clear. Contact with the female

apparently never occurs. Because of this and the lack of an obviously

correlated female response, I have excluded this act from the display

phase.

Display in L. compressicornis consists of a single act, antennal press-

ing (Fig. 1). This act is always preceded by antennal vibration. Prior

to pressing the antennae are rotated so that their anterior surface is

directed forward and, at the same time, they are moved slightly postero-

medially to a position above the midline of the female’s abdomen. Tac-

tual stimulation immediately follows and consists of the male pressing

down on the last two abdominal tergites of the female one to five times

in rapid succession. Both antennae are employed synchronously with

the anterior surface of segments IV-XI making contact. A determination

of the number of presses normally provided during each bout of display

is complicated by female activity. Most females thwart courting males

Vol. 50, No. 1, January 1974

Figs. 1-2. Courtship behavior in L. compressicornis. Fig. 1. Display phase.

Male performing antennal pressing. Fig. 2. Genital phase. Male attempting gen-

ital insertion.

by either decamping, kicking, or abruptly elevating their abdomen at

the onset of display. The majority of bouts, however, consist of three

presses. Display was not common in this study. It occurred, on the

average, only once during every four minutes of courtship.

Display is invariably followed by a genital insertion attempt (genital

phase) (Fig. 2). In attempting insertion the male maintains his pos-

terior position, elevates his fore legs and, with genitalia partially ex-

truded, curls the apex of his abdomen forward through his middle and

The Pan-Pacific Entomologist

Figs. 3-4. Scanning electron micrographs of right antenna of L. compressicornis

(anterior view). Fig. 3. Male (60x)- Fig. 4. Female (50x)-

hind legs toward the gonopore of the female. If, as is usually the case,

the female does not allow mating, he either returns to preliminary phase

activity or ceases courting entirely. Genital insertion attempts only oc-

cur after display. In my observations their duration varied from seven

to 45 seconds.

The display and genital phases appear to be tied behaviorally in this

species since insertion is always attempted immediately after pressing.

This is even true when the female decamps during display. In this case

the male quickly pursues the female and upon continuing courtship sev-

eral seconds later resumes with an insertion attempt. Situations where

the female successfully eludes the male were not observed.

Only a single mating was observed. This particular male inserted im-

mediately after display, and within less than five seconds turned to his

right and assumed the linear position without contacting the female with

his fore legs. The duration of this mating was not recorded.

Anatomical Correlates of Courtship

Antennal modifications in the male of L. compressicornis are the

most obvious correlates of courtship. As shown in Figs. 3 and 4, seg-

ments III-XI of the male are more strongly broadened than those of the

female, and their anterior surface is concave rather than convex. Ad-

ditional modification is found in antennal surface structure. Most of the

male’s segments have fewer setae on the anterior (contacting) surface,

and segment XI is entirely denuded at its base. Examination of this

surface at high magnification with the scanning electron microscope re-

Vol. 50, No. 1 , January 1974

veals an abundance of minute cuticular pores. These are especially dense

at the base of segment XI, the area of greatest density appearing as a

small ellipse at 60X (Fig. 3). The function of these pores is unknown.

Their association with chemical stimuli has been suggested for other

meloines (Matthes, 1969; Pinto, 1973).

Another modification in males of L. compressicornis is the highly

developed fore tarsal pads. The pale, apically flared setae which com-

prise the pads in this species are best developed on the first three seg-

ments. In Linsleya convexa, similarly modified fore tarsal pads in the

male are considered an adaptation for grasping the female while he is

mounted on her (Selander and Pinto, 1967). In L. compressicornis ,

however, the male does not contact the female with his legs. Instead,

the pads may aid the peripatetic males in maneuvering on the host plant

surface during their persistent search for females and during courtship.

Comparative Behavior and Discussion

L. convexa is the only other species of Linsleya for which courtship

data are available. This species shares few similarities with L. compres-

sicornis besides traits typical of meloines in general 3 . In L. convexa the

male courts entirely from a fully mounted position (dorsal position),

and display consists of rubbing the dorsum of the female with his hind

two pair of tarsi (Selander and Pinto, 1967). The only noteworthy

similarity is that males of both species fail to manipulate the antennae

of the female but stimulate her dorsum instead.

Courtship in L. compressicornis is remarkably similar to that of

Pleuropompha and certain species of Epicauta. In both species of Pleu-

ropompha and in Epicauta ( Epicauta ) puncticollis Mannerheim (Pinto,

1973), and in E. ( Epicauta ) ruidosana Fall (Pinto, unpubl.) the male

also courts entirely from an unmounted posterior position. The genital

phase is performed in the same manner in all five species, and although

the specifics of display vary, employment of the antennae by the male

is characteristic of all but E. ruidosana. In all other nominate Epicauta

studied, the genital phase and often the display phase as well are per-

formed from a dorsally mounted position (Pinto, 1973). A dorsal

mount, at least during part of courtship, is typical of Meloinae. Its com-

plete absence has been reported only in the above-mentioned species.

The act of antennal pressing in L. compressicornis is roughly similar

to display in some other species of nominate Epicauta , namely E. par-

dalis LeC., E. magnomaculata Martin, and E. ventralis Werner (Pinto,

3 Courtship traits typical to the Meloinae are discussed by Selander (1964b).

The Pan-Pacific Entomologist

MS). In these three species the posteriorly positioned male rubs or

presses the dorsum of the female with his antennae. The specifics of this

act in these Epicauta are sufficiently different, however, as to preclude

homology with antennal pressing. As in both species of Linsleya , dis-

play in all nominate Epicauta that have been studied involves the stimu-

lation of the female’s dorsum in some way (Pinto, 1973), and is never

directed toward her antennae. Antennal manipulation is found in most

other meloine taxa (Selander, 1964b; Pinto, 1972) including Epicauta

subgenus Macrobasis (Selander and Mathieu, 1969).

With respect to courtship then, L. compressicornis and certain species

of nominate Epicauta are more similar to each other than to congeners.

Unfortunately, we can not yet speculate knowledgeably as to whether

posterior courtship, their shared pattern, represents the primitive or

derived state in the Epicautina. However, it is clear that either posterior

or dorsal courtship has been derived independently in the subtribe since

the two patterns occur in both Linsleya and Epicauta. Such marked in-

terspecific variation in the position of the courting male is surprising

considering its constancy in other meloines. In other taxa (e.g. Meloe,

Lytta , Eupompha , Pyrota ) dorsal mounting is typical, and the position

assumed by the male varies only slightly between species. The adaptive

significance of posterior versus dorsal courtship can not be profitably

discussed without analyses of several aspects of reproductive behavior

and related ecological factors.

The similarities of behavior in L. compressicornis and certain Epicauta

are marked. To the contrary, there are no important parallels to court-

ship in any lyttine studied so far. In species of Lytta the male invariably

courts from a fully mounted position, and display, when present, in-

volves the manipulation of the female’s antennae (Selander, 1960; Mat-

thes, 1972; Gerber and Church, 1973; Pinto, unpubl.). A similar pat-

tern has been reported for the lyttine genus Lydus (Cros, 1912).

Conclusion

In spite of the failure of adult anatomy to clearly indicate relationship,

both larval and ethological features can now be cited in support of plac-

ing the five species of Linsleya in the Epicautina. The similarities of

the first instar larvae of Linsleya, Epicauta and Pleuropompha are nu-

merous (see MacSwain, 1956) and, in themselves, leave little doubt as

to the affinity of these genera. Selander (1964a) has also pointed out

that larvae of Linsleya sphaericollis (Say) appear to be parasitoids of

grasshopper egg pods (as in Epicauta) rather than of bee nests (as in

Vol. 50, No. 1, January 1974

Lytta and related genera). The behavioral data presented above are of

importance since they clearly support these conclusions based on larval

anatomy and bionomics, and, furthermore, suggest that Linsleya is most

similar to the nominate subgenus of Epicauta.

As mentioned above, Kaszab (1959) and Gupta (1965) continue to

place Linsleya near Lytta 4 . Kaszab ’s rationale for this is not clear. Al-

though his 1959 classification of the Meloidae is primarily based on

wing venation, the wings of Linsleya are poor indicators of subtribal

affinity as they are similar to those of both Epicauta and Lytta. Gupta’s

reassignment of Linsleya was prompted by the absence of two structures

in L. sphaericollis , V-shaped proventricular folds and well developed

stomadaeal valves. Both traits are present in Epicauta and Pleuro-

pompha ; other epicautine genera ( Psalydolytta , Denier ella and Anom-

alonchus) were not studied. To reassign Linsleya solely on this basis

is not convincing. If Linsleya is the most primitive genus of epicautine

as both MacSwain (1951) and Selander (1955) believe, its lacking cer-

tain specializations of the more derived genera should not be surprising.

Acknowledgments

Assistance by Messrs. Derham Giuliani and Gregory R. Ballmer is

gratefully acknowledged. Figures 1 and 2 were prepared by Mr. Carl

T. Conley. Identification of the host plant of L. compressicornis was

kindly provided by Mr. Oscar F. Clarke (U. C. Riverside Herbarium).

Literature Cited

Arnett, R. H., Jr. 1963. The Beetles of the United States (a Manual for Iden-

tification.) Wash. D.C., 1112 pp.

Cros, A. 1912. Lydus algiricus L. Ses moeurs — Sa larve primaire. La Feuille

Jeunes Natur., 42 : 78-86.

Gerber, G. H. and N. S. Church. 1973. Courtship and copulation in Lytta nut-

talli (Coleoptera: Meloidae). Can. Entomol., 105: 719-724.

Gupta, A. P. 1965. The digestive and reproductive systems of the Meloidae

(Coleoptera) and their significance in the classification of the family.

Ann. Entomol. Soc. Amer., 58: 442-474.

Horn, G. H. 1873. Revision of the species of several genera of Meloidae of the

United States. Proc. Amer. Phil. Soc., 13: 88-117.

Kaszab, Z. 1959. Phylogenetische Beziehungen des Fliigelgeaders der Meloiden

(Coleoptera), nebst Beschreibung neuer Gattungen und Alien. Acta

Zool. Acad. Sci. Hung., 5: 67-114.

MacSwain, J. W. 1951. A new genus of Meloidae from North America. Pan-

Pac. Entomol., 27: 58.

4 In a recent classificatory outline of the Meloidae, Kaszab (1969, Mem. Soc. Entomol. Ital., 48:

243) lists Linsleya as an epicautine.

The Pan-Pacific Entomologist

MacSwain, J. W. 1956. A classification of the first instar larvae of the Meloidae

(Coleoptera) . Univ. Calif. Publ. Entomol., 12: 1-182.

Matthes, D. 1969. Die Fachelbalz von Cerocoma schafferi (L.) (Coleopt.,

Meloidae). Zool. Anz. Suppl. — Verhandlungen Zool. Ges., 33: 316-322.

Matthes, D. 1972. Die Balz der Spanischen Fliege ( Lytta vesicaloria L.). Zool.

Anz., Leipzig, 188: 441-447.

Pinto, J. D. 1972. Comparative courtship behavior of Negalius, Phodaga and

Cordylospasta , three closely related genera of blister beetles (Coleoptera:

Meloidae). J. Kans. Entomol. Soc., 45: 459-476.

Pinto, J. D. 1973. Sexual behavior in the genus Pleuropompha LeConte: A

new mating display in blister beetles (Coleoptera: Meloidae). Can.

Entomol., 105: 957-969.

Selander, R. B. 1955. The blister beetle genus Linsleya (Coleoptera: Meloidae).

Amer. Mus. Novitates, 1730: 1-30.

Selander, R. B. 1964a. The systematic position of the genus Linsleya (Coleop-

tera: Meloidae). Proc. Entomol. Soc. Wash., 66: 216.

Selander, R. B. 1964b. Sexual behavior in blister beetles (Coleoptera: Meloi-

dae) I. The genus Pyrota. Can. Entomol., 96: 1037-1082.

Selander, R. B. 1960. Bionomics, systematics, and phylogeny of Lytta, a genus

of blister beetles (Coleoptera, Meloidae). Univ. 111. Biol. Monogr., 28:

1-295.

Selander, R. B. and J. M. Mathieu. 1969. Ecology, behavior ,and adult anatomy

of the Albida Group of the genus Epicauta (Coleoptera, Meloidae).

Univ. 111. Biol. Monogr., 41: 1-168.

Selander, R. B. and J. D. Pinto. 1967. Sexual behavior in blister beetles (Cole-

optera: Meloidae) II. Linsleya convexa. J. Kans. Entomol. Soc., 40:

396-412.

Werner, F. G., W. R. Enns and F. H. Parker. 1966. The Meloidae of Arizona.

Agr. Exp. Sta. Univ. Ariz. Tech. Bull., 175.

Vol. 50, No. 1, January 1974

Adult Behavior and Population Biology of Poladryas minuta ,

and the Relationship of the Texas and Colorado Populations

(Lepidoptera: Nymphalidae)

James A. Scott

Department of Entomology, University of California, Davis

This paper is part of a comparative study of behavior and movements

of eleven species of diurnal lepidoptera (Hesperiidae and Papilionoidea) ,

emphasizing the relationship between mate-locating behavior and move-

ments (Scott, 1973a; 1973b). Included are studies of mate-locating be-

havior by males, mating, movements and lifespan, oviposition, adult

and larval foodplants, and basking behavior in Poladryas minuta Ed-

wards. In addition I describe experiments in which the Texas sub-

species ( minuta ) was hybridized and backcrossed in nature to the Colo-

rado subspecies ( arachne Edwards), providing much of the behavioral

information.

The species was studied in 1969 on a treeless ridge just northwest of

Cripple Creek, Teller County, Colorado, and in 1972 on Green Moun-

tain, Jefferson County, Colorado. All times are 24-hour standard time.

I thank Roy 0. Kendall, San Antonio, Texas, and Kilian Roever,

Phoenix, Arizona, for helpful information, Jerry A. Powell, University

of California, Berkeley, and Glenn R. Scott, Lakewood, Colorado, for

reviewing parts of the manuscript. The University of California, Berke-

ley, provided a grant for computer time.

Methods

Felt-tipped markers were used to give each individual a different

number using the method of Ehrlich & Davidson (1960). Marks were

placed on the upperside so that the number could usually be determined

without capture. Individuals were marked and individually released

at the site of capture.

The following method of analysis of movements allows direct com-

parison between sexes and between species, detection of change of move-

ments with age, and separation of the velocity and distance aspects of

movements. The capture points for each individual are plotted on sep-

arate maps of the study site. The following statistics were determined

for each individual:

di, tj — distance in meters and time in days between capture i and cap-

ture (i + 1) ; D — sum of all d’s of an individual; R — distance between

The Pan-Pacific Entomologist 50: 9-22. January 1974

The Pan-Pacific Entomologist

the two farthest capture points (range) ; T — time between first and last

capture; V — overall velocity (D/T) ; Vj— velocity between successive cap-

tures (di/tj).

Means of the above statistics are computed for all individuals of each

sex. Midpoint age is the age midway between two successive captures

of an individual. Correlations between the above movement parameters

and age determine whether movement changes with age. Population

size, survival rates, and number of new insects per day were determined

from mark-recapture data using the method of Jolly (1966).

Mate-locating Behavior

Most butterflies use one of two strategies to bring the sexes together

for mating (Scott, 1973b) : in some species males wait at characteristic

sites such as hilltops, gullies, or treetops, and locate females by investi-

gating passing objects ( perching behavior ). In these species females

fly to the perching sites after emergence. In other species males search

for females by flying almost continuously ( patrolling behavior ) . P.

minuta is the only known species of butterfly (and perhaps insect) which

utilizes completely different strategies of mate-location depending on the

time of day. In the morning from about 0700 to about 1215 and espe-

cially from 0800 to 1130, male P. m. arachne perch on hilltops and high

points on ridges. In the afternoon until about 1600, males patrol near

flowers, usually on hillsides or flats. Male P. m. minuta in Texas also

perch in the morning on ridgetops and patrol in the afternoon on hill-

sides.

In the morning, perching males alight on bare spots or rocks and

dart out at passing fluttering objects (usually other male P. minuta , and

often males of Hesperia comma L., another perching, hilltopping spe-

cies). Between investigative flights, males often visit flowers or fly

short distances to adjacent perches. If they encounter a female during

these short flights courtship ensues, resulting in the small percentage

of morning copulations initiated by flying males. If a male investigates

an adjacent perched male, an “encounter” usually results in which both

fly vertically near each other for about 3 m, then separate and return

to the ground or repeat the vertical flight. Passing males may be pur-

sued briefly. Males usually return to the same or a nearby perch after

investigating a passing object, but sometimes move to another spot on

the ridge. In the afternoon, the behavior of males changes radically:

males patrol rapidly near flowers, briefly pursuing male P. minuta or

other butterflies and courting females.

Vol. 50, No. 1, January 1974

Experiments with females dangling from a fishing pole showed that

movement of the wings is necessary to attract a perched male. When

the female fluttered, the male approached and then courted if the fe-

male was dropped to the ground. Usually the female hung motionless,

and was ignored by males, which would sometimes pass by the motion-

less female to investigate a more distant much larger black Papilio but-

terfly ( minuta is mottled orange). Flight is not an absolute require-

ment to attract the male, since in two instances courtship commenced

when a female was carefully placed next to a perched male without dis-

turbing him.

Mating

Virgin females were released before perched males for analysis of

courtship, and many courtships of native individuals were also observed.

To obtain pure minuta matings, the male was released first, then the

female was released near the perched male. Thirty-two completed court-

ships in five male-female combinations (three pure combinations: mi-

nuta $ X minuta $ (1), minuta nympha Edwards S X m. nympha 2

(1), arachne $ X arachne 2 (3) ; two hybrid combinations: arachne

$ X minuta 2 (12), arachne $ X ( arachne S X minuta 2) 2 (15))

and many abortive courtships were observed. Courtship in these five

combinations was identical, including all the behavioral elements de-

scribed below.

In the simplest form of courtship the female alights (usually within

3 m of the male’s perch), the male alights behind her and bends his

abdomen laterally (11 right, 7 left in successful attempts) to copulate.

Four additional components may be present. Male hovering consists

of the male rapidly beating his wings at small amplitude a few cm

above and downwind of the female for a few seconds. The male then

alights behind the female. If the female flies, crawls vigorously away,

or flutters during courtship, the male may hover before alighting be-

hind her again. Male fluttering is similar to hovering except that the

male is on the ground so the wings are moved more slowly, above the

horizontal, and with greater amplitude while the body remains sta-

tionary. Male fluttering occurred rarely (only 4 successful copulations)

and only after alighting while the male was behind the female prior to

attempted copulation. Male nudging consists of the male holding the

wings about 40° above horizontal, the antennae directed backward,

and pushing his head under a spread hind wing of a female, evidently

positioning him alongside the female so that his laterally curved ab-

The Pan-Pacific Entomologist

domen is in position for copulation. If the female’s wings are nearly

closed the male immediately crawls alongside and attempts coupling.

The male usually keeps his wings about 40-60° above horizontal even

when it would appear advantageous to raise them to get closer to the

female. Male nudging was observed in many successful courtships,

especially early in courtship when the female’s wings were spread. Fe-

males usually raised their wings prior to coupling but several males

managed to couple by nudging under the female’s spread wings. Fe-

male fluttering consists of females holding the wings about 40° above

horizontal and fluttering the wings slightly. It occurs while the male is

on the ground behind the female.

Successful courtship last from 2 seconds to 2 minutes after both in-

dividuals alight, but typically requires about 10-15 sec.

Unreceptive virgin females discourage males in a variety of ways.

They may crawl away, turn, fly a short distance, or flutter their wings

(female fluttering) ; they rarely perform the stereotyped rejection dance

of mated females (performed early in three successful courtships; see

below). Occasionally they may lift the abdomen about 30° above hori-

zontal so that the male cannot couple, and they usually keep the wings

spread. Males often overcome moderate female unreceptivity by per-

sistent hovering, nudging, and attempted coupling. The male crawls

or flies behind the female, often hovering briefly when the female flies

or crawls rapidly away, or sometimes he hovers after she flutters her

wings. After many copulation attempts (up to 20-30 ), the female may

become quiescent, raising her wings nearly to the vertical, and moving

her abdomen to the horizontal or slightly below so that the male can

couple. One female whose wings remained spread raised her abdomen

slightly to nearly horizontal so that the nudging male could couple.

The main causes of courtship termination by virgin females are: 1)

the female eludes the male by crawling or flying so that the male can-

not relocate her; 2) after courting a persistently unreceptive female the

male flies away.

Table 1 shows that male hovering and fluttering are associated with

crawling, flying, or fluttering in unreceptive virgin females. In only

five courtships was male hovering or fluttering not associated with

these female responses. The function of hovering and fluttering seems

to be to inform the female of the identity of the male and thereby to

make her more receptive, but the male often hovered before landing and

before the female could respond, suggesting that male hovering has be-

come somewhat ritualized.

Vol. 50, No. 1, January 1974

Table 1. Association of some of the behavioral elements of courtship

in Poladryas minuta subspecies, based on 32 successful courtships with

virgin females. Females less than two hours old indicated by (y) ; two

days old, (o) ; other females were one day old. See text for descriptions

of male and female behavioral elements.

Male Behavior

Hover-

ing &

Flut-

tering

Other*

Total

Hovering

Flut-

tering

Female

Flying

3 ( ly)

Response

Fluttering

Crawling

2 ( ly )

Flying and

Fluttering

Flying and

Crawling

3 (ly)

Fluttering

and

Crawling

5 (3y,lo)

l(o)

None

4(ly,lo)

6(2o)

Total

* Includes nudging or absence of preliminary courtship.

Females mated at ages of one hour to two days. Table 1 shows that

young females were somewhat less receptive than older females, and

that males hover slightly less frequently over older females. Males be-

gin courting on the day of emergence, as one P. m. minuta male perched

and chased 10 objects on a hilltop in a half hour period two hours after

emergence, and attempted to copulate with a female. Another P. m.

minuta male mated one day after emergence.

Mated, unreceptive females have a rather stereotyped rejection dance.

The female, when pursued by a male, slowly flies vertically about three

m, then rapidly returns to the ground. The male frequently cannot

follow her through the downward part of this flight. The dance is re-

peated if the male follows her. This dance is almost identical to the

vertical encounter between two perched males, suggesting that the visual

similarity may function in discouraging courting males. The female

rejection dance occurs in all four taxa studied, and in P. m. nympha

in southern Arizona (Fred Thorne, written communication).

Many of the elements of behavior during courtship seem to depend

on visual cues. The perched male is attracted only to moving objects.

The Pan-Pacific Entomologist

The rejection dance of mated females visually resembles the vertical

encounter between adjacent perching males. Males sometimes hover

over red Castilleja flowers possibly because the colors of the flower and

butterfly upperside are similar ( Castilleja was not used as a nectar

source). Perching males often follow other males which have a slow,

linear flight similar to that of females, and often land beside very young

released males and occasionally hover and try to copulate. One male

grasped a newly emerged male for several minutes, but such homosexual

behavior is uncommon.

Behavior during copulation. The male usually remains motionless

during copulation but the female often opens and closes her wings a

few times after coupling, then usually remains motionless. Upon ter-

mination, the male resumes mate-locating behavior, while the female

remains motionless for several minutes.

Duration of copulation depends on the history of prior mating of

the male. If the male had terminated mating within the previous hour,

copulation lasted 9 hr. and 10 hr. 41 min. (N = 2). If the male had

mated the previous day, copulation lasted an average of 54 minutes

(20-82, N = 8) . If the male had mated more than 1 day before or

had no known history of mating, copulation lasted an average of 26

min. (18-49, N = 21). Some of the latter males may have mated pre-

viously so that duration without recent mating of the male may be sig-

nificantly less than 26 minutes.

Number of matings. Males may mate at least five times. One native

arachne male mated 5 times in 4 days, and 4 other arachne males mated

at least 3 times each. Immediately after mating, males resume perching

and courting, and two males mated only 11 and 15 minutes after ter-

minating a previous copulation, but as noted above this resulted in a

prolonged mating. Females usually mate only once, very rarely twice.

Dissections of 51 field collected females indicated 14 virgins, 36 mated

once (one spermatophore) , and only one mated twice ( a fresh female

with two abnormally small spermatophores) . The male deposits a

clear, solid plug in the female genital orifice. The plug is visible ex-

ternally, and may act as a mechanical barrier to further mating.

Time and location of mating. Courting and mating occur through-

out the day. Six spontaneous matings were observed in the morning

hours, and 29 experimental copulations were induced between 0732 to

1211 by releasing females in front of perching males. Many courtships

by perching males were observed from 0756 to 1147, and many court-

ships by patrolling males were observed from 1300 to 1530. The mating

Vol. 50, No. 1, January 1974

behavior of males and females remains constant during the day, unlike

mate-locating behavior of males, and all types of rejection behavior

of females and courtship behavior of males were observed throughout

the day. Virgin females were found at all hours, even late in the day,

suggesting that some females do not mate until the second day of adult

life.

In the morning, all observed matings and courtships except one oc-

curred on the ridgetops. A single mated pair on the hillside was found

100 m east of a Cripple Creek ridgetop indicating that courtship prob-

ably occurred on the hillside. Two female P. m. minuta two days old

left a hilltop after release but later returned and mated there. The per-

centage of virgin females on ridgetops at Cripple Creek before 1200 was

35% (9 of 26) but only 8% (2 of 24) 20 m or more from a ridgetop.

These figures suggest that until females fly to the ridgetops they remain

near the sites of emergence, which are mainly on hillsides near Pen-

stemon. After mating, females leave the ridgetops. In the afternoon,

courtships occur mainly on hillsides and flat areas near the morning

perching sites, where virgin and mated females often alight on flowers

to feed.

Both sexes usually emerge from pupae before 1100, males an average

of a day earlier than females. In the afternoon males apparently ac-

tively search for females which emerge that day, then wait on ridges

the next morning for the females which did not mate on the day of

emergence. Some females may immediately fly to ridgetops, but as

shown above females are more receptive a day or more after emergence.

Movements

In 1969 males were marked and recaptured for 10 days along the top

of a ridge at Cripple Creek, mainly in the morning (Fig. 1, Table 2).

Males flew along the entire ridgetop, moving a distance (D) up to

470 m. The moderately high density with resultant numerous chases

between males may have contributed to the high movement on the ridge,

but it is more probable that the movement occurred because the ridge

contained suitable perching spots and flowers along most of it length.

At the Bear Creek study area described below, the hilltops and perch-

ing sites were discontinuous and movement was less. Males are not

territorial, as they wandered over the entire ridge and perched wherever

suitable sites occurred.

In 1970, 20 males were marked and released at Bear Creek, Chaffee

County, Colorado. Six of these were recaptured. Over a two-day pe-

The Pan-Pacific Entomologist

Fig. 1. Maps of Cripple Creek (A) and Bear Creek (B) sites. Contour interval

3 meters (A) and 6 meters (B). Numbered hilltops correspond to foci of activity

discussed in text.

riod only one of these males stayed where marked. Two males moved

considerable distances from a hilltop to a ridge and vice versa. Three

males moved from hilltop 30 (Fig. 1) in the morning to flowers on the

hillside in the afternoon: one of these, observed on hilltop 30 in the

morning of two successive days, appeared on the hillside visiting flowers

on the afternoon of the next day, and returned to hilltop 30 the follow-

ing morning. Another male remained on hilltop 30 in the morning and

the hillside to the west in the afternoon for three days, then was found

on hilltop 33 on the sixth day. Still another male resided on hilltop 30

during the mornings of three days in a 4-day span, and moved to the

hillside west of hilltop 30 on two afternoons during this time. The last

two examples suggest that males return to the same hilltop and hillside

areas repeatedly, but this is probably because hilltop 30 and the hill-

Vol. 50, No. 1, January 1974

Table 2. Movement data for Poladryas minuta arachne at Cripple

Creek in 1969. T = time between first and last capture; t i? dj = time,

and distance respectively, between capture i and capture (i + 1) ; R =

distance between the two farthest capture points ; D = sum of all d’s of

an individual; V = D/T; Vi = dj/h; * = p ^ 0.01 (other correlations

p > 0.05). Significance of correlations was determined by tests of zero

correlation.

Movement

Parameter

Movement

T (days)

3.66

di (meters)

151

ti (days)

1.65

151

V (meters/day)

R (meters)

Vi (meters/day)

151

D (meters)

125

r (ds, ts) = .270*

r (di, age) r

= .003

r (ti, age)

= .184

Partial correlation

(di, ti) , age

constant -

.275*

r (vi, age)

= -0.006

Partial correlation

(di, age), ti

constant rz

-0.049

side to the west are the most favorable (the most prominent hilltop, and

the nearest hillside with many flowers) sites in the immediate vicinity

of the Bear Creek locality.

Effect of copulation and age on movements. Males which copu-

lated on Green Mountain were marked and their movements were stud-

ied. The results suggest that 1) males do not need to have experience

at a site to copulate there, and 2) copulation does not cause the male to

remain at that spot. For example, two arachne males and a minuta male

which were caught elsewhere on Green Mountain or laboratory reared

were released in the morning on a small ridgetop where they immedi-

ately perched and mated within 5 to 15 min. of release. Other arachne

and minuta males perched immediately upon release, showing that, in

the morning at least, males perch and mate in suitable topographic sites

regardless of “territorial” experience. That males do not remain in

areas of previous copulations is shown by their frequent movements.

For example, one male mated twice on hill A on one day, flew elsewhere

the next day, moved to hill B the third day, mated on hill A the sixth

day, then spent the remainder of the sixth day on hill B. Another male

mated on hills, A, A, and B on three successive days. Two males mated

on hills B, A, and A on successive days; one of these courted on hill A

the fourth day. Only one male mated many times on one hill (5 times

in 4 days) .

The Pan-Pacific Entomologist

Table 3. Population parameters of Poladryas minuta arachne males

at Cripple Creek, 1969, estimated from multiple recapture data using the

stochastic model of Jolly (1966). N = total population size; Phi =

probability of survival from each sampling period to the next; B =

number of new animals joining the population during that sampling

period ; SE = standard error.

Day

N ± 1.96 SE

Phi ± 1.96 SE

B ± 1.96 SE

August 12

—

.727

.342

—

■ —

154.7

120.4

.847

.381

118.9

163.9

249.1

158.6

.953

.408

7.8

160.4

245.1

131.7

.637

.314

132.7

119.8

287.1

140.8

.693

.441

146.9

199.9

300.0

228.5

.813

.494

63.4

188.5

273.0

151.1

.923

.481

81.9

136.1

267.4

127.9

—

■ —

Correlations of movement parameters with age for males (Table 2)

show positive correlations of dj with time between recaptures. Time

between captures (tj increases slightly but not significantly with age,

which may indicate less active flight, and hence smaller probability

of capture, with age.

Male Population Parameters

A 10 day mark-recapture study of males at the Cripple Creek site

indicated a population size of approximately 200, with an initial in-

crease then a decrease (Table 3). An average of about 80 individuals

per day joined the population. Average survival rates and expected life-

span were .794 (4.3 days) using method 1, and .817 (5.0 days) using

method 2 of Scott (1973a). Probably a small portion of the population

emigrated from the area, so that the survival rate should be increased

slightly. The potential lifespan is longer than this, of course. The two

longest lived males survived at least nine days and nine others seven to

eight days. Longer times might have been recorded if the mark-recapture

study had been longer.

Feeding and Oviposition

Individuals of both sexes and all ages frequently visit flowers, espe-

cially during the hottest part of the day. Males feed on flowers occasion-

ally during the morning perching period, but more frequently in after-

Vol. 50, No. 1, January 1974

noon. Both sexes visit yellow flowers most frequently, but also visit

flowers of other colors. At Cripple Creek 84 individuals were observed

on Heterotheca villosa (Pursh) Shinners and 16 were observed on other

yellow flowers, while only 19 visits were recorded to white, orange, or

blue flowers. Proportions were similar at other localities.

Oviposition occurs as the female slowly flutters among small Pen-

stemon plants which are usually without inflorescences (individuals have

never been observed feeding on the flowers, which are blue, white, or

red). Eventually she deposits a round cluster averaging 38 eggs (14—

87; N = 43 ) , usually on the lower surface of one of the lower leaves.

Daily eggs counts of 16 laboratory reared P. m. minuta and Fi females

mated on Green Mountain indicate a preoviposition period of 1-4

(usually 1-3) days. Most females deposited several clusters on the same

day or over a period of several days, but several first oviposited on

about the third day, then laid other clusters several days later.

Larval foodplants. Larvae feed on various Penstemon species in

different geographic areas, including P. alhidus Nutt, in Baylor County,

Texas (P. m. minuta ; identified by Roy 0. Kendall), P. dasyphyllis

Gray in Santa Cruz County, Arizona (P. m. nympha) , P. virgatus ari-

zonicus Heller on Mt. Graham, Arizona (P. m. near arachne ; both Lil-

ian Roever, written communication), and P. alpinus Torr. in Larimer

County, Colorado (P. m. arachne ; Sperry & Sperry, 1932). Oviposition

was on P. harbatus torreyi (Benth.) Keck in Boulder County, Colorado

(P. 772. arachne ; Emmel et al. 1971), and on P. secundiflorus Benth.

at Cripple Creek. Many unidentified Penstemon were fed to the larvae

in the laboratory, with the following general results. The herbaceous

or shrubby species with green, mostly hairless, delicate foliage were

moderately or very acceptable to the larvae. A cultivated variety (“pin-

nifolius”) with small needle-like leaves was moderately acceptable. The

herbaceous species with thick leaves covered with a whitish or bluish

bloom were completely unacceptable. P. m. minuta larvae readily ate

the leaves of P. albidus , and ate lesser amounts of other Penstemon

including P. barbatus. The Fi, backcross, and P. m. arachne larvae fed

readily on most species tested (with the exception of the glaucous spe-

cies) , including P. barbatus. The palatability of these Penstemon species

is similar for grazing animals (Forest Service, 1937), suggesting that

grazing may have caused the extinction of P. m. minuta over most of

its former range in Texas (Kendall, 1971).

First and second instars are gregarious feeders, but later instars be-

come solitary due to mortality and dispersal. Developmental period

The Pan-Pacific Entomologist

from egg to adult ranges from 46 to 57 days indoors at about 20° C

(egg stage 8-10 days, pupal stage 11-14 days). There are 4-5 broods

in Texas and 2—3 in Colorado, but emergence is asynchronous; distinct

broods were vaguely defined at Cripple Creek and almost constant num-

bers were observed on Green Mountain from early June to early Sep-

tember. Diapause may occur in half-grown larvae.

Hybridization Experiments

Hybridization of these subspecies is significant not only in providing

much of the behavioral data presented, but also because they have been

considered to be distinct species (Bauer, 1961) . P. m. minuta was reared

from larvae collected in Baylor County, Texas, and females were re-

leased in front of wild perching males of P. m. arachne on Green Moun-

tain and south of Glenwood Springs, Garfield County, Colorado. Some

resultant Fi females were later mated to wild arachne males on Green

Mountain and the resulting backcross larvae raised to adults. Sex ratio

of Fi and backcross adults was approximately 1:1. Among 11 minuta

2 ( X arachne $ ) crosses, two produced no eggs, three infertile eggs,

and six fertile eggs which produced adults. Among 14 Ft $ (X arachne

$ ) crosses, six females failed to oviposit, while eight laid many com-

pletely fertile eggs which produced many adults. Among 15 controls

(arachne $ X arachne $ ) nine females failed to oviposit, eggs of two

females were infertile, offspring of two females died as first or second

instar larve (apparently due to disease), and two females produced

many viable offspring. P. m. minuta and P. m. arachne are morpho-

logically very similar. Adults differ slightly in details of wing, antennal

and abdominal color pattern, and larvae differ in the ground color be-

tween the scoli of the dorsum of late instars (red in minuta , largely

white in arachne ). Mr. Jack Harry, Denver, Colorado, collected a large

series near Guadeloupe Peak, Culberson County, Texas, which although

more similar to minuta than arachne , is clearly intermediate in several

wing pattern characteristics. In addition the two subspecies are very

similar in foodplant preferences and in all aspects of courtship behavior,

as shown previously. These similarities, together with the high degree

of genetic compatibility, clearly demonstrate the conspecificity of these

two taxa.

Discussion

The most unusual feature of behavior in P. minuta is the alteration of

mate-locating behavior during the day. This alteration appears to be

Vol. 50, No. 1, January 1974

well adapted to the search for flowers and virgin females. In the morn-

ing, before the females emerging that day are able to fly to ridgetops,

males wait for females, which are more active and receptive after their

first day of adult life, to come to ridgetops. In afternoon, males search

throughout the habitat near flowers, often on hillsides and flats, for

virgin females which often feed on flowers at that time. In the morning,

temperatures are generally lower than in afternoon; males bask at tem-

peratures too low for prolonged flight. In afternoon higher temperature

results in more flower feeding, and enables continuous searching flight.

Summary

Males perch on hilltops and ridgetops from 0700 to about 1215 to

locate females, and dart out at passing fluttering objects. Females fly

to ridgetops to mate. In the afternoon, males patrol on hillsides and

flats near flowers in search of females. Courtship is quite variable, as

several male and female behavioral elements are present or absent de-

pending partly on the receptivity of the female, which in turn depends

partly on age. Unreceptive virgin females have a variety of means of

discouraging males, but unreceptive mated females use a stereotyped

vertical flight which resembles the encounter between adjacent perching

males, and visual stimuli are of general importance both before and

during courtship. Males are not territorial. Movement is most pro-

nounced along continuous ridges, decreasing in those habitats with

isolated hilltops. Both sexes feed predominantly at yellow flowers. Fe-

males deposit clusters averaging 38 eggs on various species of Pen-

stemon , usually beginning 2-4 days after emergence. P. m. minuta from

Texas was hybridized and backcrossed with P. m. arachne from Colo-

rado by use of natural courtships. Lack of barriers to interbreeding,

and the similarity of adult behavior, larval and adult morphology, and

foodplants, confirm the conspecificity of these two allopatric subspecies.

Literature Cited

Bauer, D. L. 1961. p. 131 In-. Ehrlich, P. R., & A. H. Ehrlich. How to know

the butterflies. Wm. C. Brown, Dubuque, la.

Ehrlich, P. and S. Davidson. 1960. Techniques for capture-recapture studies of

Lepidoptera populations. J. Lepid. Soc., 14: 227-230.

Emmel, J., 0. Shields, and D. Breedlove. 1971. Larval foodplant records for

North American Rhopalocera. Part 2. J. Res. Lepid., 9: 233-242.

Forest Service, U. S. 1937. Range Plant Handbook. U. S. Government Printing

Office, Washington.

Jae, R. 1972. Natural interbreeding of close nymphalid groups. J. Lepid. Soc.

26: 28.

The Pan-Pacific Entomologist

Jolly, G. 1966. Explicit estimates from capture-recapture data with both death

and immigration — stochastic model. Biometrika, 52: 225-247.

Kendall, R. O. 1971. Lepidoptera in the unpublished field notes of Howard

George Lacey, naturalist (1856-1929). J. Lepid. Soc., 25: 29-44.

Scott, J. A. 1973a. Population biology and adult behavior of Lycaena arota

(Lyacaenidae) . J. Lepid. Soc., in press.

Scott, J. A. 1973b. Mate-locating behavior of butterflies. Amer. Midi. Nat.,

in press.

Scott, J. A. 1973c. Mating of butterflies. J. Res. Lepid., 11: 99-127.

Sperry, G. H., and J. Sperry. 1932. Notes on the larva of Melitaea pola Bdv.

Bull. So. Calif. Acad. Sci., 31: 8.

SCIENTIFIC NOTE

Notes on Hesperus arizonicus with description of the female (Coleoptera:

Staphylinidae ). — Hesperus arizonicus was described by Moore in 1958 (Trans.

San Diego Soc. Natur. Hist. 12: 311) on the basis of a single male specimen from

Patagonia, Santa Cruz County, Arizona. A single female specimen in the collection

of the University of California at Riverside is from Cave Creek Ranch, elevation

5000', Cochise County, Arizona, taken on August 1, 1965 at ultraviolet light by

G. R. Ballmer. It differs from the male in numerous respects.

Description of the female. Color largely ferruginous; third through tenth

antennomeres dusky; elytra with yellow band across basal third hounded basally

and apically by a narrow piceous band, apical third ferruginous; fifth abdominal

segment piceous at apical two-thirds; sixth abdominal segment yellow. Head

wider than long, slightly wider than pronotum, dorsal surface feebly microreticulate,

with coarse punctures generally separated by less than their diameters except for

small elongate impunctate central area anteriorly, and area of nearly coalescing

punctures behind eyes. Pronotum subquadrate, slightly longer than wide, slightly

narrowed behind, sides somewhat sinuate before posterior angles; surface feebly

microreticulate, not impresesd, as coarsely punctured as head but more sparsely

so, with punctures generally separated by their width, and with a central impunctate

strip. Elytra quadrate, wider and longer than pronotum; humeri broadly rounded,

sides straight and hardly divergent to narrowly rounded exterior apical angles;

surface finely, densely, and rather roughly punctured. Abdomen narrower than

elytra, tapering slightly to apex; dorsal and ventral surfaces with fine, longitudinal

microstrigulation, about as finely punctured as elytra but more sparsely so and

not as roughly ; last sternite evenly arcuate at apex.

The most important structural difference between this female and the holotype

male is in the unmodified apical margin of the sixth sternite, a condition typical

of many Staphylinidae. There is also considerable difference in color, some in

ground sculpture and in the fact that the surface of the pronotum is not impressed.

These latter differences are probably within the range of individual variation for

the species and not of sexual significance. — Ian Moore, University of California,

Riverside, 92502.

Vol. 50, No. 1, January 1974

Vaejovis spicatus: A New Scorpion from California

(Scorpionida: Vaejovidae)

Richard M. Haradon

San Mateo, California

Described below is a new vaejovid scorpion species, discovered in tbe

Little San Bernardino Mountains of southern California. A remarkably

developed subaculear proturberance on the telson of this species suggests

the name.

Vaejovis spicatus Haradon, new species

(Figs. 1-7)

Diagnosis. — Males unknown. Females of V. spicatus are distinguishable from

all other known species of Vaejovis by the presence of an elongate subaculear

“tooth” (Figs. 4, 5). Females somewhat similar to those of Vaejovis joshuaensis

Soleglad (1972, p. 190), also from southern California, but differ as follows: (1)

fixed pedipalp finger about equal to or slightly shorter (not distinctly longer) than

manus; (2) six and seven interior lateral granules on fixed and movable pedipalp

fingers respectively (do not vary on each finger from four to six) ; (3) meta-

somal segment V width greater than (not about equal to) half its length, and

greater (not less) than width of segment I; (4) bristles along dorsal keels of

metasomal segments I-IV, 0, 1, 1, 2 (not 0, 0, 1, 1) ; (5) total adult length about

17 mm (not 24 mm) .

Description of holotype (adult female). — Size. — Measurements given in

Table 1.

Coloration. — Ground color brownish-yellow, fingers slightly darker; no contrast-

ing markings. Granules along carinae usually reddish-brown. Tip of aculeus red.

Carapace. — Anterior margin concave, with slight median notch. Entire surface

finely granular. Median furrows narrow, shallow; posterior laterals broad, very

shallow; all other furrows and associated keels obsolete. At least two pairs of

lateral eyes, third indefinite. Median ocular tubercle set in shallow depression,

forward of center; posterior carapace length/carapace length = 0.65.

Chelicerae. — Superior margin of fixed digit with basal bicusp, single adjacent

tooth, and terminating in single tine; inferior margin smooth. Movable digit bi-

furcate distally, superior tine about % length of inferior; superior margin with

two small subapical teeth, adjacent larger tooth, and fourth small, basal; serrula

along smooth inferior margin, not extending to apex. Long white hairs along in-

ferior and interior basal surfaces of fixed digit. Single dorsal bristle just posterior

to digital commissure.

Pedipalps. — Sparsely hirsute. Interior keels of humerus and brachium distinct

and granular; exterior dorsal keels weak, lightly granular. Intercarinal areas uni-

formly finely granular. Chelae relatively slender; manus contour somewhat

rounded; all keels weakly developed; digital smooth, inner secondary very lightly

granular, interior marginal vestigial. Movable digit distinctly longer than manus;

interior margins with closely set, pointed denticles in seven (movable digit) and

Tile Pan-Pacific Entomologist 50: 23-27. January 1974

The Pan-Pacific Entomologist

Figs. 1-5. Vaejovis spicatus Haradon, new species. Holotype female. 1. Right

chela, trichobothria ventral-exterior surfaces. 2. Right brachium, trichobothria

exterior surface. 3. Right chela, trichobothria dorsal surface. 4. Telson, lateral

view. 5. Telson, ventral view.

six (fixed digit) rows, each row marked distally by enlarged denticle and adjacent

interior lateral granule; slight even space between closed fingers. Trichobothria:

humerus, one each proximally on interior, dorsal and exterior surfaces; brachium,

two ventral, one interior, two dorsal, 14 exterior (Fig. 2) ; chela, two on interior

base of fixed digit, five on exterior surface of manus (Fig. 1), 19 on fixed digit

and manus viewed doreally (Fig. 3) .

W diking legs . — Generally smooth ; all keels very weak or obsolete. Conspicuous

bristles sparse on all segments. Single row of short white bristles ventrally in line

with unguicular spine along entire length of distal tarsomere.

Genital plate . — Completely fused medially.

Pectines . — Moderately hirsute; extending to trochanter of fourth walking leg;

11 teeth overlapping slightly in series; six oval to subcircular middle lamellae;

fulcrae subcircular.

Mesosoma . — Terga extremely finely granular; vestigial median keel on terga

four to seven ; two pairs of weak lateral keels on seven. Sterna smooth to extremely

finely granular; lateral keels on last sternum weak, granular; stigmata short,

slit-like.

Metasoma . — Segments I— II wider than long, III about as wide as long. Keels:

dorsals serrate, terminating in enlarged conical spine; superior laterals I-IV ser-

rate, with enlarged spine on I— III and fin-like process posteriorly on IV, low

Vol. 50, No. 1, January 1974

Figs. 6-7. Vaejovis spicatus Haradon, new species. Holotype female. 6. Dor-

sal view. 7. Ventral view.

rounded granules on V ; laterals granular in posterior % on I— II, posterior % on

III, absent on IV, weak with scattered rounded granules in anterior % on V. In-

ferior laterals similar to corresponding inferior medians; I weakly crenulate; II

crenulate; III crenulate to finely serrate; IV-V serrate. Inferior median keels

I-IV set with 3, 3, 3, 3 bristles; dorsal keels with 0, 1, 1, 2 bristles.

Telson . — Vesicle laterally swollen; surface extremely finely granular; 16 long

slender hairs on ventral and lateral surfaces. Aculeus short, sharply curved, with

elongate subaculear protuberance (Figs. 4, 5) .

Variation . — The only other adult specimen, also a female, does not differ sig-

nificantly from the description of the holotype. Three minute immature specimens,

of undetermined instar, were characterized most obviously by differences in ana-

tomical proportions, lack of pigmentation, and incomplete carinal development;

the subaculear tooth was well developed.

Material. — Holotype from Berdoo Canyon, 6.9 miles NE of junction with Dilllon

Road, Little San Bernadino Mountains, Riverside County, California (elevation

3600 feet), 31 March 1972, R. M. Haradon and J. L. Marks. Four paratypes from

Berdoo Canyon, 3.2 to 5 miles NE of Dillon Road (elevation 2000-2500 feet). 18

The Pan-Pacific Entomologist

Table 1 . Measurements (in millimeters) of adult Vaejovis spicatus

Haradon, new species.

Holotype

(female)

Paratype

(female)

Total length

17.30

16.10

Carapace length

2.35

2.25

Anterior width

1.15

1.10

Width at/of ocular tubercle

1.65/0.25

1.60/0.25

Posterior width

2.00

1.85

Mesosoma length (sum)

6.10

5.65

Metasoma length (sum)

6.60

6.05

segment I length/width

0.90/1.15

0.80/1.10

segment II length/ width

1.05/1.15

0.95/1.10

segment III length/width

1.15/1.15

1.05/1.10

segment IV length/ width

1.35/1.25

1.25/1.20

segment V length/width

2.15/1.30

2.00/1.20

Telson length

2.25

2.15

Vesicle length/width

1.65/1.05

1.60/1.15

Vesicle depth

0.80

Aculeus length

0.60

0.55

Pedipalp humerus length/width

2.00/0.60

1.90/0.55

brachium length/width

2.30/0.70

2.15/0.65

manus length/width

1.85/1.00

1.80/1.00

chelal/fixed digit lengths

3.65/1.80

3.50/1.70

movable digit length

2.20

2.10

Pectinal teeth (left/right)

11/11

10/11

Middle lamellae (left/right)

6/6

Margin lengths, anterior/dentate

1.35/1.10

1.25/1.00

March, 9 April and 5 May 1972, and 24 February 1973, R. M. Haradon and J. L.

Marks.

The holotype has been deposited at the California Academy of Sciences, San

Francisco.

Remarks. — The type locality and only known habitat is a narrow,

sparsely vegetated, desert canyon. All specimens of V. spicatus were

found on rocky canyon walls, by ultraviolet light at night. Also at the

type locality occur at least six other scorpion species: Hadrurus ari-

zonensis Ewing, Paruroctonus vachoni Stahnke, Vaejovis confusus

Stahnke, Vaejovis deserticola Williams, Vaejovis joshuaensis Soleglad,

and Vejovis hirsuticauda Banks.

Acknowledgments. — Stanley C. Williams, California State Univer-

Vol. 50, No. 1, January 1974

sity, San Francisco, kindly reviewed the original manuscript. Photo-

graphic credit is due R. H. Howard, Los Angeles. Joseph L. Marks,

Santa Monica, generously contributed much time and effort in the field,

and is properly credited as the co-discoverer of V. spicatus.

Literature Cited

Soleglad, M. E. 1972. Two new scorpions of the wupatkiensis group of the

genus Vejovis. Wasmann J. Biol., 30: 179-195.

BOOK NOTICE

Eastern Forest Insects. Whiteford L. Baker. U.S. Department of Agriculture,

Forest Service, Miscellaneous Publications no. 1174, 642 pp. 1972. $5.00.

This completely revised version of an earlier edition (For. Ser. Misc. Publ. 657)

contains concise but informative accounts of the life histories of the important in-

sects destructive to forest trees in eastern North America. The text has been ex-

tensively rewritten and many new photographs included. The cited literature in-

cludes over 800 entries, nearly all more recent than 1940, and both the insects and

their host plants are indexed, which should make this volume an extremely valuable

successor to the earlier work. — Editor.

RECENT LITERATURE

Ants of Deep Canyon. George C. Wheeler and Jeanette Wheeler. Philip L. Boyd

Deep Canyon Desert Research Center, University of California, Riverside. 162

pp. 1973. $3.95.

In this paperback volume the Wheelers have produced a highly interesting sum-

mary of the biology of desert ants as well as a useful taxonomic treatment of the

genera which occur in the arid regions of North America. Discussions of the eco-

logical characteristics of the North American hot deserts and some of the morpho-

logical, physiological and behavioral adaptations of desert ants are followed by de-

tailed taxonomic and ecological discussions of the species occurring in Deep Canyon.

The text is profusely illustrated with photographs of the ants, their nests, and typ-

ical habitats, and provided with a glossary of technical terms. — Editor.

The Pan-Pacific Entomologist

The Maritime Earwig in California

(Dermaptera: Carcinophoridae)

Robert L. Langston

31 Windsor Ave., Kensington, Calif. 94708

During an extensive survey of earwigs in California, particular em-

phasis was focused on the maritime or seaside earwig, Anisolabis mari-

tima (Gene). My special interest was due to the lack of material in

most California museums, and to the rather narrow ecological habitat

of this species.

Based on the classification of Popham (1965), the placement of An-

isolabis is in the subfamily Carcinophorinae under the Labiodea.

Within California, this is the only species of the genus found under

natural conditions. It is morphologically distinct, and is rarely con-

fused with any other earwig occurring in the state.

A condensed synonymy is given below to indicate only the original

references for each generic change, combined with the first records for

the United States and also California.

Anisolabis maritima (Gene)

Forficula maritima Gene, 1832, Sagg. Monogr. Forfic. Indig., p. 9. [Nice, France;

Genoa and Tuscany, Italy; along Mediterranean.]

Forficula ( Labidura ) maritima, Fischer, 1853, Orth. Eur., p. 68, pi. 6, figs. 4, 4a-d.

F orficesila maritima, Serville, 1853, Ins. Orth., p. 27.

Anisolabis maritima, Fieber, 1853, Lotos, III, p. 257. [South Carolina.] (Generic

assignment and first United States record.)

Brachylabis maritima, Dohrn, 1864, Stettin. Ent. Zeit. 25, p. 293.

Anisolabis maritima, Essig, 1922, Pomona College Jour. Ent. & Zool., p. 75. [La-

guna Beach, California,] (First California record.)

Diagnostic Features and Life History

The adult maritime or seaside earwig measures 20-32 mm including

forceps. It is wingless, with the body shiny black, or sometimes very

dark brown. The uniformly dark colored antennae vary from 20 to 24

segments in the adults, with a lesser number in the juveniles. (One to

several lighter segments are diagnostic for some other species of ear-

wigs.) The legs are pale yellow (dark banding on the legs is character-

istic of a smaller, closely related species commonly found in California) .

The forceps of the male are curved or sickle-shaped, and asymmetrical,

the right being shorter. The forceps of the female are basally parallel,

becoming curved slightly inward near the tips.

The Pan-Pacific Entomologist 50: 28-34. January 1974

Vol. 50, No. 1, January 1974

The immatures have the body shape, color and straight forceps of the

mature females, but juveniles can be differentiated by the number of

segments in the antennae; the number increases with each successive

stage. Preserved tenerals may remain brown, as in some of the faded

specimens in museum collections.

Maritime earwigs appear to be primarily predaceous. In the labora-

tory they have been observed by Fulton (1924) to catch and devour

crickets, sand-fleas and smaller earwigs, using their forceps. Foraging

occurs at night. In the daytime I found these earwigs under rocks, logs,

driftwood and various other objects washed up along shores where there

is salt or brackish water, but they quickly crawl into cracks and crevices

when exposed. Because of the flexible abdomen and the strong pinch-

ing ability, they are most easily collected with forceps.

Adults and juveniles have been collected during all seasons of the

year. They occur in definite colonies, and if a single individual is

found, usually many (up to several dozen) can be collected in the im-

mediate vicinity. In my experience, there was a predominance of fe-

males at all collecting sites.

Based upon seven years (1964-69, 73) of collecting in California,

adult females and the larger juveniles (third, fourth and fifth instars)

were found in the winter and early spring months. Depending upon the

climate, availability of food, and other factors, the instars tend to over-

lap in size, and even the adults have considerable size variability. Some

of the dwarf adults may be just slightly over half the size of the ma-

jority of individuals.

In late spring, summer and autumn, males were found in the colonies

along with females and all stages of juveniles. Eggs were found in deep

cracks or burrows from early summer to autumn, but are probably pres-

ent at other times. The female guards her egg clutch, which is probably

deposited over a period of several days. In one instance, 84 eggs were

found tended by a single female. In another case, 88 eggs guarded by

two females were unearthed from a chamber between rocks and dirt.

Females maintain the eggs in neat piles, turn them over, and keep them

free of dirt. The eggs are glistening white, nearly spherical and slightly

over 1 mm in diameter when deposited. As the embryos develop, the

eggs elongate to nearly 2 mm and darken.

In California, Anisolabis maritima is found only under very restricted

ecological conditions. Most individuals have been collected near the

high tide level at San Francisco Bay, San Pablo Bay and Carquinez

Strait (Langston, 1967). Below this level the conditions appear to be

The Pan-Pacific Entomologist

too wet, with the water often seeping into the depressions beneath rocks

or logs. In this zone the predominant animal life consists of Isopoda

(kelp sowbugs), and other Crustacea. Above the high tide level the

substrate becomes increasingly drier. In this zone the predominant

animal life consists of Isopoda (dooryard sowbugs, common pillbugs),

with the commoner insects including silverfish, crickets, European ear-

wigs, carabid and staphylinid beetles, and ants. Many of the areas where

this earwig was found were very rocky with considerable drift and de-

bris, hut very little noticeable plant life. It is probable that A. maritima

moves up and down from the high tide level to feed on most of the ani-

mal forms mentioned above, except probably the larger Crustacea.

Caudel (1913) stated that A. maritima is found only along the sea-

shore. In California it seems to prefer bays and inlets (Langston, 1967) .

Although generally distributed along the Gulf and Atlantic coasts, the

established colonies in California are mostly restricted (see Fig. 1), with

few collections from along the open ocean coasts. Absence of many

records on the immediate coast is not due to poor collecting, as two

other species of earwigs, Euborellia annulipes (Lucas) and Forficula

auricularia Linnaeus, have been taken abundantly. There are many

records of both species either on or just above the seashore in north-

ern, central and southern California.

The first published report on the occurrence of A. maritima in Cal-

ifornia was by Essig (1922) , who found it at Laguna Beach and vicinity

during June and July, 1921. In 1944, a single male was collected at

Costa Mesa also in Orange County. It has been taken in quarantine by

the California Department of Agriculture at six localities in southern

California.

Geographical Distribution

Anisolabis maritima is worldwide in distribution. It has been re-

corded from most of the continents and major islands, with the excep-

tion of the Arctic and Antarctic regions. Specifically, it has been re-

ported from France, Italy, along the Mediterranean (Hebard, 1917) ;

Canary Islands, Formosa, Japan, New Zealand, Puerto Rico, Santo Do-

mingo (Burr, 1910); and Venezuela (Rehn, 1949). Published North

American records include British Columbia, Canada (Hebard, 1933;

Heifer, 1963) ; and the Gulf and Atlantic coasts of the United States as

far north as Maine (Hebard, 1917) .

California distribution : — The detailed California records include the

term “juv.” for immature specimens, which are similar to the females

until the penultimate stage. In this stage and the adults, the forceps dif-

Vol. 50, No. 1 , January 1974

Fig. 1 . California distribution of Anisolabis maritima. Circles denote established

populations. Triangles denote quarantine records.

fer markedly between the sexes. Therefore, the specimens are not sexed

in the detailed collection records unless close to, or in the adult stage.

Established records of specimens are believed to represent naturalized

populations. These include localities where large numbers were taken

The Pan-Pacific Entomologist

or small numbers were accumulated over a period of years. A single

record may be considered established if it is within the over-all distri-

bution where there are adjacent or contiguous records. Quarantine rec-

ords indicate specimens taken at border or ports-of-entry stations by

the California Department of Agriculture. Quarantine also includes the

within-state or county records where specimens were intercepted. These

border and within-state records usually show the point-of-origin, and/

or the host material.

The following initials refer to the institutions where specimens are

on deposit: BVC = Bureau of Vector Control, California Public Health

Service, Berkeley; CAS = California Academy of Sciences, San Fran-

cisco; CDA = California Department of Agriculture, Sacramento; CIS

= California Insect Survey, University of California, Berkeley; CSH =

California State University at Hayward; LACM = Los Angeles County

Museum of Natural History, Los Angeles; SFS = San Francisco State

University, San Francisco; UCD = University of California, Davis;

and UCR = University of California, Riverside.

Alameda County: Alameda, 1 juv., No date or collector (CDA) ; Albany, 2

$ 2, X-17-1965 (R. L. Langston— CIS) ; Berkeley, 6 $ $ , 24 2 2, XII-16-1949,

40 $ $ . XII-24-1949, shore of S. F. Bay (J. W. MacSwain — CIS) ; Emeryville,

2 $ $ , 20 $ $ , 20 juv., X-17-1965, 4 $ 2 , 4 juv., 1-15-1966, mud flats (R. L. L.—

CAS, CDA, CIS, LACM, UCD, UCR) ; Oakland, VII-17-1957 (N. L. Jones— CDA).

Contra Costa County: Point Molate, Richmond, 1 $ , 4 2 2, III-23-1962 (A.

G. Raske — CIS) ,12 $ 2 , 19 juv., 11-13-1966, 24 $ $ , 26 2 2, V-19-1966, shore

of S. F. Bay (R. L. Langston— CAS, CDA, CIS, CSH, LACM, SFS, UCD, UCR),

23 S $ , 41 2 2 , 20 juv., III-26-1966 (R. L. L. & P. A. Opler — CAS, CIS) ; Point

Richmond, 4 $ $ , 12 2 2,8 juv., V-19-1966, 2 $ $, 10 2 2,6 juv., V-6-1967,

shore of S. F. Bay (R. L. L. — CIS, LACM) ; Pinole Point, 3 $ $ , 12 2 2,6 juv.,

IX-29-1973 (R. L. L. — BVC, CAS) ; Point San Pablo, Richmond, 2 $ $ , 8 2 2,

IX-16-1967, 11 S $, 8 2 2, HI-9-1968, 15 $ $ , 25 2 2 , 20 juv., IX-21-1973, shore

of San Pablo Bay (R. L. L.— BVC, CAS, LACM) ; Port Costa, 1 mi. W., 2 <$ £,

15 2 2 , V-30-1964, 7 2 2, VI-27-1964, 1 $ , 1-17-1965, 8 2 2,8 juv., 11-27-1966,

shore of Carquinez Strait (R. L. L. — CAS, CDA, CIS, LACM, UCD, UCR) . Los

Angeles County: Los Angeles, IV-10-1940, Quarantine from San Francisco on

dahlias (CDA) ; San Gabriel, III-3-1941, Taken in quarantine on chrysanthemum

(CDA). Marin County: Belvedere, 1 2, V-13-1947 (E. Tams — CDA); China

Camp, 2 2 2, XI-7-1965, shore of San Pablo Bay (R. L. Langston — CIS) ; Near

Peacock Gap, San Rafael, 3 3^,5 2 2,3 juv., XI-7-1965, 6 $ S , 20 2 2 , 84

eggs, VI-2-1966 (R. L. L.— CAS, CDA, CIS, LACM, SFS, UCR) ; Sausalito, 1 $,

14 2 2 , 10 juv., XI-7-1965, shore of S. F. Bay (R. L. L. — CAS, CIS, LACM, UCR).

Orange County: Costa Mesa, 1 $, III-6-1944, ex firewood (R. G. Bumgardner

— CDA) ; Laguna Beach and vicinity, VI & VII-1921 (Essig, 1922 — first California

record). San Diego County: Chula Vista, 1-18-1933, ex lawn (CDA) ; Del Mar,

III-9-1932, Quarantine on Araucaria sp. (J. Adams — CDA) ; El Cajon, IV-10-1940,

Quarantine from San Francisco on dahlia tubers (CDA) ; Escondido, VI-29-1934,

Vol. 50, No. 1, January 1974

Quarantine from OREGON (M. England — CDA) . San Francisco County:

“Frisco Bay, Cal.” 1 $, 4 9 9, XII-10-1949, under rock (“W. V. G.”— CIS) ; San

Francisco, 1 $, IX-1935 (E. S. Ross — CAS). Solano County: Benecia, 5 $ $,

12 $ $ , VI-2-1966 (R. L. Langston — CAS, CIS) ; Benecia, 1 mi. W., 7 $ $ , 13

9 9,8 juv., 88 eggs, VI-2-1966, 13 $ $ , 32 9 9 , 27 juv., X-16-1966, shore of Car-

quinez Strait (R. L. L. — CAS, CIS, CSH, SFS) ; Glen Cove, 1 $ , 1 9, VI-2-1966,

1 $, 4 9 9, VI-11-1967, 1 $, 4 9 9, VI-15-1969, shore of Carquinez Strait (R.

L. L. — BVC, CIS, LACM) .

Limited searches with negative results were made on parts of the im-

mediate coast within the boundaries of the inset in Fig. 1. It is possible

that severe wave action excludes the earwig from the Pacific Ocean sea-

shore, in the western part of this area. It appears to be absent along

the northern periphery of San Pablo Bay (Black Point, Vallejo, mouths

of the Napa and Petaluma Rivers), possibly because the water is too

fresh from the river outflows. It may also be absent from the west delta

area east of Benecia or Port Costa (Martinez, Port Chicago, Antioch,

Bethel Island, Brannan Island, Rio Vista) because the water may be

too fresh, although seasonal increases in salinity occur depending on

the tides and time of year. No examples were taken south of Alameda

or San Francisco. Perhaps the water is too salty or polluted. However,

two other earwigs ( Euborellia annulipes and Forficula auricularia ) oc-

cur abundantly around the South Bay.

Acknowledgments

The help of Mr. George Buxton, California Department of Agriculture,

Sacramento is greatly appreciated. Complete freedom in the use of his

files, examination of specimens, and prompt notification of new records

as they appeared in quarantine were especially helpful. Thanks are ex-

tended to Dr. J. A. Powell of the California Insect Survey, Berkeley, for

revisions and suggestions in relation to a more complete manuscript

on the Dermaptera, from which much of the data for this paper was ex-

tracted. I gratefully acknowledge the cooperation of Dr. P. H. Arnaud,

Jr., for the study of the collection at the California Academy of Sciences,

San Francisco, which is particularly rich in specimens from many parts

of the world.

Literature Cited

Burr, M. 1910. The Dermaptera (earwigs) of the United States National Mu-

seum. Proc. U.S. Nat. Mus., 38: 443^167.

Caudell, A. N. 1913. Notes on Nearctic orthopterous insects. I. Nonsaltatorial

forms. Proc. U.S. Nat. Mus., 44: 595-614.

The Pan-Pacific Entomologist

Essig, E. 0. 1922. Insect notes from Laguna Beach, California. Pomona College

J. of Entomol. & Zool., 14: 75-78.

Fulton, B. B. 1924. Some habits of earwigs. Ann. Entomol. Soc. Amer., 17 :

357-367.

Hebard, M. 1917. Notes on the earwigs (Dermaptera) of North America, north

of the Mexican boundary. Entomol. News, 28: 311-323.

Hebard, M. 1933. Dermaptera in collection of the California Academy of Sci-

ence. Pan-Pac. Entomol., 9: 140-144.

Helfer, J. R. 1963. How to know the grasshoppers, cockroaches and their allies.

Pictured-key Nature Series. Wm. C. Brown Co., Dubuque, Iowa, v -p

354 pp.

Langston, R. L. 1967. The maritime earwig, Anisolabis maritima (Gene). Pan-

Pac. Entomol., 43: 88.

Popham, E. J. 1965. A key to dermapteran subfamilies. The Entomologist,

June, 1965: 126-136.

Rehn, J. W. H. 1949. Catalogue of the Dermaptera of Venezuela. Bol. Entomol.

Venezolana, 8: 1-9.

RECENT LITERATURE

The Butterflies of Southern California. Thomas C. Emmel and John F.

Emmel. Natural History Museum of Los Angeles, Science Series, 26, 148 pp.

1973. $4.00.

Amid the plethora of regional natural history manuals published recently this

volume is distinguished by its uniform professional quality, combined with a pop-

ular, easily read style. The introductory discussion includes a concise review of

the seasonal and geographic distribution of southern California butterflies in terms

of Merriam life zones. Brief but extremely interesting historical sketches of the

more important lepidopterists of the area are followed by informative accounts

of the species, including summaries of distributions and larval food plants and

short descriptions of immature stages. The color plates of adults are excellently

reproduced, and many larvae and pupae are illustrated in text figures. The host

plant index and a pertinent list of references are useful features not always found

in such faunal treatments.- — Editor.

Vol. 50, No. 1, January 1974

Notes and Descriptions of Some Lycid-like

Neotropical Lepturine Cerambycidae

(Coleoptera)

John A. Chemsak and E. G. Linsley

University of California, Berkeley, California

The genus Megachoriolaus was proposed by Linsley (1970) for a

Mexican species with lycid-like coloration. At the same time a number

of other species were transferred to this genus from Euryptera, all of

which are probably also mimetic. In 1971 Linsley and Chemsak added

two of Bates’ species and described one additional new species. Subse-

quent collections have yielded a number of additional new species which

are described below because of their possible significance in Neotropical

mimetic complexes.

During the study of the Euryptera-Megachoriolaus complex (Linsley

and Chemsak, 1971), a paper by Fuchs (1956) was inadvertently over-

looked. In this publication, three new species of Euryptera were de-

scribed and other species previously included in the genus were char-

acterized. Judging by the original description, E. unilaticollis Fuchs is

probably assignable to Euryptera as defined by Linsley and Chemsak;

E. rotundipennis Fuchs to Choriolaus Bates; and E. brasiliensis to Mi-

miptera Linsley.

The National Science Foundation through grant #GB-4944X is grate-

fully acknowledged.

Genus Megachoriolaus Linsley

Megachoriolaus Linsley, 1970, Pan-Pac. Entomol., 46:128; Linsley and Chemsak,

1971, Arq. Zool., 21:3.

Type species: M. chemsaki Linsley, by original designation.

All of the known species of this genus fall into one of two color types,

yellow and black or red and black. Since few of the mimicry models

are known, the significance of these patterns remains to be determined.

The yellow and black species are: chemsaki Linsley; breviceps Linsley;

spiniferus Linsley; patricia (Bates) ; filicornis Linsley and Chemsak;

and unicolor (Bates). The red and black species include: flammatus

Linsley; ignitus (Schaeffer) ; sabinoensis (Knull) ; imitatrix Linsley;

cruentus (Martin) ; texanus (Knull) ; and the four new species described

below.

The Pan-Pacific Entomologist 50: 35-43. January 1974

The Pan-Pacific Entomologist

Megachoriolaus lineaticollis, new species

Female: Form moderate sized, elytra slightly expanded posteriorly; elytra dark

Nopal red (Ridgeway, 1912), pronotum red orange except for broad, median,

longitudinal black band, vertex with a black band extending onto neck, appendages

black, prosternum black at sides, mesosternum orangish on coxae and mesosternal

process. Head with vertex impressed behind antennal tubercles, finely, confluently

punctate, pubescence fine, dense, recumbent; front moderately elongate, ratio of

distance between apices of antennal tubercles and line midway between them and

apex of labrum, 1.0:1. 8; antennal tubercles shallow, very finely punctate to large

glabrous median triangle, pubescence fine, rather dense, depressed; clypeus with

a few irregular punctures at base; labrum with a few setiferous punctures; genae

finely punctate; antennae extending to about middle of elytra, segments to base

of fifth shining, clothed with coarse black hairs, outer segments opaque, poriferous

areas absent, segments from fifth thickened, third shorter than first, fourth shorter

than third, fifth longer than third but shorter than first. Pronotum densely, rather

finely punctate except for vague linear area at center behind middle; pubescence

dense, reddish, appressed; sides sinuate, posterior angles acutely produced over

humeri; prosternum subglabrous, shallowly impressed before coxae; mesosternum

with intercoxal process extending to level of coxae, abruptly declivous in front;

metasternum very finely, shallowly punctate, finely pubescent. Elytra about 2.2 as

long as width across humeri; punctures very fine, dense, separated by about one

puncture diameter; pubescence dense, long, reddish, subdepressed; apices shallowly

emarginate truncate, outer angles strongly dentiform. Legs with posterior tarsi

shorter than tibiae, first segment as long as remainder of tarsus; femora slender,

densely, shallowly punctate. Abdomen very densely punctate except at middle;

last sternite subtruncate at apex. Length, 13 mm.

Holotype female (United States National Museum) from Volcan de Chiriqui,

1000 M., Panama. (Friedr. Tippmann, Wien).

This is the only known species of reddish Megachoriolaus with a black pronotal

band. The rather fine dense punctation of the pronotum and the dense pubescence

of the elytra also make this species distinctive.

Megachoriolaus nigricollis, new species

Male (?) : Form moderately robust, elytra subparallel, slightly tapering toward

apices; integument black, elytra almost Nopal red (Ridgeway, 1912). Head with

vertex vaguely impressed on each side of midline, punctures vague, pubescence

dense, golden, appressed; front elongate, narrow, ratio of distance between apices

of antennal tubercles and line between them and apex of labrum, 1.0:2. 3; antennal

tubercles not prominent, minutely shallowly punctate, median glabrous triangle

large; clypeus finely punctate at base, labrum with several small punctures at sides;

genae very finely, sparsely punctate; antennae extending to about apical one-third

of elytra, segments to fourth shining, remainder opaque, segments from fifth thick-

ened, poriferous areas vague on outer segments, third segment slightly shorter

than first, fourth shorter than third, fifth equal to third. Pronotum moderately

densely punctate, punctures a little larger than those of elytra; pubescence red-

orange, dense, depressed; sides sinuate; posterior angles acute, extending over hu-

meri; prosternum finely punctate over apical %, pubescence very sparse; meso-

sternum with intercoxal process broad, protruding slightly above level of coxae,

Vol. 50, No. 1, January 1974

abruptly declivous in front; metasternum glabrous at middle, sides and margins

very finely, densely punctate, pubescence sparse. Elytra a little more than twice

as long as broad; punctures rather fine, well separated; pubescence moderately

dense, suberect, reddish; apices slightly obliquely truncate, outer angle obtusely

dentate. Legs with posterior tarsi about as long as tibiae, first segment as long

as remainder together; front tibiae densely pubescent internally; femora moderately

densely clothed with bronze pubescence externally. Abdomen very finely, densely

punctate at sides, glabrous at middle, last sternite almost all punctate, apex very

broadly emarginate, angle dentate. Length, 13-16 mm.

Holotype male (?) (California Academy of Sciences) and six male (?) para-

types from 30 miles NE Tehuantepec, Oaxaca, Mexico, 8 July, 1955 (Derham

Giuliani) .

No other known species of Megachoriolaus has the entire body except

the elytra black. The type series shows no variation in this color pattern

and all specimens are very similar. In addition to the black head and

pronotum, the characteristics of the head, mesosternum and abdomen

will separate M. nigricollis from other species.

This species has no obvious secondary sexual characters.

Megachoriolaus celestae, new species

(Fig. 1)

Male (?) : Form moderately robust, elytra subparallel before tapering at apices;

elytra a scarlet Brazil red (Ridgeway, 1912), head and pronotum orangish, mouth-

parts, antennae, eyes, legs and most of underside black, abdomen often partially

pale. Head with vertex shallowly impressed on each side of midline, finely, con-

fluently punctate, finely clothed with brownish depressed pubescence ; front elon-

gate, ratio of distances between apices of antennal tubercles and line midway be-

tween them and apex of labrum, 1.0:2. 3; antennal tubercles prominent, minutely

opaquely punctate to large glabrous median triangle, pubescence very dark, brown-

ish, depressed; clypeus rather coarsely, shallowly punctate over basal %, apex

impunctate, labrum impunctate; genae minutely, vaguely punctate; antennae ex-

tending to about apical one-third of elytra, basal segments to middle of fifth

shining, rather sparsely clothed with coarse, suberect black setae, outer segments

opaque, minutely densely punctate and pubescent, poriferous areas of outer seg-

ments small, third segment slender, about as long as first, fourth slender, shorter

than third, segments from apex of fifth very slightly thickened, fifth about as long

as third, eleventh bluntly tapering at apex. Pronotum very densely, moderately

coarsely punctate except for median impunctate line; pubescence dense, subde-

pressed, colored similarly to elytra; sides sinuate, posterior angles acute, extending

over humeri; prosternum impunctate, very sparsely clothed with very short pubes-

cence; mesosternum very finely, densely punctate, moderately densely clothed with

short, depressed, bronze pubescence, mesosternal process lying below surface of

coxae, gradually arcuate anteriorly; metasternum finely densely punctate, moder-

ately densely pubescent. Elytra about twice as long as broad ; punctures fine, dense,

separated by about two puncture diameters; pubescence dense, rather short, sub-

depressed, colored as surface; apices obliquely truncate, outer angles dentiform.

Legs with posterior tarsi slightly shorter than tibiae, first segment longer than

The Pan-Pacific Entomologist

Fig. 1. Megachoriolaus celestae Chemsak and Linsley.

Vol. 50, No. 1 , January 1974

remainder of tarsus; femora finely densely punctate externally, densely clothed

with short, dark and golden pubescence. Abdomen shining, finely densely punc-

tate basally, less densely at middle toward apex; last sternite narrowly rounded

at apex. Length, 13-15 mm.

Holotype male (?) (California Academy of Sciences) from SE side of Cerro

Tres Picos, Montane rain forest, 1524-1829 M., Chiapas, Mexico, 25 May 1972

(D. E. Breedlove) ; one male (?) paratype from N. slope of Cerro Bola, N. Cerro

Tres Picos, 1524-2134 m., Chiapas, Mexico, 5 May 1972 (D. E. Breedlove).

M. celestae is very distinctive by the unique scarlet red coloration of

the elytra; none of the other reddish species of Megachoriolaus has this

particular hue. The species also differs by the non-expanded elytra

and form and color of the head. Of the other red-headed species, M.

ignita (Schaeffer) has the pronotum more coarsely punctate; M. imi-

tatrix Linsley has shorter antennae and sinuate-truncate elytral apices,

and M. sabinoensis (Knull) has a much shorter front of the head.

As in most species in this genus, there are no apparent secondary sex-

ual differences. It is equally possible that both our specimens may be

females or that both sexes are represented. The shape of the apex of

the last abdominal sternite in M. celestae is quite unlike most other spe-

cies which have the apex emarginate truncate and the angles dentate.

We take pleasure in dedicating this beautiful species to Celeste Green

in appreciation for the preparation of the illustrations which accompany

this and many of our prior publications.

Megachoriolaus gracilis, new species

Male (?): Form moderate sized, slender, tapering posteriorly; tips of mouth-

parts, eyes, antennae, tarsi, tibiae except bases of front pair, hind femora at

apical one-half, middle femora at apical one-third, and front pair narrowly at

apices, black, elytra approaching Nopal red (Ridgeway, 1912) in color, head,

pronotum, underside and parts of femora orangish. Head with vertex impressed

behind antennal tubercles, finely confluently punctate, pubescence very fine,

golden; front moderately elongate, ratio of distance between apices of antennal

tubercles and line midway between them and apex of labrum, 1.0 :2.2 ; antennal

tubercles shallow, very finely, shallowly punctate to large glabrous median tri-

angle, pubescence short, fine, subdepressed; clypeus with a few larger, shallow

punctures irregularly dispersed; labrum with a few shallow setiferous punctures;

genae almost impunctate; antennae almost as long as elytra, segments to middle

of fifth shining, clothed with coarse black hairs at apices, outer segments opaque,

poriferous areas obsolete, basal segments slender, segments from apex of fifth very

slightly thickened, third a little longer than first, fourth equal to first, fifth equal

to third. Pronotum moderately coarsely densely punctate except for midline be-

hind middle; pubescence moderately dense, reddish, depressed; sides sinuate, pos-

terior angles acute, extending over humeri; prosternum glabrous; mesosternum

with intercoxal process tapering, lying below surface of coxae, very gradually arcu-

ate anteriorly ; metasternum very finely, densely punctate, densely clothed with

The Pan-Pacific Entomologist

short depressed pubescence. Elytra a little less than two and one-half times as

long as broad; punctures fine, dense, separated by about one puncture diameter;

pubescence dense, moderately long, depressed, colored as surface; apices obliquely

rounded, outer angles dentiform. Legs with posterior tarsi about as long as tibiae,

first segment longer than remainder of tarsus; femora slender, densely punctate.

Abdomen shining, finely densely punctate at side; last sternite narrowly rounded

at apex. Length, 11-13 mm.

Holotype male (?) (California Academy of Sciences) from La Pacifica, 4 Km

N.W. Canas, Guanacaste Province, Costa Rica, 30 May 1972 (P. A. Opler) ;

three male (?) paratypes with same data.

The moderately slender, tapering body form of this species makes it

easily recognizable. The coloration of the body is also distinctive. In

this respect there is no variation within the type series.

Eurylemma, new genus

Form moderately robust; elytra expanded posteriorly. Head with front short,

deeply impressed transversely; palpi unequal, apical segments cylindrical; eyes

small, finely faceted, deeply emarginate, upper lobes small, widely separated above ;

tempora prominent, elongate, parallel; antennae short, segments moderately thick-

ened, basal segments with coarse setae at apices, poriferous areas vague, scape

robust, longer than third segment, fourth shorter than third, fifth equal to first.

Pronotum rather flattened, sides sinuate; hind angles acute, not extending over

humeri; apex about as broad as head, basal margin broadly lobed; disk impressed

at sides; prosternum narrow, deeply impressed at apex, intercoxal process slender,

expanded at apex, coxal cavities partially open behind, coxae rather small; meso-

sternum with intercoxal process lying below surface of coxae. Elytra with broad,

elevated lines; apices broadly rounded. Legs slender, short; posterior tarsi shorter

than tibiae, first segment a little shorter than remainder of tarsus together. Ab-

domen normally segmented.

Type species: Eurylemma auricollis, new species.

This genus is apparently related to the group of genera presently re-

ferred to as the Euryptera-Megachoriolaus complex. It superficially re-

sembles Euryptera but differs by the more slender antennae, longer

tempora, broader front of the head, non-elevated mesosternal process,

and the absence of dentate angles at the apex of the last abdominal ster-

nite. The shorter front of the head, prominent tempora, shape of the

pronotum, and costate elytra will distinguish Eurylemma from Mega-

choriolaus.

A single lycid-like species is presently known.

Eurylemma auricollis, new species

(Fig. 2)

Female: Form moderate sized, depressed, elytra expanding posteriorly; integu-

ment black with a bluish overcast, pronotum broadly yellow down sides, pubescence

dark, golden on head and sides of pronotum. Head with front rather short, antennal

tubercles short, finely punctate; clypeus with a few scattered punctures, lahrum

Yol. 50, No. 1, January 1974

Fig. 2. Eurylemma auricollis Chemsak and Linsley

The Pan-Pacific Entomologist

almost impunctate; vertex broad, minutely punctate, each side with a band of

dense, golden, appressed pubescence extending from antennal tubercles to neck;

antennae extending to about middle of elytra, segments slightly thickened, segments

to base of fifth shining with a metallic caste, remainder opaque. Pronotum broader

than long; sides yellowish, densely clothed with golden recumbent pubescence;

disk impressed at sides near base; punctures fine, contiguous, middle with a glab-

rous longitudinal line; pubescence at middle short, dark, depressed; prosternum

vaguely punctate, pubescence sparse; meso- and metasternum very finely, separ-

ately punctate, moderately clothed with fine subdepressed pubescence. Elytra about

three times as long as basal width, broadly expanding toward apices; each elytron

rather strongly tricostate and with a feeble costa toward lateral margin; punctures

minute, dense; pubescence short, depressed; apices broadly rounded. Legs slender,

short, finely pubescent. Abdomen shining, almost glabrous; last sternite truncate

at apex. Length 17 mm.

Holotype female (California Academy of Sciences) from Cerro Huitepec, W.

of San Cristobal de las Casas, 2591 M., Chiapas, Mexico, 23 May, 1972 (D. E.

Breedlove) .

This species has a very distinctive appearance as a result of the dark

integument with a bluish caste and the golden recumbent pubescence of

the head and pronotum.

Euryptera albosterna, new species

Female: Form moderate sized, elytra strongly expanding posteriorly; color

black, head with parts of front and vertex yellow orange, pronotum with two broad

red bands down sides extending over humeri, humeri red, red band extending

acutely back to about basal %, underside of head, middle of prosternum and

mesosternum yellow; pubescence black on dark areas, bright red and appressed on

red bands of pronotum and elytra. Head moderately elongate, ratio of distance

between apices of antennal tubercles and a line midway between them and apex

of labrum, 1.0:1.75; front moderately impressed, indistinctly punctate, median tri-

angle vague, clypeus sparsely, moderately coarsely punctate, labrum sparsely, mod-

erately coarsely punctate, pubescence fine, subdepressed; vertex plane, median

line becoming a carina to neck, punctures vague, pubescence short, subdepressed;

genae finely, sparsely punctate; palpi very unequal, last segment of maxillary very

elongate; antennae rather stout, extending to about basal % of elytra, basal seg-

ments shining, clothed with coarse black setae, segments from sixth opaque, mi-

nutely pubescent, scape conical, longer than third segment, third longer than

fourth, fifth longer than third. Pronotum minutely punctate; pubescence long,

appressed, colored as background; sides sinuate, hind angles acutely produced over

humeri; base lobed at middle; prostemum glabrous, not impressed, intercoxal pro-

cess fairly broad, rounded; mesosternum with intercoxal process broad, extending

above coxae, abruptly declivous in front; metasternum minutely punctate, moder-

ately densely pubescent. Elytra about 2% times as long as basal width, broadly

expanding toward apices; punctures minute, dense; pubescence dense, suberect,

depressed along humeral red area, sides fringed; each side of suture at basal %

with a strong costa terminating in a sharp spine at apical inner angle; epipleurae

strongly margined; apices broadly emarginate, outer angles dentate. Legs short,

slender; posterior tarsi shorter than tibiae, first segment shorter than remainder

Vol. 50, No. 1, January 1974

of tarsus. Abdomen minutely, densely punctate, segments narrowly glabrous pos-

teriorly; last segment sinuate at apex, outer angles spinose, Length, 12 mm.

Holotype female (California Academy of Sciences) from Turrialba, Costa Rica,

31 January 1973 (V. M. Kirk) .

This species shares most of the generic characteristics of Euryptera

latipennis Serville, and the only relative differences of E. albosterna are

the wider prosternal process and reduced costae of the elytra.

E. albosterna may be readily recognized among the Central American

eurypteroid lepturines by the appressed and suberect pubescence of the

elytra, the distinctive red and black dorsal coloration, and by the sutural

costae and spines of the elytra.

We wish to thank V. M. Kirk for making this specimen available for

study.

Genus Mimiptera Linsley

Mimiptera Linsley, 1961, Jour. New York Entomol. Soc., 69:139. Type species:

Euryptera fulvella Bates, by original designation.

Specimens of this genus are extremely rare in collections.

Since the genus was described only one additional example has been

seen, from Hacienda Comelco, 24 Km. NW Canas, Inter-Am. Hwy.,

Guanacaste, Costa Rica, on flowers of Casearia, 16 November, 1971

(E. R. Heithaus). Based on the coloration, it would be regarded as M.

fulvella (Bates) but structurally it is identical with M. costaricensis

(Melzer). As in other mimetic species, such as those of the genus Ely-

troleptus (Chemsak and Linsley, 1965), M. fulvella apparently has more

than one lycid-like color form, that is with the elytra transversely black

banded or wholly yellow. M. costaricensis is therefore considered synon-

ymous with M. fulvella.

Literature Cited

Chemsak, John A. and E. G. Linsley. 1965. A revised key to the species of

Elytroleptus with notes on variation and geographical distribution. Pan-

Pacific Entomol., 41: 193-199.

Fuchs, E. 1956. 2. Beitrag zur kenntnis der neotropischen Cerambyciden. En-

tomol. Art. Mus. G. Frey, 7: 567-576.

Linsley, E. G. 1961. The North and Central American species of Euryptera and

a related new genus. Jour. New York Entomol. Soc., 69: 131-141, 1 fig.

Linsley, E. G. 1970. New genera and species in the lepturine complex related

to Euryptera and Choriolaus. Pan-Pacific Entomol., 46: 128-141, 4

figs.

Linsley, E. G. and John A. Chemsak. 1971. An attempt to clarify the generic

status of some Neotropical species currently assigned to Euryptera,

Chontalia and Ophistomis. Arq. Zool., S. Paulo, 21: 1-40.

Ridgeway, R. 1912. Color Standards and Color Nomenclature. 43 pp., 53 pis.

The Pan-Pacific Entomologist

Larvae and Pupae of Some Wrack Dipterans

on a California Beach

(Diptera: Coelopidae, Anthomyiidae, Sphaeroceridae)

Helen Kompfner

Hopkins Marine Station of Stanford University

Introduction

The kelp (wrack) flies, Coelopa vanduzeei Cresson (1914), Fucellia

costalis Stein (1910), Fucellia rufitibia Stein (1910), and Leptocera

johnsoni Spuler (1925) are common on California beaches (Cole,

1969). The life histories of such related European species as Fucellia

maritima and Coelopa frigida have been studied (Egglishaw, 1960a and

b), but little is known about the immature forms of the Pacific coast

species. Preliminary observations of fly larvae found living in wrack

suggested that a study of distribution and biology of larvae on the beach

would be valuable in further understanding how different species of

flies relate to the wrack, as a food source and habitat. This investiga-

tion was carried out April— June, 1972, on two pocket beaches in Mon-

terey Bay, California: (1) the west beach of Mussel Point, Pacific

Grove, and (2) the Point Lucas beach, Pacific Grove. My studies of

larvae were accompanied by a study of the activity patterns and be-

havior of adult wrack dipterans by another investigator (Hyatt, 1972)

at Mussel Point.

Characteristics of Larvae and Pupae

(Fig. 1 and 2)

Larvae were collected in the field and raised on samples of damp

wrack kept in the laboratory at room temperature. Initial identifications

were made on the basis of emerged adults. Samples of these adult flies

were examined and identified by Dr. Paul Arnaud of the California

Academy of Sciences, and George C. Steyskal of the U.S. Dept, of

Agriculture.

Coelopa vanduzeei. — Third instar larva. Length 9.0-12.0 mm, great-

est width and height 1.5 mm at caudal end, body narrowing anteriorly,

second segment 0.5 mm wide. Color white, with grey ventral spines on

abdominal segments. Caudal plates yellow with black borders, sur-

rounded by pale hairs; 3 spiracles. Anus four-lobed, located ventro-

posteriorly on segment 12. Anterior prothoracic spiracles bi-lobed, each

The Pan-Pacific Entomologist 50: 44-52. January 1974

Vol. 50, No. 1, January 1974

Fig. 1 . Larval structures of wrack dipterans discussed in text. A. Coelopa

vanduzeei B. Fucellia costalis C. Fucellia rufitibia D. Leptocera johnsoni. (1)

left lateral view of mouthparts (2) left lateral view of entire third instar larva

(3) caudal view, showing caudal plates and posterior spiracles.

lobe marginally scalloped to form 10 petaldike processes. Antennae

smooth, finely pointed.

Puparium. Length 3. 5-6.0 mm; greatest width at segment 8, about

half this ; greatest height at segment 8, approximately one third length ;

ventral surface and dorsal surface of anterior three segments flattened;

dorsum arched; lateral edges of each segment indented; color yellow-

brown to brown-black.

Fucellia costalis . — Third instar larva. Length 9.0-12.5 mm; body

widest at caudal end, 1.5 mm, tapering to 0.5 mm at anterior end of

second segment. Color white, or off-white. Caudal plates white with

three amber spiracles, no caudal plate hairs. Anterior prothoracic spi-

racles each with a single cluster of 8-10 lobes. Antennae blunt-tipped

with annular ridges.

Puparium. Length 6.0-7.0 mm; greatest width and height at seg-

ments 7 and 8, 3.0 mm. Shape fusiform, circular in cross-section. Color

dark reddish brown, segments 1-11 with wrinkled surface, segment 12

matte.

The Pan-Pacific Entomologist

1mm

Fig. 2. Puparia, dorsal views. A) Coelopa vanduzeei , B) Fucellia costalis, C)

Fucellia rufitibia D) Leptocera johnsoni.

Fucellia rufitibia . — Third instar larva. Length 8.5—10.5 mm, greatest

width 1.2 mm, at caudal end, tapering to 0.3 mm at anterior end of seg-

ment 2. Color white, grey at twelfth segment. Caudal plates white with

yellow or red spiracles, no caudal plate hairs. Anterior prothoracic spi-

racles each one-lobed, with a single cluster of 7-11 processes. Antennae

blunt-tipped, with annular ridges.

Puparium. Length 4.0-5. 0 mm, greatest width and height at segments

7 and 8, 1.5 mm. Shape fusiform, circular in cross-section. Color red-

dish brown, surface texture smooth, shiny.

Leptocera johnsoni . — Third instar larva. Length 2. 0-3. 5 mm. Color

white, with black spines on abdominal ridges at anterior end of segments

6-12. Caudal plates on caudal stalks, plates black surrounded by long,

black hydrophobic hairs in tufts of 4 or 5. Prothoracic spiracles with

5 slender radiating marginal processes.

Puparium. Length 2. 0-2. 5 mm, width and height greatest at seg-

ments 5-11, 0.8 and 0.5 respectively, dorso-ventrally flattened especially

segments 1-2 and 12. Color yellow brown, segments 1-4 often translu-

cent. Segment 12 has two caudal stalks.

Field Studies

Distribution of fly larvae was studied on beaches at Mussel Point

and Point Lucas. Samples were taken at high, mid and low sections

of the beach. Since most larvae were found in wrack banks (piles greater

than 15 cm in depth, as defined by Backhand, 1945), vertical distribu-

tion within the banks was also studied. Samples of wrack from various

elevations on the beach, and samples taken at depths of 10, 20, and

Vol. 50, No. 1, January 1974

111

I-

111

Fig. 3. Distribution of larvae on beach in relation to tidal height.

30 cm within wrack banks, were collected in plastic bags and returned

to the laboratory. Larvae were extracted from the wrack by shaking

the samples vigorously with water, then capturing the smaller larvae

with a fine aquarium net. The wrack sample was then carefully examined

for any remaining larger larvae.

Fig. 3 shows the distribution of fly larvae from wrack banks stranded

at various elevations above mean lower low water. The majority of

wrack deposits at Mussel Point centered around the +3.0 tidal level

and consisted of mixed red and brown algae and the surf grass Phyl-

lospadix. The major deposits at Point Lucas consisted of banks of the

brown alga Macrocystis concentrated on the upper beach at tidal heights

of +5.0 to 6.5 feet. Coelopa vanduzeei larvae were found exclusively

in lower beach wrack banks. Fucellia costalis larvae were found almost

exclusively in the upper beach banks. F. rufitibia larvae were found at

all levels, but predominately on the mid and upper beach. L. johnsoni

larvae also occurred widely on the beach but were more evenly dis-

tributed. The occurrence of larvae in wrack corresponds well with the

distribution of the adult flies found on the same beach in studies by

Hyatt (1972).

Vertical distribution of fly larvae within wrack banks is shown in

Fig. 4A, B. In banks on the lower beach Coelopa vanduzeei predomi-

The Pan-Pacific Entomologist

nated in the top 10 cm, and L. johnsoni at the lowest levels. A small

number of F. rufitibia were found, largely at intermediate levels. F.

costalis was absent. Observation of a lower beach bank before and

after complete immersion during a high tide showed that numbers of

C. vanduzeei present did not change ; numbers of L. johnsoni diminished,

however, suggesting a loss accompanying wave action. In the upper

beach bank (Fig. 4B) Fucellia costalis larvae predominated, and F.

rufitibia outnumbered L. johnsoni at the lowest levels. C. vanduzeei

was absent. High beach wrack banks tended to dry out, and as this

happened, larger larvae were observed moving down in wrack over

time.

Observations on Feeding, Behavior, and Temporal Variation

Larvae appeared to eat the algae on which they were found. Frag-

ments of Macrocystis were found in the guts of all species. Larvae were

never observed on very fresh wrack; apparently some decomposition

must occur before the algae are eaten. Nematodes were abundant where

fly larvae were found and may be involved in initial algal breakdown,

as suggested by Egglishaw (1960b).

Wrack consisting of the kelp Macrocystis seemed to be preferred

above other algae by all larvae collected from natural wrack. When

wrack was experimentally sorted out into separate piles containing

only brown algae, only reds, only Phyllospadix, or a mixture of all

types, and placed on the beach for several days, larvae were only found

on brown and mixed wrack. Larvae placed on red algae and Phyllo-

spadix died within a day in the lab, while those on Macrocystis flour-

ished. Those kept without food died within two days, and larvae kept

without moisture died overnight.

Of the four species observed, the larvae of C. vanduzeei and L. john-

soni were the most active and moved most rapidly.

Over the 5 week period of the study some temporal variation was

observed in the numbers of C. vanduzeei larvae. In the last two weeks

of April none were found on the beach at Mussel Point. By May 2, they

were found in small numbers and in the week of May 8-15, occurred

in multitudes in buried wrack on the lower beach there and further south

along the coast at Pebble Beach and Carmel. The latter period coincided

with neap tides, and formation of major wrack deposits on the beaches.

By May 17 no more larvae were found and the wrack was also gone

from the beach. Fucellia costalis , F. rufitibia , and Leptocera johnsoni

were found in fairly constant numbers, but never as abundantly as

Vol. 50, No. 1, January 1974

C. vanduzeei

F ruf itibia

L. johnson i

F. costal i s

13% 13%

74 /

86 %

14 1

59 %

27 % 1 4 %

N=15

N=7

N^5t

377o . 62%

1 %

N=73

N=61

17%

52% 31% N=142

10 20

1 0

20 30

WRACK HEIGHT IN CENTIMETERS

Fig. 4. A) Vertical distribution of larvae in lower beach wrack bank. B) Ver-

tical distribution of larvae in upper beach wrack bank.

Coelopa vanduzeei at its peak, over the time interval noted. Some adult

L. johnsoni were observed mating in all wrack deposits observed over

5 weeks.

Laboratory Rearing of Larvae

Attempts to obtain eggs and subsequent larval stages in the laboratory

from captive populations of flies maintained on wrack in large plastic

bags were not successful. However, larvae of C. vanduzeei , F. costalis ,

F. rufitibia , and L. johnsoni were reared successfully on Macrocystis

stipes (examined first for foreign larvae or eggs) in Petri dishes at

room temperature. Cultures were wetted and new Macrocystis added

every two days. Pupae were removed, placed in corked vials, and

checked daily for hatching. This method of rearing does not eliminate

the possibility that some larave may have fed on each other. Some

species of Fucellia are known to be predaceous as larvae, i.e. F. rejecta

Aldrich.

The picture of development obtained, while incomplete, provides the

beginning of a timetable for the life history. Data available are shown

in Fig. 5. Duration of the third instar was shortest in Coelopa vanduzeei ,

longest in Fucellia costalis. The former lives lowest in the intertidal

zone, the latter highest on the beach. Duration of pupal period showed

considerable variation among species and within C. vanduzeei. For the

latter, there may be some connection between intertidal height on the

beach and length of pupal period. Although later development took

The Pan-Pacific Entomologist

unknown

N Larva second I | Pupa

i nstar

Fig. 5. Developmental period of lab reared larvae.

place in the laboratory, the 12 individuals of C. vanduzeei which hatched

after a 4 day pupal period were collected from buried wrack at the +3.0

foot tide level, while the individual which spent 9 days as a pupa was

taken from an upper beach wrack bank. L. johnsoni had a pupal period

of 6-7 days, F. rufitiba and F. costalis 12—15 days. F. costalis larvae

may remain in the third instar for considerable periods; one batch of

10 remained in this stage for 25 days before pupation.

Discussion

The flies appear to use different strategies in exploiting the available

food and habitat resources for larvae on the beach. C. vanduzeei is

found almost exclusively in lower beach wrack which is wetted at least

once a day and seldom remains on the beach longer than two weeks.

The main pressure here is for quick larval development and a short

pupal period. The caudal plate hairs allow the larva to breathe from

air bubbles trapped in the wrack during periods of tidal immersion.

Immature F. costalis and R. rufitibia, found mainly on the mid and

upper beach levels, are able to use wrack for longer periods. Caudal

hairs are lacking. The danger of being washed out to sea is less than

that of dessication. The pupal period reflects this, being relatively long.

L. johnsoni larvae are found in large numbers and are distributed both

high and low on the beach. Their caudal plate hairs allow them to re-

spire when submerged in wrack beds low on the beach, but their small

size makes them vulnerable both to being swept out of low immersed

wrack, and to desiccation in higher beach wrack. Perhaps this is why

Vol. 50, No. 1 , January 1974

L. johnsoni tends to occur well below the surface of wrack banks. A

comparison of mean tidal height of distribution and mean pupal period

for each species suggests that species found higher on the beach may

have longer pupal periods, although the variation within each species

is large enough to eliminate meaningful statistical comparison with

present data.

Summary

Larvae and pupae of Coelopa vanduzeei , Fucellia costalis, Fucellia

rufitibia, and Leptocera johnsoni were studied on two California beaches.

Third instar larvae and puparia are described. Studies of distribution

of larvae on the beach showed C. vanduzeei occupies lower beach wrack

banks and F. costalis the upper beach banks. F. rufitibia larvae, while

widespread, occur more in mid and upper beach wrack, while those of

L. johnsoni are found at all levels. Larvae were observed to prefer the

kelp Macrocystis both as a habitat and food source. C. vanduzeei larvae

were observed in large numbers only during a two week period coin-

ciding with a major wrack deposit. F. costalis, F. rufitibia, and L. john-

soni were found in constant numbers over 5 weeks. Duration of pupal

periods in the laboratory were: C. vanduzeei, 4-9 days; L. johnsoni,

6-1 days; F. costalis and F. rufitibia 12-15 days. Average duration of

the pupal period appears to increase with increasing tidal height of the

larval habitat.

Acknowledgments

I would like to thank Dr. Paul Arnaud of California Academy of Sci-

ences and George C. Steykskal of the U.S. Department of Agriculture

for identifying the adult flies of this study, and Joel Hyatt, for many

helpful discussions about flies and larvae on the beach. I am especially

indebted to Dr. Donald P. Abbott of Hopkins Marine Station, for his

unfailing patience and encouragement with this study and the editing

of this manuscript. Dr. I. A. Abbott also provided much appreciated

encouragement during this study.

Literature Cited

Backlund, H. O. 1945. Wrack fauna of Sweden and Finland. Ecology and

chorology. Opuscula Entomol. Suppl., 5: 1-238, Lund, Sweden.

Cole, F. R. 1969. The flies of Western North America. University of California

Press, Berkeley.

Egglishaw, H. 1960a. The life history of Fucellia maritima (Haliday) (Diptera,

Muscid.) Entomologist, 93: 225-231.

The Pan-Pacific Entomologist

Egglishaw, H. 1960b. Studies of the family Coelopidae (Dipter) . Trans. R.

Entomol. Soc., 112: 109-140.

Hyatt, Joel. 1972. Behavior of wrack dipterans, F. rufitibia (Anthomyiidae)

and C. vanduzeei (Coelopidae) on a California beach. Unpub. MS on

file at Hopkins Marine Station Library.

SCIENTIFIC NOTE

The Occurrence of Trichocorixa reticulata in the Gulf of California

(Hemiptera: Corixidae). — During the course of ground observations in sup-

port of the NASA Earth Resources Satellite Program 1 , plankton tow samples were

taken in the delta region of the Colorado River at the extreme northern end of

the Gulf of California. One of these tows (Plankton Station D-4 at N. end of Isla

Montague in the Colorado River Delta, 31° 49.9' N. latitude, 114° 48.5'W. longi-

tude, 16 December 1972, J. R. Hendrickson) included a female of Trichocorixa

reticulata (Guerin-Meneville) .

While the occurrence of this species in salt water has been noted previously

(see Williams 1944, Proc. Haw. Ent. Soc., 12: 186-196, and Sailer 1946, Proc.

Haw. Ent. Soc., 12 : 617-620) , it has not been recorded from ocean waters. Wil-

liams found it in ponds more saline than sea water, accounting for its success in

colonizing salt water habitats.

The locality where the specimen was collected is one of strong tidal currents

produced by semidiurnal tides which can vary by as much as 9 meters between

extreme high and extreme low. The water carries a very heavy silt load (standard

Secchi disk visibility range is 2-15 cm depth) ; salinity has been measured as low

as 37 and as high as 44 parts per thousand, depending upon tidal state and season

(world normal salinity is about 35 parts per thousand, normal for the northern

Gulf of California is about 37 parts per thousand) . Accompanying the specimen

were the normal components of a marine plankton tow (copepods, arrow worms,

jellyfish, marine fish juveniles) . The specimen was swimming actively in the con-

centrated plankton sample when first noted. — J. T. Polhemus, Martin Marietta

Aerospace, Denver, Colorado 80201 and J. R. Hendrickson, Department of Bio-

logical Sciences, University of Arizona, Tucson 85721.

1 NASA Contract NAS5-21777, Identification No. UN603. This work was also supported by NSF

Grant GB34675.

Vol. 50, No. 1, January 1974

Observations on Primary and Secondary Parasitoids of

California Oakworm, Phryganidia calif ornica. Pupae

(Lepidoptera: Dioptidae)

David J. Horn 1

Department of Entomology, The Ohio State University, Columbus 43210

Introduction

The California oakworm, Phryganidia californica Packard, periodi-

cally defoliates evergreen and deciduous oaks ( Quercus ) in coastal Cal-

ifornia. In the San Francisco Bay region, irruptions occur at 4- to 7-

year intervals and last 2-3 years each, followed by spectacular declines

in numbers. The oakworm normally has two generations a year. Larvae

overwinter on evergreen oaks, particularly coast live oak, Quercus agri-

folia Nee, and feed on new foliage in April and May. Pupation occurs as

a chrysalid on branches and trunks of oaks, and in surrounding vegeta-

tion, during June. The weak-flying adults emerge in a few weeks, mate,

and oviposit on both evergreen and deciduous oaks, especially Q. lobata

Nee, valley oak. Eggs hatch, and larvae feed in July and August, to pu-

pate in September and emerge as adults in October and November.

These oviposit on evergreen oaks, eggs hatch, and the larvae may feed

but are usually inactive until the onset of warmer weather and new

foliage in late March and April (Harville, 1955, and my observations).

Early workers (Kellogg and Jack, 1895; Doane, 1912) attributed

oakworm population declines to parasitization because dense infestations

usually yielded high percentages of parasitized oakworm pupae. Har-

ville (1955), following exhaustive field study in which pupal parasitism

never exceeded 75%, concluded that population declines were caused

by a variety of factors, particularly cold winter weather killing over-

wintering larvae.

Among the pupal parasitoids, Harville found Itoplectis behrensii

(Cresson) most commonly. It is probably an obligate parasitoid of

oakworm pupae (though Muesebeck et al., 1951, cite its presence also

in Recurvaria milleri Busck) . It is bivoltine, like its host, and the long-

lived adults can be collected throughout the year (Townes and Townes,

1960). Harville (1955) observed that I. behrensii females hunt tac-

tilely on oak branches and trunks, “stumbling” onto potential hosts, and

being unable to distinguish previously parasitized hosts. He found that

1 Formerly Department of Biological Science, California State University, Hayward.

The Pan-Pacific Entomologist 50: 53-59. January 1974

The Pan-Pacific Entomologist

Table 1. Population density, parasitization, and survival of Phryga-

nidia californica pupae. Hayward, Alameda Co., Calif. May-July.

Year

Pupae/ft 2

Sample

size

Total

parasitization

Moth

emergence

1969 1

23.4

270

110 (41%)

100 (37%)

1970

30.3

478

339 (71%)

56 (12%)

1971

.01

2 (13%)

12 (80%)

1972

0.4

2 (6%)

26 (74%)

1 Sampling

commenced 20 June.

Parasitization

probably lower, and moth

emergence greater, for

whole season, than indicated value.

/. behrensii in turn was parasitized by 3 secondary parasitoids: Gelis

tenellus (Say), Mastrus aciculatus (Provancher) , and Dibrachys cavus

(Walker) ; the total incidence of parasitism by all three on I. behrensii

was less than 1% in eight study sites though D. cavus was sometimes

locally more abundant. Harville also found Brachymeria ovata (Say)

as a primary pupal parasitoid of the oakworm. It has no secondary

parasitoids. B. ovata is facultative, having been recorded from over

100 species of lepidopterous pupae (Burks, 1960), and Harville sug-

gested that it might survive on alternate hosts, possible species of Mala-

cosoma , during times of oakworm scarcity.

My preliminary observations of oakworm and parasitoid populations

near Hayward, California in 1969 revealed more extensive secondary

parasitization of 1. behrensii than that Harville reported. This suggested

that the interrelationships between oakworm populations and those of

the several parasitoid species warranted further study.

Methods

I selected a wooded hillside adjacent to the Ecological Field Station,

California State University, Hayward. The vegetation of the study area,

described in detail by Cogswell (1966) is a broadleaf evergreen wood-

land, dominated by coast live oak and California laurel ( Umbellularia

californica ) . No insecticides have ever been used here. In 1969-70,

but not in 1971—72, the spring generation of oakworms severely defoli-

ated oaks in the study area.

On each visit to the study plot in 1969-70 and 1972, I counted fresh

oakworm pupae in each of 10 1ft 2 areas of tree-trunk undersurface

chosen randomly from a group of 5 trees. Pupae were so few in 1971

that I obtained no meaningful estimate of density.

I determined the extent of parasitism by collecting a sample of 50-

Vol. 50, No. 1, January 1974

Table 2. Emergence of parasitoid species from Phryganidia cali-

fornica pupae. Hayward, Alameda Co., Calif. June-July 1969, May-

July 1970.

Year

Total

parasitoids

behrensii

ovata

tenellus

aciculatus

Bathy-

thrix

cavus 1

1969

110

42 1

1970

339

107

72 1

1 Number of hosts parasitized, not number of emerging parasitoids. D. cavus is gregarious, with

3-10 parasitoids/host.

100 pupae randomly from lower, accessible, oak limbs and trunks, and

kept pupae individually in 4-oz jars until moths or parasitoids had

emerged. I repeated the procedure every 5 days from the onset of pu-

pation until no new pupae appeared. Pupae from which nothing emerged

were visually inspected, then dissected. While collecting pupae, I spent

several hours observing the activities of I. behrensii and B. ovata. Wlien

I encountered pupae of other Lepidoptera, I saved these for possible

parasitoid emergence. These were primarily Hemerocampa vetusta Bois-

duval (Liparidae), Hemihylaea edwardsi Packard (Arctiidae), Mala-

cosoma calif ornicum (Packard), and M. constrictum (Edwards) (Lasio-

campidae) .

Results

Six species of parasitoids emerged from field-collected oakworm pu-

pae; 5 were those Harville (1955) found; the sixth, Bathythrix sp., a

secondary parasitoid of I. behrensii , represents a new host record.

Table 1 shows trends in oakworm numbers, pupal parasitization, and

moth emergence during 1969-72. Oakworm density and parasitization

increased from 1969 to 1970, then the oakworm population declined

dramatically. Fewer moths were produced in 1971-72 than in 1969-70,

but the percentage produced was much greater.

Table 2 shows relative proportions of the several parasitoid species in

1969-70. Parasitoids of I. behrensii were much more abundant in 1970

than in 1969. In 1971-72 I found only two parasitized oakworm pupae

each year; all yielded I. behrensii. No secondary parasitoids or B. ovata

were found in oakworm pupae in 1971 or 1972. I reared B. ovata from

one pupa of Hemerocampa vetusta and one of Hemihylaea edwardsii.

The latter is a new host record. From 35 pupae of Malacosoma spp. in

1972, I reared two G. tenellus but no B. ovata.

Fig. 1 illustrates seasonal trends in the incidence of parasitization

The Pan-Pacific Entomologist

Fig. 1. Trends in oakworm pupal density and parasitization at Hayward, Ala-

meda County, California, 1970.

Vol. 50, No. 1, January 1974

in 1970, the year for which I obtained most data. Parasitization by

B. ovata, though never exceeding 10%, was greatest on earlier oakworm

pupae, with a small second generation in early July. Percent parasitiza-

tion by 7. behrensii increased from 23% in May to over 90% in July.

Over the entire sampling period from May to July, oakworm pupal

mortality was 88%, as follows: B. ovata 4%, 7. behrensii 67%, and un-

known 17%. Some of the unknown mortality may have been due to

fungal infections and some may have resulted from oviposition by para-

sitoids or feeding by predaceous Hemiptera (Horn, 1973) . Occasionally,

I encountered desiccated parasitoid larvae within oakworm pupae. These

are included as unknown mortality because I could not determine the

species of parasitoid involved.

The incidence of secondary parasitization of I. behrensii increased

from 0 in May to 100% in late June and July. G. tenellus , D. cavus , and

Bathythrix sp. were equally abundant and showed no seasonal separation,

while Mastrus was much less common (Table 2). Together, secondary

parasitoids killed 69% of Itoplectis present in the samples over the sea-

son. A few B. ovata may have been parasitized in early July.

I observed searching behavior of 10 I. behrensii females in 1970 for

periods up to 45 minutes each. My observations agreed with those of

Harville (1955), that 7. behrensii searches very rapidly and efficiently,

running over the branches and trunk, and ovipositing readily in pre-

viously parasitized hosts. In every case, females turned away when en-

countering their own trail, indicating the presence of a trail odor (Price,

1970). Two B. ovata females that I observed moved about more slowly

than 7. behrensii and oviposited in nearly all oakworm pupae encount-

ered. One moved directly from an oakworm pupa to oviposit in a co-

coon of 77. vetusta. Superparasitism apparently occurs readily in each

species.

Discussion

Conclusions based upon observation of a local population through

only a portion of a cycle are tentative at best, but the increase in para-

sitization immediately preceding the oakworm population decline sug-

gests that 7. behrensii may be partially responsible for oakworm pop-

ulation declines.

Of interest are the interactions among the parasitoids and their use

of a fluctuating resource. 7. behrensii, having no alternate hosts, must

disperse readily and search quickly, as its probability of survival is low,

especially in years when oakworm pupae are scarce. It averages twice

the size of B. ovata, has much longer wings, and adults are relatively

The Pan-Pacific Entomologist

long-lived. Other parasitoid species display these characteristics in un-

stable environments (Price, 1972). B. ovata, being facultative, moves to

alternate hosts, at least to Hemerocampa vetusta and Hemihylaea ed-

wardsii, when oakworms are scarce. It is smaller and stockier than I.

behrensii and is thus capable of searching a wider variety of habitats,

including ground litter and bark crevices, as well as the surfaces of

branches and trunks. This befits a parasitoid with over 100 recorded

hosts. Malacosoma may not be as likely a reservoir of alternate hosts

as Harville (1955) suggested. I did not find B. ovata in a small sample

(N = 35) of Malacosoma pupae, nor did Langston (1957) discover it

in a large survey of parasitoids of California Malacosoma, though Stehr

and Cook (1968) recovered it from Malacosoma elsewhere.

I. behrensii thus has some attributes of an r-strategist whereas B.

ovata, relatively, is a K-strategist (Force, 1972) ; populations of the

former fluctuate in number along with their fluctuating resource, while

numbers of the latter probably remain relatively constant due to the

supply of alternate hosts. B. ovata occurs earlier and later than the

bulk of I. behrensii in oakworm pupae. This suggests that I. behrensii

may be the superior competitor when both species parasitize a single

host pupa. In the laboratory I attempted to induce females of each spe-

cies to oviposit in each of 20 oakworm pupae, from which I recovered

14 I. behrensii and 5 moths. However, I was not certain whether that

B. ovata actually oviposited, so the results are inconclusive.

The facultative secondary parasitoids, at least Gelis tenellus and Di-

brachys cavus, also move to alternate hosts including parasitoids of

Malacosoma (Langston 1957) when /. behrensii is rare. Mastrus

aciculatus has been recorded from several hosts, mostly Tenthredinidae

(Bobb 1965, Price, 1970), as have species of Bathythrix (Muesebeck

et al. 1951). There was no seasonal separation of their occurrence on

I. behrensii, but they do differ in size and this may affect competition

among them. More must be known of their relative abundance on other

hosts to ascertain their competitive interactions, which may be rapidly

changing. For instance, Bathythrix occurred commonly in my 1970

field samples yet was not present in any of Harville’s study populations.

It may have recently broadened its host preference to include I. behrensii.

The total incidence of parasitization on I. behrensii by all 4 secondary

parasitoids (52% in 1969 and 69% in 1970) was far greater than the

1% reported by Harville (1955). This suggests that significant local

changes may occur in the parasitoid complex of the oakworm, with po-

tentially important consequences for its population dynamics.

Vol. 50, No. 1, January 1974

Acknowledgments

The work was partially supported by a grant from the Research

Foundation, California State University, Hayward. C. E. Dasch assisted

with identification of parasitoids. R. S. Horn helped search for host

pupae and she and J. P. Harville read drafts of the manuscript. The

insatiable appetites of C. A. and R. B. Horn kept me abundantly sup-

plied with 4-oz jars.

Literature Cited

Bobb, M. L. 1965. Insect parasite and predator studies in a declining sawfly

population. J. Econ Entomol., 58: 925-926.

Burks, B. D. 1960. A revision of the genus Brachymeria Westwood in America

North of Mexico (Hymenoptera: Chalcidae) . Trans. Amer. Entomol.

Soc., 86: 225-273.

Cogswell, H. L. 1966. Thirtieth breeding bird census 40. Broadleaf evergreen

forest with shrub-filled openings. Audubon Field Notes, 20: 637-643.

Doane, R. W. 1912. Some insect pests of the California live oaks. J. Econ. En-

tomol., 5: 346.

Force, D. C. 1972. r- and K-strategists in endemic host-parasitoid communities.

Bull. Entomol. Soc. Amer., 18: 135-137.

Harville, J. P. 1955. Ecology and population dynamics of the California oak

moth Phryganidia calif ornica Packard (Lepidoptera) Microentomol.,

20: 83-166.

Horn, D. J. 1973. Leptocoris rubrolineatus, an occasional predator of the Cali-

fornia oakworm, Phryganidia calif ornica. Pan-Pac. Entomol., 49: 196.

Kellogg, V. K. and F. J. Jack. 1895. The California phryganidian. Proc. Calif.

Acad. Sci. (Ser. 2), 5: 562-570.

Langston, R. L. 1957. A synopsis of hymenopterous parasites of Malacosoma

in California. Univ. Calif. Publ. Entomol., 14: 1-50.

Muesebeck, C. F. W., K. V. Krombein and H. K. Townes. 1951. Hymenoptera

of America north of Mexico. Synoptic catalog. USDA Agr. Monog., 2:

1-1420.

Price, P. W. 1970. Trail odors: recognition by insects parasitic on cocoons.

Science, 170: 546-547.

Price, P. W. 1971. Parasitoids utilizing the same host: adaptive nature of dif-

ferences in size and form. Ecology, 53: 190-195.

Stehr, F. W. and E. F. Cook. 1968. A revision of the genus Malacosoma Hubner

in North America (Lepidoptera: Lasiocampidae) : systematics, biology,

immatures, and parasites. U.S. Nat. Mus. Bull., 276: 1-321.

Townes, H. and M. Townes. 1960. Ichneumon flies of America north of Mexico.

2. Subfamilies Ephialtinae, Xoridinae, Acaenitinae. U.S. Nat. Mus.

Bull., 216: 1-676.

The Pan-Pacific Entomologist

Biology of Three Callirhytis Gall Wasps

from Pacific Slope Erythrobalanus Oaks

(Hymenoptera: Cynipidae)

D. Charles Dailey, Tim Perry, and Christine M. Sprenger

Sierra College, Rocklin, California 95677

This paper presents life histories of two species of alternating unisex-

ual and bisexual generation gallmaking cynipids, Callirhytis eldoradensis

(Beutenmuller) (= Andricus eldoradensis Beutenmuller, 1913) and

Callirhytis flora Weld $2 (1922a) ( = Callirhytis milleri Weld 9,

1922b) new synonym, and a description of the males of a third species,

Callirhytis per foveata (Kinsey) $ 2 (1922).

All three of these gallmaking cynipid species occur on all three species

of Pacific slope oaks in the subgenus Erythrobalanus : Quercus agrifolia

Nee, Quercus kelloggii Newberry, and Quercus wislizenii A. DeCandolle.

Callirhytis eldoradensis (Beutenmuller), new combination

Callirhytis eldoradensis emerges from integral acorn hull galls in

aborted acorns (Fig. 1). The original description included only males

and erroneously associated them with galls of Callirhytis milleri Weld.

The bisexual generation female and correct gall were described by Weld

(1922). This generation is unusual because the male is larger than the

female and emergence is during late September and October rather than

in the spring or early summer. The previously undescribed unisexual

generation emerges in late spring or early summer from a spherical bud

gall (Fig. 2).

Biology. — During October 1970 a collection of C. eldoradensis was

reared indoors and individuals of both sexes were confined together

in bags on three native Q. wislizenii. On the following two days seven

naturally emerged (not escaped) males were collected on the bags’ ex-

teriors. This suggests the possibility of a female emitted pheromone.

When the natural female emergence was beginning and bagged females

began oviposition, three consecutive days of rain and cold weather inter-

fered with observations and caused a high mortality of reared and pos-

sibly naturally emerged insects.

When the bags were removed in late March, 1971 the twigs contained

green spherical bud galls about four mm in diameter with white basal

hairs. The galls abort the apical meristem and are surrounded basally

by the bud scales. Four of these galls figured as numbers 191—194 by

The Pan-Pacific Entomologist 50: 60-67. January 1974

Vol. 50, No. 1, January 1974

Figs. 1-2. Callirhytis eldoradensis. Fig. 1. Bisexual generation gall, 5X-

Fig. 2. Unisexual generation gall, 15X-

Houard (1935) were erroneously identified as Andricus attractans Kin-

sey, now in Callirhytis. They do not secrete honeydew as do C. at-

tractans galls. They contain a succulent layer of nutritive tissue around

the larval cell which provides a food and moisture reserve for the larva

after the gall falls to the ground in April. Emergence is during the late

spring or early summer, apparently the next year. One specimen reared

at the U.S. National Museum emerged 27 April 1923 from galls collected

May 1922. Another, reared indoors by the author, emerged prior to 15

June 1971 from galls collected April 1970. None of the adjacent limbs

or trees, including the original source of insects, harbored the galls,

probably due to both previous intensive collecting and inclement

weather. Nor did any other unisexual generation galls occur on the

isolated twigs.

Morphology of the unisexual generation requires transfer of the

species from its previous assignment in Eumayria Ashmead to Calli-

rhytis Foerster group B, which includes almost exclusively species as-

sociated with acorns.

The following additional circumstantial evidence also substantiates

this association of alternate generations. Bisexual generations usually

emerge about two to three months later than the unisexual generation,

which in this case suggests natural emergence and oviposition of the uni-

sexual generation about May or June. By this time the trees are usually

completing seasonal growth and though galls can be induced in some

differentiated nearly mature plant tissue (Lyon, 1970), one of the few

The Pan-Pacific Entomologist

plant tissues still growing sufficiently to be an optimum oviposition

site for this unisexual generation female is the rapidly enlarging acorns.

Callirhytis flora $ 2 , C. milleri £ , and Callirhytis carmelensis Weld

$ are the only other gallmaking cynipids known to be associated with

Pacific Slope Erythrobalanus acorn galls. The first two are alternate

generations of the same species and thus neither could be an alternate

generation of C. eldoradensis. It is highly unlikely that both generations

of a cynipid would occur on a tissue so limited in availability as acorns,

especially since a large brood of C. carmelensis would abort the acorns

needed for a brood of C. eldoradensis. Also though acorns of both Q.

kelloggii and Q. wislizenii require two seasons to mature and might re-

ceive eggs during the first % of their time on the tree and sustain the

bisexual generation of C. eldoradensis , Q. agrifolia matures acorns in

only one season. Thus the typical winter emerging unisexual generations,

such as C. carmelensis, cannot be part of the life cycle of C. eldoradensis

because on Q. agrifolia they would have no acorns available in which

to oviposit. There are no other known spring or early summer emerging

unisexual generations associated with acorns which might potentially

be considered as the alternate generation of C. eldoradensis.

Systematics. — Female. Head: nearly massive, broadly triangular in front view,

widest at antennal socket level, slightly narrower than thorax. Occiput darker,

elevated about % width of ocellus above head contour. Cheeks widened behind

eyes. Malar groove absent, malar area pubescent with fan-shaped striae radiating

from clypeus corners. Antenna 13 segmented, last segment twice the length of the

preceding. Thorax: mesoscutum sparsely pubescent, with fine transverse rugosity.

Notaulices weakly complete, distinct only posteriorly. Anterior parallel lines and

parapsidal lines extending nearly % length of mesoscutum. Median groove polished,

Vs length of mesoscutum. Scutellum coriaceous centrally, reticulate laterally;

foveae reticulate medially, polished smooth laterally. Mesopleuron ridged longi-

tudinally. Tarsal claws simple. Wings: veins pale yellow-brown; faintly clouded,

surface with short pubescence, margin bare. Radial cell open, aerolet % length

of open cubital cell. Abdomen; first tergite polished smooth with dorsally inter-

rupted pubescent ring at base, succeeding tergites polished micropunctate, three

tergites visible dorsally. Ventral spine 12 times as long as wide, sparsely pubescent,

forming 90° angle with ventral valves, equal to length of entire hind tarsus. Color:

amber, abdomen darker.

Type Deposition . — Plesiotype female in U.S. National Museum.

Callirhytis flora Weld

Callirhytis flora, which was described from females (though Weld

also reared males) that emerged from an integral hard midrib leaf

gall (Fig. 3) during May and June, is the alternate bisexual generation

of Callirhytis milleri, new synonym which emerges between late Feb-

Vol. 50, No. 1, January 1974

ruary and mid March from lumpy aborted acorn cotyledons (Fig. 4).

Unisexual generation galls have been illustrated by Beutenmuller (1913)

and Weld (1922b). The former erroneously associated them with Cal-

lirhytis eldoradensis. Bisexual generation galls are also figured by Weld

(1922a, 1957). Both generations belong to Weld’s group B, which in-

cludes primarily unisexual generations from hard acorn galls. C. flora

and C. milleri are hereby synonymized, with Callirhytis flora Weld

having priority.

Biology. — On 20 March 1969 numerous unisexual generation females

placed on Q. wislizenii with rapidly elongating buds readily oviposited

in the midrib of the lower surface of young leaves. Bisexual generations

usually take two or three months to develop, and the only known bi-

sexual group B Callirhytis emerging from a leaf gall on California

Erythrohalanus oaks during May to June is C. flora.

The long ovipositor of C. flora (9.3 X head width) caused Weld to

believe it oviposited in acorns. Such an ovipositor would only be nec-

essary for ovipositing deep in plant tissue, such as the center of a ma-

turing acorn. The senior author collected ten bisexual generation fe-

males of C. flora while they were ovipositing in acorns of Q. agrifolia

and in second year acorns of Q. wislizenii during May and June 1967

to 1969. These acorns drop to the ground in the fall and the unisexual

generation larvae remain in the galls at least IVz years before pupation.

Weld (1922b) has recorded a larval period lasting 2% years.

C. flora and C. milleri key out to the same couplet (Weld, 1922b).

This structural similarity of alternate generations is paralleled by that

of Callirhytis pomiformis (Bassett) (Lyon, 1959). The only other

known Pacific Slope gallmaking cynipids reared from Erythrohalanus

acorn galls are C. eldoradensis and Callirhytis carmelensis Weld. Neither

of these could result from the oviposition of C. flora because of the fol-

lowing reasons. C. eldoradensis from acorns is also a bisexual gen-

eration, and the alternate unisexual generation emerges from a spherical

bud gall. Acorn pip galls of C. carmelensis develop during March and

April and drop to the ground about May. The acorns of Q. kelloggii and

Q. wislizenii , which require two years to mature, could receive eggs

nearly any time during the first or second years and produce galls of

C. carmelensis. However, Q. agrifolia produce mature acorns between

February and October of the same year. Since the bisexual generation of

C. flora emerges in June, any Q. agri folia acorns in which they might

oviposit would drop to the ground that autumn and not remain on the

tree until the following spring. C. flora bisexual generation individuals

The Pan-Pacific Entomologist

Figs. 3-4. Callirhytis flora. Fig. 3. Bisexual generation gall, 2.5 X- Fig. 4.

Unisexual generation gall, 2.5 X-

do not live through the winter until the following spring to be able to

oviposit in the newly forming crop of acorns. C. carmelensis could

therefore not be the alternate generation of C. flora.

Systematics. — Male. Head: massive, not widened behind eyes, malar space

grooved; antennae 15 segmented, third excavated dorsally, last equal to preceding.

Thorax: scutum coriaceous, coarse posteriorly, notaulices complete, anterior par-

allel lines distinct; foveae shallow, ridged, weakly separated; scutellum rugose,

smoothest centrally; mesopleuron smooth dorsally and ventrally, coriaceous med-

ially; wings pubescent, margins ciliate, veins pale beyond second crossvein. Ab-

domen: smooth, bare, brown, lightest dorsally, not pedicellate. Length: 1.7 mm.

Color: frons, genae, antennae, and legs yellow-brown, remainder red-brown.

A unisexual generation male sport which emerged 22 January 1969 differs from

bisexual generation males by its larger size (2.8 mm) ; uniformly brown wing-

veins; last antennal segment slightly longer than preceding; scutum finely rugose

anteriorly, coarser posteriorly; polished coriaceous abdominal tergites.

Type Deposition . — Plesiotype bisexual generation male and unisexual generation

male sport in U.S. National Museum. Plesioparatype males in the Weld collection

in the possession of Robert J. Lyon, Los Angeles City College (3), California

Academy of Sciences (10), senior author’s collection (60), and U. S. National

Museum (5) .

Vol. 50, No. 1, January 1974

Figs. 5-6. Callirhytis perfoveata, bisexual generation galls. Fig. 5. Position

of galls on leaf, 2X- Fig. 6. Detail of gall structure, 7X-

Distribution. — Probably found throughout the range of the host

plants.

Parasite Biology. — Euderus craw fori Peck (Hymenoptera: Eulophi-

dae) adults were dissected from larval cells of bisexual generation C.

flora which contained partly consumed adult gallmakers indicating the

parasites did not attack the gallmakers until the latter completed pu-

pation.

Callirhytis perfoveata (Kinsey)

When Kinsey described the rapidly developing spring cynipid C.

perfoveata from six females (1922) he stated, “such species of cynipids

are usually bisexual, so the male may yet be discovered for the species.”

This has been confirmed with the rearing of males from Quercus wis-

lizenii.

Biology. — The green, nearly spherical, succulent, monothalamous in-

tegral leaf galls (Figs. 5 & 6) were noted by the senior author in the

new growth of Q. wislizenii at Folsom Lake, Placer Co., California, 19

March, 1967. They were fully developed and contained mature larvae.

On 23 March some contained white pupae. Galls collected then and

maintained indoors 65-75° F yielded two females on 1 April. Galls

still on the trees showed no evidence of natural emergence. From galls

collected 15 April both males and females emerged from 20 to 29 April

(indoors). By May over half of the galls examined on the trees had

emergence holes. Sprenger collected C. perfoveata from Q. wislizenii

in Rocklin, Placer Co., California on 13 and 14 April, 1973. Insects

The Pan-Pacific Entomologist

emerged 13 to 27 April while being maintained indoors. Neither the

oviposition site of the females nor the alternate unisexual generation is

known. Galls have also been seen at Lemoncove, Tulare Co., and Clear-

lake, Lake Co., California.

Systematics. — Female. Similar to Kinsey’s 2 description, but with the following

variation: proximal antennal segments yellow-hrown darkening to deep brown

distally; cheeks and malar space reddish-brown to black; mandibles yellow-brown

to reddish-brown; median groove sometimes indistinct. Using maximum width of

the head as a base, the mesonotum length ratio is 1.3, antennae 2.3, wings 3.7.

Length of 48 specimens, 2.1-2.6 mm. Average 2.4 mm.

Male. Head coriaceous; from above transverse, as broad as thorax, not broad-

ened behind eyes; in anterior aspect, interocular space twice as wide as high, malar

space 0.2 eye height without groove; antennae 15 segmented. Mesonotum cori-

aceous, notaulices complete, broader behind, median groove less distinct than in

females; scutellum rugose, smoother medially, fovea smooth and polished, separated

by ridge; mesopleuron nearly bare, more striate ventrally. Wing hyaline, margin

ciliated, veins brown, radial cell open, aerolet % wing length. Claws simple.

Abdomen non-pedicellate, shorter than head plus thorax, tergite II smooth. Using

maximum head width as base, mesonotum length is 1.4, antennae 3.3, wing 4.1.

Length of 27 specimens, 1.8-2.4 mm, average 2.1 mm. Body, head, and antennae

black, abdomen dark brown, lightest dorsolaterally on tergite two.

Type Deposition. — The series contains 27 males and 48 females. Plesioallotype

male in U.S. National Museum. Plesiotype insects of both sexes and freeze dried

galls have been deposited at the U.S. National Museum, California Academy of

Sciences, Weld collection in the possession of Robert J. Lyon of Los Angeles City

College, University of California at Davis, and in the authors’ collections.

Acknowledgments

The authors are grateful to Mr. Robert J. Lyon of Los Angeles City

College for the use of Weld’s collection and field notes, to Dr. B. D.

Burks of the U.S. Natl. Museum for identifying the chalcid parasites,

and to Linda Campbell and Barbara Young for the illustrations.

Literature Cited

Beutenmuller, W. 1913. The North American acorn galls with descriptions of

new species. Bull. Brooklyn Entomol. Soc., 8: 101-105.

Houard, C. 1935. Les collections cecidologiques du laboratoire d’Entomologie du

Museum d’Histoire naturelle de Paris: galles des Etats-Unis. Marcellia,

28: 162-163.

Kinsey, A. C. 1922. New Pacific coast cynipidae. Amer. Mus. Nat. Hist. Bull.,

46: 287-288.

Lyon, R. J. 1959. An alternating sexual generation in the gall wasp Callirhytis

pomiformis (Ashm.). Bull. So. Calif. Acad. Sei., 58: 33-37.

Lyon, R. J. 1970. Heterogony in Callirhytis serricornis (Kinsey) . Proc. Entomol.

Soc. Wash., 72: 176-178.

Vol. 50, No. 1, January 1974

Weld, L. H. 1922a. Notes on cynipid wasps, with descriptions of new North

American species. Proc. U.S. Natl. Mus., 61: 20.

Weld, L. H. 1922b. Notes on American gallflies of the family cynipidae pro-

ducing galls on acorns, with descriptions of new species. Proc. U.S.

Natl. Mus., 61: 1-32.

Weld, L. H. 1957. Cynipid galls of the pacific slope. Privately Printed, Ann

Arbor, Mich., 64 pp.

SCIENTIFIC NOTE

Recovery of Anax junius from a glacier in the Sierra Nevada (Odonata:

Aeschnidae). — On August 16, 1972, while traversing the Lyell Glacier in Yosemite

National Park with Charles Dickson, ranger-naturalist at Tuolumne Meadows, I

discovered several specimens of a dragonfly, Anax junius (Drury), frozen into the

ice at an altitude of 12,200 feet. The winter of 1971-72 had been unusually mild,

resulting in one of the lightest snowpaclcs of this century. As a consequence, many

of the Sierran glaciers retreated, and surfaces of ice deep within them were ex-

posed. These particular dragonflies were undoubtedly swept onto Mount Lyell by

upslope winds to perish on the glacier, where they have remained embedded for

uncounted years.

R. W. Garrison of the Dept, of Entomological Sciences at the University of Cal-

ifornia, Berkeley, who confirmed the identification, also noted that one of the

females was teneral and was therefore probably trapped on the glacier during her

first week of life, possibly on her maiden flight.

Anax junius is a very widespread species, occurring throughout North and Cen-

tral America. During previous summers in the high Sierra, I have taken numerous

specimens in Dana Meadows at Tioga Pass, where it undoubtedly breeds in sub-

alpine lakes there. — R. P. Papp, Department oj Entomological Sciences, University

of California, Berkeley 94720.

The Pan-Pacific Entomologist

Observations on the Behavior of Mallophora fautrix

Osten Sacken

(Diptera: Asilidae)

John Alcock

Department of Zoology, Arizona State University, Tempe 85281

Mallophora fautrix subspecies bromleyi Curran (Cole and Pritchard,

1964) is a robust asilid with a close mimetic resemblance to the bumble-

bee Bombus sonorus Say (see Plate 53, Linsley 1960). Linsley has pro-

vided a brief account of the ethology of this species (under the name

M. bromleyi ) with emphasis on the prey taken. The purpose of this

paper is to offer additional information, largely on the mating and

oviposition behavior of the fly. My observations were made during the

first two weeks of August 1973 at a site near Cave Creek, approximately

2 km east of Portal, Arizona and 1 km north on the road to San Simon.

The flies were abundant in an area of dense vegetation watered by a

small irrigation ditch; the predominant plant was Cortyza coulteri Gray

although scattered cockleburrs, Xanthium saccharatum Wallroth, and

other desert vegetation were also present (see Linsley and Cazier, 1972,

for a description of the locality and its flora).

Mating behavior. — Asilid activity at the site began at about 0900-

0930 Mountain Standard time and continued at a high level until about

1230. The morning was apparently devoted primarily to reproductive

behavior in as much as I never saw a fly with prey during this time

despite the presence of many individuals. Males flew from one perch

to another with brief intervals of rest. In the course of their flights

they frequently hovered with the hindlegs widespread near conspicuous

burrs, large seed heads, and perched individuals of the same species.

They then darted at these objects sometimes striking a plant part re-

peatedly before flying away. They also pursued conspecific individuals

in the air in a noisy, bobbing “roller-coaster” flight which often carried

them 3-4 m high.

Although the onset of copulation was not observed, I did see five

attempted matings (1020-1245) and four coupled pairs (1000-1205).

Males struck and attempted to grasp perched or egg-laying females.

If successful they would fall to the ground holding their would-be mate

firmly. They then probed with their abdomen in an effort to achieve

genital coupling. In the cases of attempted matings that I watched the

male was often oriented with respect to the female in ways that made

The Pan-Pacific Entomologist 50: 68-72. January 1974

Vol. 50, No. 1, January 1974

Fig. 1. Two positions assumed by copulating Mallophora fautrix.

copulation impossible (for example, one male held the female by her

abdomen while probing her head and thorax) and sooner or later the

the pair broke apart. Mating pairs (Fig. 1) were typically observed

clinging to the vegetation in the tail-to-tail position, while disturbed

pairs flew from one area to another with one fly trailing behind the

other.

BEE

0830 1000 1200 1400

Fig. 2. The flight periods of the bumblebee B. sonorus and its mimic, Malloph-

ora fautrix, at the Portal site. Data collected on 10 August (0830-1100 Mountain

Standard Time) and 12 August (1100-1400) 1973. Black boxes indicate that at

least one flying bee or asilid was seen during a five min. interval within five m.

of the observer. On 13 August observations between 1400-1700 produced just

three sightings of a bumblebee and one of the asilid.

The Pan-Pacific Entomologist

Fig. 3. A female Mallophora fautrix in the act of producing an egg case. Note

that the case is constructed of thin coils of white oothecal material.

Flight activity. — The period of considerable flight activity coincided

closely with the time when the fly’s abundant model, B. sonorus, was

also active in the area (Fig. 2) . Whether the synchronous flight periods

reflect a common thermal constraint on energetic flight for both species

or whether selection has favored flies which time their conspicuous

aerial reproductive activities to match the flight period of the model

could not be determined. Perhaps both factors are responsible for the

daily pattern of activity of M. fautrix.

Egg-laying. — Females laid eggs between 1035-1245 (N = 10) . As

Linsley surmised (1960), this species places its egg cluster at the tip

of dried weed stems (especially Conyza coulteri ) at a height of 1-2 m

above the ground. Females first alighted head-up on a suitable site and

then moved about probing with their abdomen, apparently searching

Vol. 50, No. 1, January 1974

for an open area on which to place the egg case. Having located such

a site, the females then curled their abdomen and began to exude a

thin coil of white foam (Fig. 3). By moving their abdominal tip back

and forth in a semi-circle the insects created a half-cylinder with a cavity

which was filled with masses of eggs at short intervals. Egg cases ranged

in size from 0.5-2. 0 cm; females were sometimes interrupted in the midst

of egg-laying by a male (three observations).

Discussion

Lavigne and Holland (1969) have summarized the information avail-

able on the courtship and oviposition behavior of the Asilidae (see also

Lavigne, 1970a, b; Rogers and Lavigne, 1972, for additional details).

Although complex courtship displays have evolved in a few genera of

asilids, most members of this family omit this component of reproduc-

tive behavior. Mallophora fautrix appears to belong with the majority

in this regard although, as noted, males will hover near certain objects

{e.g. burrs) with their hind legs extended. This behavior could con-

ceivably provide a female with a courtship signal. In addition, the func-

tion of the oscillating flights may also be related to courtship although

this remains speculative.

The oviposition behavior observed for this species is unusual for the

Asilidae. Apparently only species of Mallophora and Mallophorina

produce chalky egg cases and place them on open stems. However, a

small number of asilids, including some species of the closely allied

genus Protomachus , oviposit on vegetation, generally in some protected

site, such as the florets of a grass or the space between a grass stem and

leaf (Lavigne and Holland, 1959).

Acknowledgments

Mr. Joe Wilcox identified specimens of the fly for me. Dr. Mont A.

Cazier directed me to the relevant literature and identified the plants

named in the paper. This study was made possible by a faculty Grant-

in-Aid from Arizona State University.

Literature Cited

Cole, F. R. and A. E. Pritchard. 1964. The genus Mallophora and related

asilid genera in North America (Diptera: Asilidae). Univ. Calif. Pubs.

Entomol., 36: 43-93.

Lavigne, R. J. 1970a. Courtship and predatory behavior of Cyrtopogon auratus

and C. glarealis (Diptera: Asilidae). J. Kans. Entomol. Soc., 43: 163-

171.

The Pan-Pacific Entomologist

Lavigne, R. J. 1970b. Courtship and predation behavior of Heteropogon macu-

linervis (Diptera: Asilidae) . J. Kans. Entomol. Soc., 43: 270-273.

Lavigne, R. J. and F. R. Holland. 1969. Comparative behavior of eleven species

of Wyoming robber flies (Diptera: Asilidae). Univ. Wyo. Agr. Exp.

Sta., Sci. Monog., 18: 1—61.

Linsley, E. G. 1960. Ethology of some bee- and wasp-killing robber flies of

southeastern Arizona and western New Mexico (Diptera: Asilidae).

Univ. Calif. Pubs. Entomol., 16: 357-392.

Linsley, E. G. and M. A. Cazier. 1972. Diurnal and seasonal behavior patterns

among adults of Protoxaea gloriosa (Hymenoptera, Oxaeidae) . Amer.

Mus. Novit, 2509: 1-25.

Rogers, L. E. and R. J. Lavigne. 1972. Asilidae of the Pawnee National Grass-

lands in northeastern Colorado. Univ. Wyo. Agr. Exp. Sta., Sci. Monog.,

25: 1-35.

SCIENTIFIC NOTE

Lepturine Longhorns Collected on Chinquapin Bloom (Coleoptera:

Cerambycidae) . — The forested regions of western Oregon are moderately rich

in members of the cerambycid tribe Lepturini. Adults of most species of these

anthophilous beetles appear in the spring and early summer and collectively visit

the blossoms of a wide variety of herbaceous and woody plants. Periods of activity

vary with altitudinal distribution but usually by late July many species and their

host flowers have disappeared. However, there are some lepturines which fly

throughout July and August in the middle elevations of the Cascades, and for

these chinquapin appears to be a favored and important source of pollen. Giant

chinquapin, Chrysolepsis chrysophylla (Dougl. ex Hood.) Hjelmquist, is widely

distributed in the Cascade and Coast ranges and its showy catkins of creamy white,

heavily fragrant flowers are commonly in evidence from July to early September.

Eleven species of longhorns were collected on chinquapin blossoms as follows:

Anoplodera amabilis LeC., A. canadensis Oliv., A. chrysocoma Kirby, A. crassi-

cornis LeC., A. crassipes LeC., A. dehiscens LeC., A. dolorosa LeC., A. tibialis

LeC., Leptura obliterata Hald., L. propinqua Bland, and Stenocorus flavolineatus

(LeC.). Species diversity and abundance varied with locality except for A. cana-

densis and S. flavolineatus which were represented by single specimens. Collection

records include: % mile east of Suttle Lake Junction, Jefferson County, August

1, 1973; three miles east of Santiam Pass, Jefferson County, August 1, 1973 and;

13.2 miles south of Marion Forks, Linn County, August 4, 1973. A. crassicornis

has been found abundantly on chinquapin on Doak Mountain, Klamath County,

along with a few specimens of A. amabilis and A. dehiscens. — Richard L. Penrose

and Richard L. Westcott, Oregon Department of Agriculture, Salem 97310.

Vol. 50, No. 1, January 1974

Unusual Caterpillar-Prey Records and Hunting Behavior

for a Podalonia Digger Wasp: Podalonia valida (Cresson)

(Hymenoptera: Sphecidae)

A. L. Steiner

Department of Zoology, University of Alberta, Edmonton, Alberta, Canada

Observations on Podalonia valida (Cresson) were made in late sum-

mer and fall 1972, in an area immediately adjacent to the Chiricahua

Mountains in extreme southeastern Arizona, about one mile west of the

Chiricahua National Monument entrance, and about 35 miles south of

Willcox, Cochise County.

The study area, described in more detail elsewhere (Steiner, in prep-

aration) , was situated in a riparian habitat along a temporary creek

(Bonita Creek), in the Upper Sonoran/Transition Zones. This oasis-like

area was surrounded by more arid desert scrub and grassland. Within

the study area, a lush herbaceous vegetation, tall trees, and a shrubby

understory were present.

Four female Podalonia valida were observed in the study area be-

tween September 10 and October 9, 1972. Detailed prey records were

made on only one individual. This individual paralyzed and stored in

her nests, large, very hairy caterpillars of the genus Estigmene (Arcti-

idae), exclusively. In seven observed cases, all prey were large hairy

caterpillars ranging from a black color, with rufous-reddish latero-

ventral parts, to brownish-yellowish. All caterpillars were probably

Estigmene acraea (Drury), or very closely related forms.

Predatory Behavior of Caterpillar Hunting Wasps

Lepidopterous larvae of various species, shapes, sizes, colors and

habitats are preyed upon by digger wasps of the genera Ammophila and

Podalonia (and some other genera including some solitary Vespidae).

The majority of these wasps prey on naked caterpillars only (see for

instance Murray, 1940; Muesebeck et al ., 1951; Evans, 1959; Powell,

1964; Krombein, Burks et al., 1967, for Nearctic species, and Berland,

1925; Roth, 1928; Tsuneki, 1968, for Palearctic species). A few rec-

ords of odd prey species are available, however. Thus according to Ap-

pleton (in Murray, 1940), Podalonia occidentalis Murray preys on the

tent-caterpillar. Ammophila pilosa pilosa (Fernald) (= azteca Cam-

eron) was found to store a Lycaenid larva, about 14 mm length, densely

covered with short secondary setae, and also a Pterophorid larva (about

The Pan-Pacific Entomologist 50: 73-77. January 1974

The Pan-Pacific Entomologist

15.5 mm length), with numerous tufts of elongate secondary setae;

these caterpillars have rather concealed feeding habits (Powell, 1964) .

Neither Lycaenids nor Pterophorids seem to have been reported as prey

of other North American Ammophila (Powell, 1964). According to

Ferton (1901, 1914), the Palearctic Podalonia hirsuta (Scopoli) (var.

mervensis Radoszkowsky : “Ammophila ebenina” Costa) takes naked

prey such as Agrotis , Cucullia chamomillae, but also hairy caterpillars,

for instance Epinephele jurtina (Berland, 1925).

A number of Podalonia species of known biology prey on Agrotidae-

Noctuidae (cutworms), sheltered underground during daytime, e.g.,

Podalonia robusta (Cresson) : Evans, 1963; luctuosa and/or communis

(Smith) : Hicks, 1931, 1932, for Nearctic species — hirsuta (Scopoli) :

Fabre, 1879, 1882; affinis (Kirby) : Marchal, 1892; tydei (Le Guillou) :

Fabre, 1879; Picard, 1903; atrocyanea Eversmann ( = chalybea Kohl),

microhirta Kohl and/or caucasica Morawitz: Tsuneki, 1968, for Pale-

arctic species). The wasps excavate these nocturnal prey from their

underground shelters.

Podalonia valida , in contrast, hunts diurnal lepidopterous larvae in

the open, usually on dense herbaceous or low brushy vegetation such

as white horsemint ( Monarda pectinata ) , Helianthus (Compositae) , and

goldweed ( Verbesina encelioides) . In the area considered, these lepi-

dopterous larvae were found to move from one plant species to another

quite extensively. Whether this was related to polyphagous feeding be-

havior or to escape behavior, in response to disturbances by hunting

wasps, was not determined, but captive specimens readily accepted some

plants of the sunflower family. Podalonia valida was capable of follow-

ing with great accuracy the fresh scent trails left by the caterpillars, while

moving on the ground from one plant to another. The wasp proceeded

head down, tapping her antennae on the ground, especially where the

caterpillar had stopped and rested for some time (mostly at the bases

of plants), or had made a sudden and sharp change in direction. This

ground level search behavior often led the wasp to the base of one plant,

where a concentrated search would be undertaken, the excitation of the

wasp increasing steadily in the process. There is little doubt that the

wasp was picking up more and more fresh olfactory evidence of a re-

cent occupancy of the site by the prey. Often, the wasp would then climb

up the plant and investigate it thoroughly. At other times the wasp

would fly directly onto the plants, without first searching the ground.

Discovery of the caterpillar did not necessarily follow, however, even if

the wasp came very close to the prey. Such failure to capture the prey

Vol. 50, No. 1, January 1974

Fig. 1 . Podalonia valida carrying a large, paralyzed lepidopterous larva ( Estig -

mene acraea ) (Arctiidae), using her mandibles and fore legs to hold the prey.

Locomotion, with two pairs of legs only, is assisted by wing beats.

was particularly frequent at early stages of hunting phases. This sug-

gests that “motivational” deficiencies, rather than deficiencies in the

ability to detect or locate the prey, were then involved. Such incomplete

behavior is very frequent at early stages of the different phases of the

nesting cycle in digger wasps (Steiner, 1962). These abortive activities

are particularly conspicuous during the nest-digging phase, resulting

in one or several incomplete burrows abandoned at once. Such trun-

cated behavior also occurs at early stages during other phases of the

nesting cycle, including the hunting and prey-stinging stages (Steiner,

1962), and several aborted pursuits, attacks or stinging of prey by

Podalonia valida were recorded in this study. In these cases the wasp

did not make any further attempt at pursuing the escaping prey. At

such early stages, in contrast to later stages of the hunting phase, the

wasp would not usually accept a suitable prey, even if dropped right

in front of her. Such apparent unwillingness to capture a detected prey

has also been reported from other wasps, including other Podalonia

species. For example observations made on Podalonia luctuosa (and/

or communis ) (Newcomer, 1930; Hicks, 1931, 1932, in Murray, 1940),

which preys on cutworms (Noctuidae), concealed beneath the soil by

day, revealed abortive termination of digging behavior during hunting.

No evidence of hunting by sight was found in P. valida , except per-

haps at very close range. Little information on prey stinging was ob-

The Pan-Pacific Entomologist

tained, since this occurred mainly in very dense vegetation and was ac-

complished within a very short period of time. The heavy, cumbersome

prey is straddled and carried over the ground in typical Podalonia

fashion, using the mandibles and also the front legs, which embrace the

larva (Fig. 1). Wing beats provide some additional propulsive force.

A more detailed account of the biology and behavior of this Podalonia

species will be presented elsewhere.

Acknowledgments

I wish to express my gratitude to the Directors, staff and guests of the

Southwestern Research Station, Portal, Arizona; of The American Mu-

seum of Natural History (New York) ; and of the Chiricahua National

Monument, Arizona, for their hospitality, facilities provided, help and

useful suggestions at various stages of this project. The wasp specimens

were identified by R. M. Bohart, University of California, Davis, and

the prey specimens by D. M. Weisman, U.S.D.A., Agriculture Research

Center, Beltsville, Maryland (R. H. Foote, Chief Systematic Entomology

Eaboratory). Some of the plants were identified by S. Mouat. Special

thanks are due to B. and E. Erickson and L. Riggs, for kindly giving

me free access to their property and ranch. This work was supported

in part by NRC grant A-3499.

Literature Cited

Berland, L. 1925. Faune de France — Hymenopteres vespiformes. I (Sphegidae,

Pompilidae, Scoliidae, Sapygidae, Mutillidae) . Lechevalier, Paris.

364 pp.

Evans, H. E. 1959. Observations on the nesting behavior of digger wasps of the

genus Ammophila. Amer. Midi. Nat., 62: 449-473.

Evans, H. E. 1963. Notes on the prey and nesting behavior of some solitary

wasps of Jackson Hole, Wyoming. Entomol. News, 74(9): 233-239.

Fabre, J. H. 1879. Souvenirs entomologiques. Paris.

Fabre, J. H. 1882. Nouveaux souvenirs entomologiques. Paris.

Ferton, C. 1901. Notes detachees sur l’instinct des Hymenopteres melliferes et

ravisseurs. Ann. Soc. entomol. Fr., 70: 83-148.

Ferton, C. 1914. Notes detachees..., 8e serie. Ann. Soc. entomol. Fr., 83: 81-

118.

Hicks, C. H. 1931a. On the digger wasp, Podalonia luctuosa (Smith). Pan-

Pacif. Entomol., 8: 49-51.

Hicks, C. H. 19316. The hunt and capture of the prey of a digger wasp. Bull.

South. Calif. Acad. Sci., 30: 75-82.

Hicks, C. H. 1932. Notes on the prey and inquilines of Podalonia violaceipennis

form luctuosa (F. Smith). Psyche, 39: 150-154.

Krombein, K. V., B. D. Burks et al. 1967. Hymenoptera of America North of

Vol. 50, No. 1, January 1974

Mexico. Second supplement. U.S. Department of Agriculture, Washing-

ton, 584 pp.

Marchal, P. 1892. Notes sur la vie et les moeurs des Insectes. Observations sur

Y Ammophila affinis Kirby. Arch. Zool. exper., 23-36.

Muesebeck, C. F. W., K. V. Krombein, H. K. Townes et al. 1951. Hymenop-

tera of America north of Mexico — Synoptic Catalog. U.S. Dept. Agric.

(Agriculture Monograph No. 2), Washington. 1420 pp.

Murray, W. D. 1940. Podalonia (Hym. : Sphecidae) of North and Central

America. Entomol. Amer. (n.s.), 20: 1-82.

Newcomer, E. J. 1930. Notes on the habits of a digger wasp and its inquiline

flies. Ann. Entomol. Soc. Amer., 23: 552-563.

Picard, F. 1903. Moeurs de YAmmophila tydei Guill. Feuille des jeunes natural-

istes, 34: 15-17.

Powell, J. A. 1964. Additions to the knowledge of the nesting behavior of North

American Ammophila (Hym.: Sphecidae). Jour. Kans. Entomol. Soc.,

37: 240-258.

Roth, P. 1928. Les Ammophiles de l’Afrique du nord. Ann. Soc. entomol. Fr.,

97: 153-240.

Steiner, A. L. 1962. Etude du comportement predateur d’un Hymenoptere

Sphegien: Liris nigra V.d.L. ( — Notogonia pompiliformis Panz.). Ann.

Sci. Nat. Zool. Biol. Anim. Ser. 12, 4: 1-126.

Tsuneki, K. 1968. The biology of Ammophila in East Asia (Hym., Sphecidae).

Etizenia, 33: 1-64.

The Pan-Pacific Entomologist

The Larval Nutrition of Minettia jlaveola and

Phaonia parviceps and its Significance to

the Hawaiian Leaf -breeding Drosophila

(Diptera: Lauxaniidae, Muscidae, Drosophilidae)

J. F. McDonald, W. B. Heed 1 and Mary Miranda

Departments of Genetics and Food Sciences and Technology,

University of California, Davis, California 95616

The purpose of the present report is to point out the similarities and

differences in the larval ecology and nutrition of several species in the

two Dipteran families, Lauxaniidae and Drosophilidae. The habit of

mining in decaying detached leaves by representatives of these two fam-

ilies presents an unusual opportunity to study the dynamics of host plant

selection and related phenomena such as competition and nutrition. The

leaf-breeding habit in the Drosophilidae is known only in the Hawaiian

Islands and is universal in the genus Antopocerus and three related

groups of species within the genus Drosophila (Heed 1968). Species

in other groups and in the genus Scaptomyza also breed in fermenting

leaves and the habit appears to have evolved independently several

times. In total there are approximately 80 to 100 leaf-breeding species

endemic to the Islands. Holarctic members of the Lauxaniidae have

long been known to occupy this niche (Hering, 1951; Oldroyd, 1964).

However until the present report no one has compared larval nutrition

in the drosophilids and the lauxaniids. The overall similarity of the

rotting leaf niche available for the two groups is so striking that a de-

tailed account of recent observations and tests is given below.

The breeding biology of the Hawaiian leaf-breeders has been com-

pared previously (Heed 1971) to the unusual biology of Rhagoletis

(Tephritidae) , as observed by Bush (1969).

Results and Discussion

On October 10, 1972, eggs and first-instar Dipterous larvae were dis-

covered in a redwood forest by one of us (WBH) at Prairie Creek Camp-

ground, north of Orick, Humboldt County, California, on the surface

of fermenting leaves littering the ground beneath a California Bay tree

( Umhellularia californica (H. & A.) Nutt.). The majority of eggs and

larvae were acalyptrate but 1 predatory larva and an egg of a species

of Muscidae were also noted. The leaves were stored on moist sand in

1 Permanent address : Department of Biological Sciences, University of Arizona, Tucson, Arizona

85721.

The Pan-Pacific Entomologist 50: 78-82. January 1974

Vol. 50, No. 1, January 1974

jars at 68° F at U. C. Davis, and adults were first noticed on 24 Jan-

uary, 1973. They were identified as Minettia flaveola (Coquillet) (Laux-

aniidae) . Many puparia and puparial cases were found within the

leaves, but only 3 puparia were washed from the sand. On 28 December,

1972, another collection was made at the same tree and large third-instar

larvae were observed mining within the leaves. On 21 June, 1973, no

larvae were found and the leaves were mostly dry.

Three other collection localities produced fermenting Bay leaves that

contained larvae: (1) along the Carmel River at Carmel Valley, Mon-

terey County, 25 November, 1972; (2) McDonald Ranch at Pope Val-

ley, Napa County, 22 January and 5 February, 1973; and (3) 5 miles

west of Lake Berryessa on State Route 128, Napa County, 1 February,

1973. In addition to M. flaveola, the Napa County samples produced

another lauxaniid which proved to be a new species of Homoneura.

Predaceous larvae were observed in all of the collecting sites, and one

was seen to attack and consume a lauxaniid larva in the latter McDonald

Ranch sample. The adult of this larva was reared and identified as

Phaonia parviceps Malloch (Muscidae). The larva of P. keilini, has

been reported to prey on mosquito larvae (Oldroyd 1964).

The Prairie Creek collection in October was assayed for yeasts. Sam-

ples were taken from the surface of 3 leaves (2 of which had tiny larvae

on the surface) and innoculated on acid-malt- agar plates in the field by

direct plating and by previous dilution in 2 ml. sterile water (Table I).

Five of the species were represented by less than 10 colonies each in all

the plates. Rhodotorula minuta var. minuta was abundant (1000 col-

onies) on 1 leaf and Torulopsis fujisanensis was common on 2 leaves

(100 colonies each). Most of the species are considered to be wide-

spread and all are nonfermentative. T . fujisanensis, however, has not

been reported from the Pacific Northwest (Phaff et al. 1972). In ad-

dition to the yeasts, bacteria and mold colonies were abundant on the

original plates.

The collection from Carmel Valley was assayed for yeasts from the

somewhat fluid interior of the fermenting leaves. Five leaves, 4 with a

large larva in each, were sampled by direct plating and by the addition

of sterile water in the leaf. All slants were made on acid-malt- agar plates.

Only 28 yeast colonies appeared among many mold colonies in 9 plates.

Twenty-four of the colonies originated from the leaf with no larvae

(Table 1). Two leaves produced 2 colonies each and the two remaining

leaves produced no yeast at all.

The significance of these observations and tests lies in the similarities

The Pan-Pacific Entomologist

Table 1. Yeasts isolated from fermenting leaves of the California

Bay Tree.

Species

Total no.

colonies

California

localities

Yeast

niche

Cryptococcus albidus (Saito) Skinner

var. diffluens (Zach) Phaff et Fell

Prairie Creek

Surface

Cryptococcus laurentii var. laurentii

(Kufferath) Skinner

(3 leaves)

Leucosporidium scottii Fell, Statzell,

Hunter et Phaff

Rhodotorula minuta var. minuta

(Saito) Harrison

1000 1

Torulopsis fujisanensis Soneda

230 2

1 1

Candida moscorum di Menna

Candida buffoni (Ramirez)

Van Uden et Buckley

Pink colonies (lost)

43 s

Cryptococcus albidus var. diffluens

Carmel Valley

Interior

Cryptococcus infirms-miniatus

(Okumuki) Phaff et Fell

26 4

(5 leaves)

Debaryomyces hansenii (Zopf)

Lodder et Kreger-van Rij

1 1 leaf.

2 2 leaves.

3 Probable identification : Cryptococcus infirms-miniatus.

i 24 colonies from one leaf, no larvae; 2 colonies from feces of single larva in a second leaf.

and differences they exhibit with the identical larval leaf niche in the

Hawaiian Archipelago which is occupied by a large number of endemic

species in the genus Drosophila (Drosophilidae) (Heed, 1968; Kircher,

1969; Kircher and Heed, 1970; Carson et al., 1970). There are 4 major

similarities between the 2 regions. (1) Both the coastal region in north-

ern California and the windward slopes in the Hawaiian Islands are

belts of equable temperatures and high humidity for a major portion

of the year. This is important for the constancy of the resource (ever-

green foliage) and the slow soft-rot decay of leaves by mold and bac-

teria. (2) The fresh leaf of the Bay tree has a familiar aromatic fra-

grance when crushed. When the leaf is fermenting on the ground its

scent is prevalent in a somewhat modified form. All of the leaves utilized

by Drosophila larvae in Hawaii such as Cheirodendron, T etraplasandra ,

Vol. 50, No. 1, January 1974

Ilex, Clermontia, Pelea, Myoporum , Myrsine and others have their own

characteristic aroma produced by the breakdown products and their

essential oils. These distinctive scents are believed to play a major role

in the very high host plant specificity observed in the Hawaiian leaf

breeders and probably are also important to the lauxaniids discussed

here. Mr. Ray Miller (personal communication) relates that 7 species

in 5 genera of the Lauxaniidae have been reported in rotting leaves.

Dr. B. A. Foote, Kent State Univ., and R. Miller have reared species of

the genera H omoneura, Lyciella and Minettia from decaying leaves of

sugar maple, sassafras, alder, elm, and wild black cherry in deciduous

forests in Ohio (personal communication). (3) The nutrition of the

Hawaiian leaf-breeding larvae is known to be based chiefly on bacteria

and broken down plant parts for the reason that no yeast of any conse-

quence has been isolated from the interior of the leaves (Robertson et al.

1968). Also the leaf-breeding species are notoriously difficult to raise

on laboratory media containing yeasts. However, a variety of yeasts

have been isolated from the rotting leaf exterior in Hawaii (H. J. Phaff

and M. W. Miller, personal communication). The observations in the

present report are similar to the extent that the interior of the rotting

leaves significantly lack the variety and density of the yeasts found on

the leaf surfaces. Hering (1951) considers that the lauxaniids which

are found in rotting leaves are modified leaf miners that had a sa-

prophagous origin. (4) Both fly-substrate systems in California and

Hawaii support larval predators in the family Muscidae. In Hawaii the

genus is Lispocephala (Heed 1968) .

One of the major differences between the lauxaniids and the drosoph-

ilids is that the former oviposit on the leaf surface and the first-instar

larva presumably develops there for a while before entering the leaf.

Up to this time, the major nutritional source could come from yeasts.

The Hawaiian Drosophila insert the eggs into the mesophyll of the leaf

with only the filaments emerging (Kambysellis and Heed 1971).

Another difference is that pupation takes place within the leaf for Mi-

nettia and Homoneura but in Drosophila it occurs several inches below

the soil surface. Finally, Drosophila apparently do not use the leaf niche

in North America (Carson 1971). Presumably the absence of native

lauxaniids in Hawaii (Hardy personal communication) may have been

a contributing factor for the occupation of this niche by Drosophila.

Acknowledgments

We wish to express our appreciation to the following investigators

for the identification of specimens in the Lauxaniidae and Muscidae:

The Pan-Pacific Entomologist

Marius Wasbauer, California Department of Agriculture; Ray Miller,

Iowa State University; R. H. Foote and R. J. Gagne, U. S. Department

of Agriculture. Professor Herman J. Phaff verified the identification

of the yeasts.

The second author wishes to express his appreciation to Drs. Herman

J. Phaff, Martin W. Miller, Herman T. Spieth and Th. Dobzhansky for

the use of their laboratories during his sabbatical leave at U. C. Davis.

This study is supported in part by NSF Grant GB-28953X1 to W. B. H.

and H. W. Kircher and in part by NIH Trainee Grant STOIGM00701-13

to J. F. M.

Literature Cited

Bush, G. L. 1969. Sympatric host race formation and speciation in frugivorous

flies of the genus Rhagoletis (Diptera, Tephritidae) . Evolution, 23: 237-

251.

Carson, H. L. 1971. The ecology of Drosophila breeding sites. Univ. Hawaii

Harold L. Lyon Arboretum Lecture, 2: 1-28.

Carson, H. L., D. E. Hardy, H. T. Spieth, and W. S. Stone. 1970. The evolu-

tionary biology of the Hawaiian Drosophilidae. In Essays in Evolution

and Genetics in Honor of Theodosius Dobzhansky, M. K. Hecht, W. C.

Steere, eds. Appleton-Century-Crofts, New York, pp. 437-543.

Heed, W. B. 1968. Ecology of the Hawaiian Drosophilidae. In Studies in Genetics

IV. M. R. Wheeler, ed. Univ. Texas Publ., 6818, pp. 387-420.

Heed, W. B. 1971. Host plant specificity and speciation in Hawaiian Drosophila.

Taxon, 20: 115-121.

Hering, M. E. 1951. Biology of the Leaf Miners. N. V. Drukkerij Hooiberg,

EPE, Netherlands. 420 pp.

Kambysellis, M. P., and W. B. Heed. 1971. Studies of oogenesis in natural

populations of Drosophilidae. I. Relation of ovarian development and

ecological habitats of the Hawaiian species. Am. Natur., 105: 31-49.

Kircher, H. W. 1969. Sterols in the leaves of the Cheirodendron guadichaudii

tree and their relationship to Hawaiian Drosophila ecology, J. Insect.

Physiol., 13: 1869-1874.

Kircher, H. W., and W. B. Heed. 1970. Phytochemistry and host plant speci-

ficity in Drosophila. In Recent Advances in Phytochemistry III, C.

Steelink and V. C. Runeckles, eds. Appleton-Century-Crofts, New York,

pp. 191-209.

Oldroyd, H. 1964. The Natural History of Flies. W. W. Norton & Co., Inc.,

New York. 324 pp.

Phaff, H. J., M. W. Miller, M. Yoneyama, and M. Soneda. 1972. A compara-

tive study of the yeast florae associated with trees on the Japanese Is-

lands and on the West Coast of North America. Proc. IV IFS: Ferment.

Technol. Today: 759-774.

Robertson, F. W., M. Shook, G. Takei, and M. Gaines. 1968. Observations on

the biology and nutrition of Drosophila disticha Hardy, an indigenous

Hawaiian species. In Studies in Genetics IV. M. R. Wheeler, ed. Univ.

Texas Publ. 6818, pp. 279-299.

Vol. 50, No. 1, January 1974

BOOK REVIEW

The Bark and Ambrosia Beetles of California (Coleoptera: Scolytidae and

Platypodidae) . D. E. Bright, Jr. and R. W. Stark. 1973 Bull. Calif. Insect

Survey, 16: 1-169. University of California Press. $8.50.

Curtis G. Benjamin, in an article in Science (1973, 183(4122) : 282-284) has

lucidly examined the economic afflictions of scientific monographs. Since 1957,

per page prices have risen from 2.50 to 6.10, an increase of 144 percent. Simul-

taneously, average sales of such works declined from about 5000 to 3000 copies.

The present treatise on bark and ambrosia beetles provides a case in point — for a

paperback volume of 175 pages, $8.50 will be considered expensive by most people.

More unfortunately, even at the currently inflated rates, publishers expect such

short-run issues to be economic failures. (The average run for California Insect

Survey Bulletins is about 575.)

Economic matters aside, the Bark and Ambrosia Beetles of California is a valu-

able compilation of biological and distributional information. An introductory

review of taxonomic characters is helpful in working the keys. Summaries of the

distribution, bionomics and the ecological and economic importance of bark beetles

provide a general overview of the detailed information in the species treatments.

A useful adjunct to the text and index is a list of host plants and the scolytids

attacking them.

Keys to the genera and species are well constructed, and non-specialists should

be able to use them proficiently. In a minority of couplets overlapping size ranges

are used as primary characters, but are almost always combined with more easily

distinguished secondary characters. I had little difficulty in correctly keying five

species from different genera.

The bulk of the text consists of summaries of the geographic distribution and

host ranges for each species, with discussions of the biology when that information

is available. For the more important economic species these biological treatments

may be rather extensive, marking an important digression from most earlier Survey

Bulletins, which consisted largely of keys and distributional records. If the present

format is continued, the Bulletin Series should enjoy more general interest than

in the past.

One rather disconcerting feature is the large number of distribution maps (93),

which occupy nearly 25 percent of the text space devoted to species treatments.

Since the majority of species feed on conifers whose ranges in California are sim-

ilar, there is much repitition of distributional patterns. On several of the maps the

symbols are very difficult to discern, and on a few others as few as four to seven

collection records are indicated.

Illustrations include excellent photographs of typical egg galleries of at least

one species of most of the genera treated. Probably of less utility is the page of

line drawings of antennal clubs and funicles which is referred to in the key. My

impression was that the antennal characteristics were frequently clear without visual

aids, while some of the other key characters were less well defined.

On the whole these criticisms are minor compared to the general completeness of

the work, which will certainly be valuable to those dealing with forest insects in

California. At $8.50 it may even be a bargain if the relative price of scientific

monographs continues to inflate at an exponential rate. — John T. Doyen, University

of California, Berkeley.

The Pan-Pacific Entomologist

SCIENTIFIC NOTES

Martarega hondurensis and Buenoa antigone as predators of mosquito

larvae in Costa Rica (Hemiptera: Notonectidae). — Buenoa antigone Kirkaldy

inhabits freshwater pools in southern United States, Mexico, Central and South

America (Truxal, 1953 Univ. Kansas Sci. Bull. 35: 1351-1523). Martarega

hondurensis Bare has been recorded from Mexico, Central America and Peru

(Truxal, 1949, J. Kansas Entomol. Soc. 22: 1-24) and is found aggregated into

schools in deep sheltered eddies of rivers, as is M. mexicana Truxal (Menke and

Truxal, 1966 Los Angeles Co. Mus. Contr. Sci. 106: 1-6). Under certain circum-

stances both species prey on mosquito larvae, and one shows potential as a

biological control agent.

In July, 1972, I surveyed the fauna of two dozen small water-filled depressions

(2m 2 or less) in rock outcroppings along the banks of the Corobici and Tenorio

Rivers, 5 km from Canas, Guanacaste, Costa Rica. Other than notonectids, these

pools seemed to lack potential mosquito predators. Six pools which contained

Buenoa (predominately B. antigone ) lacked mosquito larvae, while the others

containing no Buenoa had large aggregations of mosquito larvae.

At each pool I recorded a number of habitat parameters (size of the pool, depth

and character of the bottom sediments, water turbidity, temperature, pH, vegeta-

tion, amount of shade, water depth and presence of other aquatic organisms) .

The distribution of backswimmers and mosquito larvae was similar with respect

to these parameters, the only difference between habitats being the presence of

the backswimmers and mosquito larvae themselves. Whether the distribution of

Buenoa in pools is a fortuitous event or is dependent upon some undetermined

factor remains unclear, but it seems probable that the absence of mosquito larvae

in these pools is related to the presence of Buenoa.

In Santa Rosa National Park, Guanacaste, Costa Rica, Buenoa were especially

abundant and were never found together with mosquito larvae in several large

ponds that were surveyed. Other potential mosquito predators in these ponds

may have reduced numbers of larvae, but the high density of Buenoa suggests that

the backswimmers may have had a significant part in mosquito elimination.

Laboratory experiments tended to confirm the predator-prey relationship sug-

gested by field observations. B. antigone were collected and kept in jars (110 ml)

for 24 hours at air temperature. Each jar contained two B. antigone as well as

five mosquito late instars. In twelve repetitions (six late instars, six adults) all

mosquito larvae offered were eaten. Martarega hondurensis from river sites ad-

jacent to the rock outcroppings where B. antigone occurred were offered mosquito

larvae in the same format as used for Buenoa. After 24 hours adult Martarega

had consumed an average of 2.6 of 5 larvae offered in each of 33 repetitions, while

late instars ate an average of 1.4 of 5 in each of 39 repetitions. Clearly, under

these conditions, M. hondurensis ate mosquitoes at a slower rate than did B.

antigone.

Martarega seemed more inclined to cannibalism than to mosquito predation.

In 3.5 liter glass aquaria containing from 15 to 20 M. hondurensis, 15 to 20

mosquito larvae were offered but not eaten, but earlier instar M. hondurensis were

often cannibalized. Under similar conditions B. antigone rapidly ate mosquito

larvae, with no cannibalism.

Prey preference tests showed that B. antigone ate mosquito larvae and water

boatmen before ostracods, blood worms, mosquito pupae and ants (P ^ .05)

Vol. 50, No. 1, January 1974

(Gittelman, in preparation). Probably Baenoa are capable of subsisting on a

variety of prey, allowing them to be effective predators on mosquito larvae while

not totally depending on them for their distribution. When offered a similar

selection of prey types, M. hondurensis ate ants struggling at the water surface

before all other prey species including mosquito larvae (P^.01).

B. antigone is readily maintained in the laboratory but breeding under laboratory

conditions has not been attempted. It is likely that these insects lay their eggs on or

in plant material. In either case, eggs could be collected by harvesting plants or

some artificial oviposition substrate (Toth and Chew, 1972b). Since B. antigone

does not cannibalize even in crowded laboratory enclosures, little difficulty would

be expected in mass rearing. In contrast, M. hondurensis would necessitate separate

containers, for each individual.

The suggestion that backswimmers be used for mosquito control (Toth and

Chew, 1972, Ann. Entomol. Soc. Amer. 65: 1270-79; 1972. Env. Entomol. 1:

534-5; Ellis and Borden, 1970, Ann. Entomol. Soc. Amer. 63: 963-73) seems well

applied to B. antigone but not to M. hondurensis. Although both species will eat

mosquito larvae, M. hondurensis do so at a slower rate than do B. antigone. In

addition it would appear impractical to distribute M. hondurensis as a biological

control agent because it tends to inhabit moving water where mosquito larvae

are not prevalent. On the other hand, B. antigone seem to prefer mosquito larvae

as food, and their microhabitats generally overlap.- — -Steven H. Gittelman,

Division of Biological Sciences, U-42, University of Connecticut, Storrs 06268.

Biological observations on darkling ground beetles from western North

America (Coleoptera: Tentyriidae). — Coelosattus fortineri Blaisdell (1927)

was originally described from the Algodones Sand Dunes of California; this genus

was placed in synonymy under Eusattus (Doyen, 1972: Quaest. Ent., 8: 357-376).

Coelomorpha maritima Casey (1890) was originally described from “lower Cali-

fornia.” This genus was later placed in synonymy under Coelus (Doyen, 1972).

Subsequent to their original descriptions, these beetles have remained largely un-

known in collections, although both are locally abundant.

Seventy-six individuals of Eusattus fortineri were collected from the Algodones

Dunes, 2 mi. N. Glamis, Imperial County, California, during April, 1972. Four

of these were encountered on the sand surface between 1900-2300 hrs. (PST).

The remainder were excavated from the sand around the bases of creosote bush

( Larrea divaricata Cavanilles) , at depths of 6-20 cm. The distribution of the

beetles was highly aggregated, with 28 individuals buried beneath a single plant.

A typical assemblage consisted of six to eight beetles, and only three groups of

less than three individuals were discovered. Beetles were found only beneath

plants with canopies close to the sand. Suitability of plants as shelters depends

on the degree of shading of the sand about their bases, since exposed substrates in

the Algodones Dunes reach midday temperatures well above 40°C during warm

months.

E. fortineri appears to be abundant within its restricted geographic range,

which coincides with the distribution of exceptionally fine sand composing the

Algodones Dunes and adjacent dunes in southwestern Arizona and northwestern

Mexico. Its allopatric counterpart in other areas of western North America is

the morphologically similar E. muricatus LeConte, which occurs on a variety of

The Pan-Pacific Entomologist

sandy substrates, including dunes, over a broad geographic area from eastern

Washington to Texas and northern Mexico. It has been recorded from numerous

localities within 50 mi. of the Algodones Dunes, but never in actual sympatry with

E. fortineri.

Coelus maritima has been recorded from scattered coastal localities in northern

and central Baja California, but fewer than 30 specimens were known prior to

1972. During late March, 1972 and 1973, and mid-June, 1973, approximately 500

adults of C. maritima were collected by screening sand at the maritime dunes at

Santa Maria Beach, San Quintin Bay, Baja California Norte. In March both

adults and larvae were extremely abundant to depths of 3 cm in the sand

canopied by perennial plants, especially Abronia maritima Nuttal ex Watson,

where they were associated with small numbers of other tenebrionids Cryptadius

inflatus LeConte, Eusattus laevis LeConte, E. erosus Horn, and Micromes ovipennis

(Horn). In June only adults were encountered, many teneral, suggesting that

adult emergence peaks in late spring or early summer.

The local abundance of C. maritima is probably conditioned by the characteristics

of the sand in which it lives. The beach sand at San Quintin Bay is extremely

fine and friable, allowing entry by small organisms such as Coelus. South of San

Quintin, the dunes extend inland a distance of 3-4 miles, but the sand becomes

progressively coarser and more consolidated inland. Collections away from the

immediate coast contained no Coelus, but the larger species, E. laevis was present,

and Micromes was abundant on the sand surface beneath plants. In extreme

northern Baja California, C. maritima is replaced by C. globosus LeConte, which

occurs in the same habitat. In extensive collections from dunes near Cantamar,

25 mi. S. Tijuana, C. globosus was very abundant, while C. maritima was absent,

and the two species do not appear to occur sympatrically. John T. Doyen, Division

of Entomology and Parasitology, University of California, Berkeley 94720.

Differential predation of darkling ground beetles (Coleoptera: Tene-

brionidae) by the Channel Islands Fox. — During April, 1972, 73 scats of the

Channel Islands Fox ( Urocygon littoralis) were collected from the sand dunes near

Wilson’s Cove on the northwestern end of San Clemente Island, Los Angeles

County, California. The diet of the fox is known to be extremely varied, including

berries, acorns, and carrion, as well as living invertebrates and small vertebrates

(von Blocker, 1967, in Philbrick, Proceedings of the Symposium on the Biology of

the California Islands, pp. 245-253) . All of these items were represented by the

fragments recovered from the San Clemente scats, but insect remains were partic-

ularly abundant. What is of interest is the relative numbers of some of the beetle

prey (Table 1). By far the most abundant were corpses of Trigonoscuta sp. (Cur-

culionidae), June beetles (Scarabaeidae) , and the tenebrionids Coelus remotus Fall

and Eusattus robustus LeConte. All these species are relatively abundant on or

about the sand dunes, but two of them ( Trigonoscuta and Coelus) are relatively

small bodied. Large numbers would be required to sustain an animal as large as

a fox. The large tenebrionid Eleodes laticollis LeConte is also abundant at Wil-

son’s Cove, but was rare in the scats. Three other small tenebrionids, Apsena

grossa LeConte, Coniontis latus LeConte, and Helops bachei LeConte were also

uncommon in the scats, though relatively abundant on or about the dunes. With

Vol. 50, No. 1 , January 1974

Table 1. Beetle contents of 73 scats of the Channel Islands fox. The left column

of figures lists the numbers of scats containing each species. The right column lists

total numbers found in all scats combined. Numbers of tenebrionids and of Trigo-

noscuta were estimated by counting prothoraces, which usually remained intact.

Numbers of Scarabaeidae were estimated by counting elytra.

Scats

Total number of

individuals

Tenebrionidae

Coelus remotus Fall

196

Coniontis latus LeConte

Eusattus robustus LeConte

Apsena grossa LeConte

Eleodes laticollis LeConte

Helops bachei LeConte

Scarabaeidae ( Phobetus and Serica )

Curculionidae ( Trigonoscuta )

270

the exception of Coniontis, which was the least numerous of the beetles in the

habitat, the tenebrionid species which were infrequently eaten possess quinonoid

secretions which they release at the abdominal tip when disturbed. Eleodes lati-

collis is particularly well supplied with secretions, which are stored in large res-

ervoirs in the abdomen and forcibly ejected as a fine spray which may travel over

30 cm. Coelus, Coniontis, and Eusattus, as well as the weevil and scarabs, lack

secretions. Although quinonoid secretions of tenebrionids are popularly referred

to as defensive secretions (Eisner and Blumberg, 1959, An. Rec., 134: 558-559),

little evidence of differential predation is available. The data presented here sug-

gest that species possessing secretions are ignored, at least when other food is

present. Quantitative estimates of the population densities of the various species

are unavailable, and nothing is known of the searching strategy of the fox. How-

ever, the differences in the numbers of individuals taken, especially between Eleodes

and Coelus, are so marked that selective predation to avoid the beetles with de-

fensive secretions seems likely. — John T. Doyen, Division of Entomology and Para-

sitology, University of California, Berkeley 94720.

A note on the Nesting Biology of Dianthidium pudicum. pudicum

(Cresson) (Hymenoptera : Megachilidae).- — A nest of Dianthidium pudicum

pudicum (Cresson) was discovered on 26 March, 1967, 9.3 miles south of Quartsite,

Arizona. The nest contained 10 cells located about one meter off the ground in

a forked branch of a creosote bush, Larrea tridentata Sesse & Mocino. The branch

was in a horizontal position and about 6 mm in diameter at the nest location.

The nest assumed the triangular shape of the branches (Fig. 1), with the sides

27-28 mm long. The dorsal surface extended 6 mm above the horizontal branches

and the bottom extended 1-2 mm below the branches, averaging about 13 mm

thick.

A dark brown resin, with a strong scent of Larrea, cemented numerous small

pebbles together to form the outer surface of the nest. Internally, the resin became

The Pan-Pacific Entomologist

Fig. 1 . A nest of Dianthidium pudicum pudicum.

softer and yellowish-green in color. The small pebbles were tightly embedded

throughout the interior of the nest, with occasional pieces of masticated leaf

material.

In constructing the 10 vertically fused cells, the female bee appeared to have

built the first cell in the base of the forked branches and then added rows of

two, three, and four cells as she increased the triangular shape of the nest. The

cells measured 5-5.5 mm in width and 9-10 mm in length. The cell walls averaged

0.5-0.75 mm in thickness. Cell surfaces adjacent to the branches had only a

thin layer of resin, whereas the other surfaces contained numerous small pebbles

which extended through the inner surface of resin into the cell cavity. Cell caps

of the downward opening cells averaged 0.5-0.75 mm in thickness. The caps of

6 cells consisted of a circle of small pebbles with a resin filled center, while the

other 4 cell caps did not show this construction pattern.

The dark brown cocoons were thin walled and transparent and the bees were

visible through the cocoons. Anteriorly, the cocoons were thicker and heavily

interwoven with white, silk threads (0.03-0.05 mm thick) which gave rise to the

outer median mammillary projection. The white mammillary projections measured

0.75 mm in length and 0.75-1.0 mm wide at the base. Cocoons ranged from 8-8.5

mm in length from the tip of the projection to the base and they were about 4

mm wide. An air space approximately 1 mm in length separated the tip of the

projection and the inner surface of each cell cap.

Individual orange fecal pellets were about 0.75 mm in length and 0.25 mm in

width, without surface grooves or ridges. The cylindrical pellets were smeared

against the inner surface of the cell walls, forming a continuous reddish-brown

layer 0.25 mm thick around all but the anterior quarter of the cell. This fecal

layer smooths the rough inner surface of the cell.

When the nest was opened in early April, the 3 basal cells and one of the

third row cells contained post-defecating larvae and the remaining cells held 4

Vol. 50, No. 1, January 1974

male and 2 female pupae. Three of the pupae in the outer row of cells had their

heads pointed downward, whereas the other 7 bees were oriented with their heads

up. Adults emerged 10-15 days later.

There are several differences between the nests observed by Hicks (1927, Psyche

34: 193-198) and the nest we have described. All nests described by Hicks were

attached to rocks found on the surface of the ground. The cells were larger,

measuring 15 mm in length, and the cocoons were larger than those we observed. All

of the cells examined by Hicks contained bees oriented downward and we observed

only 3 of 10 bees oriented downward. Grigarick and Stange (1968, Bull. Calif.

Insect Surv. 9: 1-113) briefly mentioned the discovery by F. D. Parker in Reno,

Nevada of a nest of D. p. puclicum in the crotch of a small fruit tree.

Dr. A. A. Grigarick kindly identified the Dianthidium for us. — S. L. Clement,

University of California, Davis 95616 and R. W. Rust, University of Delaware,

Newark 19711.

The Pan-Pacific Entomologist

PACIFIC COAST ENTOMOLOGICAL SOCIETY

J. A. Chemsak C. B. Philip M. R. Gardner P. H. Arnaud, Jr.

President President-Elect Secretary Treasurer

Proceedings

Three Hundred and Fiftieth Meeting

The 350th meeting was held Friday, February 23, 1973 in the Morrison Audi-

torium of the California Academy of Sciences, Golden Gate Park, San Francisco,

with President Chemsak presiding.

Members present (34) : P. H. Arnaud, Jr., D. Bacon, R. G. Blair, T. Briggs,

J. G. Brady, R. M. Brown, D. Burdick, J. A. Chemsak, J. G. Edwards, F. Ennik,

W. E. Ferguson, L. Floyd, N. E. Gary, A. J. Gilbert, A. & A. Gillogly, B. W.

Grushkowitz, K. S. Hagen, A. Hardy, L. S. Hawkins, T. E. Hewton, H. B. Leech,

D. W. Moss, R. P. Papp, C. B. Philip, B. Potts, J. Powell, C. Radus, C. Rees,

E. S. Ross, S. L. Szerlip, R. W. Thorp, T. Toren, M. S. Wasbauer.

Visitors present (29) : L. Bezark, N. Blair, A. Bush, W. D. Carstens, M. R.

Ebertz, S. Ferguson, K. Gary, M. Gilbert, D. Grushkowitz, W. Hardy, S. Jordan,

A. I. B. Kaplan, G. J. Keutz, C. Kitayama, B. Langston, C. Langston, R. L. Morr-

ison, G. M. Nishida, K. Nishida, N. E. Papp, W. Rawscha, R. G. Repke, M. Robey,

E. Schmidt, J. Thorp, R. T. Wian, G. L. Willey, P. Willey, A. Yuaz.

The minutes of the meeting held 15 December were summarized.

The following names were proposed for membership in the Society. Student

Membership: Clifford L. Radus, James P. Anduiza, Robert W. Pemberton, War-

ren E. Savary, Laurence A. Lacey, Steven K. Ault, Larry G. Bezart, Gary L. Willey,

Gordon M. Nishida, Regular Membership: Glenn M. Yoshimura, Regina M.

Burke, Charles D. Hunter, Fred Punzo, Kilian Roever, Ernest Anderson and Dr.

Carl 0. Mohr.

The following note was presented:

Polistiopsis mima Townsend (Diptera: Tachinidae) in Costa Rica. — Two

species are presently assigned to the genus Polistiopsis Townsend — the type species

mima Townsend 1915 (recorded from Mexico — the holotype male from Tehuantepec

and a second male from Fortin de las Flores) and williamsi Arnaud 1966 (recorded

from Paragray — the holotype male from Paso Yobay and the allotype female from

Villarica) . Townsend considered mima to be a polistid counterfeit. When I re-

vised the genus in 1966 (American Museum Novitates, No. 2241, pp. 1-12, figs.

1-20) only the four specimens cited above were available for study from the major

North American entmological collections.

This past summer while studying in Paris at the Museum National d’Histoire

Naturelle (supported by Grant No. 6351, Penrose Fund, American Philosophical

Society), it was a pleasure to discover a series of 15 specimens of Polistiopsis

among the miscellaneous unstudied Tachinidae. I would like to thank Monsieur

Loic Matile for their loan. A recent examination of these has shown that they

represent 12 males and 3 females (the latter the first known of P. mima Townsend.

This series of specimens is labeled with the data “Costa Rica/Paul Serre 1920”;

unfortunately without more precise locality data. Two of the males have the ad-

ditional datum that they were collected during the month of “Juillet”. Several

Vol. 50, No. 1, January 1974

specimens of this series are displayed this evening. — Paul H. Arnaud, Jr., Cali-

fornia Academy of Sciences, San Francisco.

The principal speaker of the evening was Dr. Norman E. Gary, Professor of

Entomology, University of California, Davis. His illustrated talk was entitled:

“Magnetic Retrieval of Honeybee Labels.”

Coffee and other refreshments were served during a social hour held in the

entomology rooms following the meeting. — M. R. Gardner, Secretary.

Three Hundred and Fifty-First Meeting

The 351st meeting was held Friday, 30 March 1973 in the Morrison Auditorium

of the California Academy of Sciences, Golden Gate Park, San Francisco, with

President Chemsak presiding.

Members present (36) : F. G. Andrews, C. Armin, P. H. Arnaud, Jr., J. Anduiza,

F. L. Blanc, R. G. Blair, T. Briggs, R. M. Brown, D. Burdick, K. S. Corwin, D.

Denning, J. Denk, J. G. Edwards, F. Ennik, M. R. Gardner, J. Guggole, R. Hall,

L. S. Hawkins, T. E. Hewton, A. I. Kaplan, C. Kitayama, R. L. Langston, H. B.

Leech, R. W. Pemberton, C. L. Radus, F. C. Roberts, E. S. Ross, W. Savary, M. V.

Silveira, R. Stecker, T. J. Toren, R. L. Tassan, R. W. Thorp, P. Welles, G. L.

Willis, D. W. Whitman.

Visitors present (12) : S. Anduiza, T. Andrews, J. R. Batch, F. Blanc, A. Bush,

R. Greek, R. Morrison, P. Neuenschwander, P. Peterson, T. Slay, J. Thorp, P.

Willey.

The minutes of the meeting held 23 March were summarized.

The following note was presented:

Insects and a mite associated with stored Cannabis sativa Linnaeus. —

On January 23, 1970 I visited the Hall of Justice in San Francisco, California, at

the request of Mr. Terry L. Coddington and Mr. John F. Willims to inspect an

insect problem associated with confiscated stocks of Cannabis sativa Linnaeus.

In offices adjoining room vaults in which these stocks were kept, the personnel

were pestered by insects in their offices.

One problem area was a trunk, thought to be from the midwestern United States,

in which a plastic liner kept the plant materials moist. From this source a series

of 145 specimens of Desmometopa sp. (Diptera: Milichiidae, det. C. W. Sabrosky),

11 Scatopse fuscipes Meigen (Diptera: Scatopsidae, det. Alan Stone), 20 Bradysia

sp. (Diptera: Sciaridae, det. R. J. Gagne), and 1 Drosophila busckii Coquillett

(Diptera: Drosophilidae, det. W. W. Wirth) were collected. These flies are pri-

marily scavengers.

On and in packages of the marijuana, 6 adults, 1 larva, and 1 pupal skin of

Plodia interpunctella (Huhner) (Lepidoptera: Pyralidae, det. D. M. Weisman) ,

and specimens of six species of Coleoptera (det. F. G. Andrews) — Ahasverus advena

(Waltl), the foreign grain beetle, Cryptolestes ferrugineus (Stephens), the rusty

grain beetle, Cryptolestes pusillus (Schonherr), the flat grain beetle, and Oryzae-

philus surinamensis (Linnaeus), the sawtoothed grain beetle of the family Cucuji-

dae, Microgramme arga (Reitter) of the family Lathridiidae, and Typhaea ster-

corea (Linnaeus), the hairy fungus beetle of the family Mycetophagidae were

collected. This group is associated with many types of stored products.

A mite Macrocheles muscadomesticae (Scopoli) (family Macrochelilae, det.

R. L. Smiley) was also collected. I would like to acknowledge my thanks to the

The Pan-Pacific Entomologist

specialists noted above for the identification of the species mentioned.— P aul H.

Arnaud, Jr., California Academy of Sciences, San Francisco.

The principal speaker of the evening was Dr. Werner Loher, Professor of En-

tomology, University of California, Berkeley. His illustrated talk was entitled:

“Acoustical communication in insects.”

Coffee and other refreshments were served at a social hour in the entomology

rooms following the meeting.- — M. R. Gardner, Secretary.

Three Hundred and Fifty-Second Meeting

The 352nd meeting was the annual field day and picnic. It was held on Sat-

urday, 12 May, 1973 at Castle Rock Park near Mount Diablo, Contra Costa County,

California.

Members present (9) : P. H. Arnaud, Jr, R. Blair, H. V. Daly, Jr., H. V. Davis,

W. E. Ferguson, M. R. Gardner, R. W. Pemberton, C. B. Philip, H. Reinhard.

Vistors present (15+) : M. Arnaud, P.-H. F. Arnaud, N. Blair, K. Blair, R. Blair.

Mrs. H. V. Daly, Jr., S. Ferguson, N. Milliet, C. Pemberton, W. J. Reinhard, En-

tomology class of Dr. H. V. Daly, Jr.

The day was warm and sunny. Picnicking and insect collecting were the major

activities of the participants. Judging from the full cyanide vials, flying insects

were found in abundance by the enthusiastic students. — M. R. Gardner, Secretary.

Three Hundred and Fifty-Third Meeting

The 353rd meeting was held Friday, 19 October 1973, in the Morrison Auditorium

of the California Academy of Sciences, Golden Gate Park, San Francisco, with

President Chemsak presiding.

Members present (44) : R. P. Allen, F. G. Andrews, C. Armin, P. H. Arnaud,

Jr., L. Bezark, R. Blair, T. Briggs, D. Burdick, J. A. Chemsak, C. R. Dutton, J. G.

Edwards, B. Ehreth, F. Ennik, B. K. Eya, M. R. Gardner, A. J. Gilbert, A. R.

Gillogly, B. Grushkowitz, K. Hagen, R. Hall, T. E. Hewton, D. Jamieson, C. Kita-

yama, R. L. Langston, H. B. Leech, K. L. Lipps, J. McCarthy, R. Morrison, D. Moss,

G. Nishida, R. P Papp, R. W. Pemberton, C. B. Philip, W. W. Pilke, H. V. Rein-

hard, W. E. Savary, M. V. Silveira, S. L. Szerlip, R. W. Thorp, T. J. Toren, B.

Villegas, D. Whitman, G. Willey, R. L. Wong.

Visitors present (41) : A. Allen, J. Baier, G. M. Bass, Mrs. Blair, P. S. Crane,

W. Craven, W. Diest, A. Edwards, S. Ferguson, T. Gabel, J. Gelhaus, R. Genberg,

M. Gershery, M. Gilbert, D. Green, D. Gropman, K. S. Horn, H. W. Horn, J. S.

Hjelle, H. Implom, S. Jordan, H. Kent, B. Kidwell, G. Krolmer, V. Mayer, D.

Meyers, B. Michalski, K. Nishida, N. Papp, P. Pitcher, M. Roberts, G. Rolfec,

J. Schager, T. Schager, J. T. Sorenson, R. Stockhill, J. Thorp, B. Tilden, J. Wells,

P. Willy, E. Yasukawa.

The minutes of the meetings held 30 March and 12 May were summarized.

The following persons were elected to membership in the Society. Student mem-

bership: Thomas Lugaskii, Thomas M. Maxwell, Mark G. Torassa, Philip D. Perkins,

Bruce Tilden. Regular membership: Karen Lipps, Fred Barnes, Richard Rust,

Paul Peterson, Philip Hubert.

The principal speaker of the evening was Dr. Edward S. Ross. His illustrated

talk was entitled: “Memorable insect encounters.”

Vol. 50, No. 1, January 1974

Coffee and other refreshments were served during a social hour in the entomology

rooms following the meeting. — M. R. Gardner, Secretary.

Three Hundred and Fifty-Fourth Meeting

The 354th meeting was held Friday, 16 November 1973, in the Morrison Audi-

torium of the California Academy of Sciences, Golden Gate Park, San Francisco,

with President Chemsak presiding.

Members present (40) : P. H. Arnaud, Jr., J. W. Bass, F. R. Barnes, L. H.

Bezark, R. G. Blair, R. M. Brown, J. A. Chemsak, T. Delgman, J. T. Doyen, J. G.

Edwards, F. Ennik, B. Eya, W. E. Ferguson, J. R. Gabel, M. R. Gardner, J. Gug-

golz, B. W. Grushkowitz, T. E. Hewton, C. Kitayama, R. L. Langston, R. Main,

R. Morrison, D. Moss, G. M. Nishida, R. W. Pemberton, C. B. Philip, W. W.

Pitcher, R. W. L. Potts, J. A. Powell, F. Radus, M. J. Robey, E. S. Ross, R. Schick,

M. V. Silveira, E. L. Smith, C. Tauber, R. W. Thorp, B. A. Tilden, G. Willey,

M. Wasbauer.

Visitors present (12) : N. Blair, A. Bush, L. P. Brower, D. Grushkowitz, M. H.

Lobisser, C. Moffitt, G. Roefes, D. Sherman, J. Thorp, M. Tauber, P. Willey, A.

Wood.

The minutes of the meeting held 19 October were summarized.

Dr. Arnaud mentioned that over 100 persons were in attendance at the meeting

of 19 October, although not everyone may have signed the register.

President Chemsak announced the nominees for the auditing and nominating

committees. For auditing committee he appointed Mr. H. Vannoy Davis Chairman,

and Dr. Edward S. Ross committeeman. For nominating committee he appointed

Dr. Marius S. Wasbauer chairman, and Drs. Robin W. Thorp and E. Gorton Lins-

ley committeemen. The nominations were unanimously approved by the mem-

bership.

The following note was presented:

Paradejeania rutilioides nigrescens Arnaud reared from Hemihyalea sp.

(Diptera: Tachinidae; Lepidoptera: Arctiidae ). — Paradejeania rutilioides

nigrescens has not had a recorded host even though it is the largest species of Cal-

ifornia Tachinidae and is often collected at flowers in the late summer and fall.

Essig in his book (1915. Injurious and beneficial insects of California, second

edition, p. 330) stated that “Caterpillars of various species are the host,” hut no

specific names are given. It has been suggested that the host would be Heteroceran

(Arnaud, 1968. Pan-Pac. Entomol., 44: 85).

It is now possible to record the first rearing of Paradejeania rutilioides nigres-

cens from the arctiid genus Hemihyalea based on a rearing made by the late Hans

Peter Allmendinger and R. Maddux. Larvae of a species of Hemihyalea were col-

lected by them at Los Gatos, Santa Clara County, California on 17 June 1972 and

the arctiids subsequently pupated. An adult female of nigrescens emerged from

one of the Hemihyalea pupae on 1 September 1972. Mr. Allmendinger had com-

mented that it was not possible to locate the exact pupa from which the nigrescens

was reared since this was a mass rearing. — Paul H. Arnaud, Jr., California Acad-

emy of Sciences, San Francisco.

The principal speaker of the evening was Dr. Lincoln P. Brower, visiting pro-

fessor of environmental studies, University of California, Davis. His illustrated

talk was entitled: “East versus West Coast Monarchs — a Question of Taste.”

The Pan-Pacific Entomologist

A social hour was held in the entomology rooms following the meeting. — M. R.

Gardner, Secretary.

Three Hundred and Fifty-Fifth Meeting

The 355th meeting was held Friday, 14 December 1973 in the Morrison Audi-

torium of the California Academy of Sciences, Golden Gate Park, San Francisco,

with President Chemsak presiding.

Members present (45) : J. Anduiza, F. G. Andrews, P. A. Arnaud, Jr., F. R.

Barnes, J. W. Bass, L. Bezark, R. G. Blair, J. W. Brady, R. M. Brown, J. A.

Chemsak, T. Delgman, D. G. Denning, J. T. Doyen, D. Durbin, J. G. Edwards, B.

Ehreth, F. Ennik, W. E. Ferguson, J. R. Gabel, A. J. Gilbert, P. A. Golen, B. W.

Grushkowitz, R. Hall, S. E. Heston, C. Kitayama, R. L. Langston, K. L. Lipps, J.

McCarthy, A. E. Michelbacher, R. W. Pemberton, P. A. Peterson, C. B. Philip,

W. W. Pitcher, J. A. Powell, H. G. Real, H. I. Scudder, M. V. Silveira, E. L.

Smith, R. E. Somerby, R. E. Stecker, S. L. Szerlip, R. W. Thorp, B. Tilden, W. H.

Tyson, G. L. "Willey.

Visitors present (23) : F. Baker, M. Brady, T. Brum, M. A. Chemsak, J. Da-

monte, L. Denning, L. Doyen, T. D. Eichlin, D. Eschen, S. Ferguson, T. Gabel,

M. Gilbert, K. Ghandi, D. Green, D. Grushkowitz, K. S. Heston, P. Heston, M.

Michelbacher, N. Mamas, M. A. Santana, P. Tyson, J. Thorp, P. Willey.

The minutes of the meeting held 16 November were summarized.

The following people were elected to membership in the Society. Regular mem-

bership: Rev. Andre Larochelle and Dr. Howard E. Evans.

The following note was presented:

Collection of adult Deuterophlebia coloradensis Pennak at Boulder Falls,

Colorado (Diptera: Deuterophlebiidae) . — The Deuterophlebiidae or mountain

midges are a rarely collected family of flies, with seven named species, and several

unnamed taxa, all assigned to the genus Deuterophlebia Edwards. They are known

only from the northern hemisphere, with one species each from Kashmir ( D .

mirahilis Edwards, 1922), Japan ( D . nipponica Kitagami, 1938), Korea ( D . tyo-

senensis Kitagami, 1938) , and four species from the western United States ( D .

coloradensis Pennak, 1945; D. inyoensis Kennedy, 1960; D. nielsoni Kennedy, 1958;

and D. shasta Wirth, 1951) . Probably unnamed taxa have been reported from the

Altai Mountains in Siberia; tributaries of the Alsea and Marys Rivers, Benton

County, Oregon; Yellowstone National Park; and finally several localities in the

Sierra Nevada of California (Kennedy, 1958, Trans. Amer. Miscro. Soc., 77: 201-

228).

Adult Deuterophlebia are rare in collections, with most of the earlier descriptive

work being based on the larval and pupal stages, with adults obtained from the

dissection of pupae. Kitakami (1938. Annot. Zool. Jap., 17: 501) collected fe-

males of D. nipponica from spider webs at Kurama and Mt. Hira and reported a

swarm of females of this species in the early morning at Simasima on the island

of Honshu. Pennak discovered on July 19, 1948 (1950. Entomol. News, 61: 36)

at 8:00 A.M. 200 dying, dead and disintegrating adult males and two adult fe-

males of D. coloradensis floating in small side eddies of North St. Vrain Creek,

near the town of Lyons, Colorado. He suggested that the imagoes emerge in the

early morning hours and flutter about feebly and briefly above the water, but soon

falling to the surface of the water and dying. Kennedy (1958) using rearing cages

Vol. 50, No. 1, January 1974

placed in Convict Creek, California over pupae of D. nielsoni, observed that ima-

gines emerged between 7 to 9 A.M. (P.S.T.), and that in an hour’s flight copula-

tion and deposition of the eggs had taken place. Between 8:00 A.M. and 9:00

A.M. the imagines began to die.

On the field trip day for the International Congress of Systematic and Evolu-

tionary Biology held on August 8, 1973, my first collecting stop was at Boulder

Falls, on north Boulder Creek, about 11 km west of Boulder, Colorado, at an ele-

vation of 2135 meters. On the western bank of the river, within 30 or 40 meters of

the base of the falls, insects were noted in flight and by sweeping an insect net

from water level to a meter and a half above the surface from the edge of the river,

D enter ophlebia were collected. The area was shaded and the rays of the morning

sun had not penetrated the area. Collecting was probably under way at 9:00 A.M.

or shortly thereafter, but the time was unfortunately not recorded. A total of 72

males presumably of D. coloradensis were collected. The type locality for D.

coloradensis (the holotype being a mature larva) is Boulder Creek, at an elevation

of 1810 meters, so that it would be a few kilometers below this Boulder Falls col-

lection. The series of 20 point mounted and 52 alcohol preserved specimens are

deposited in the collection of the Department of Entomology, California Academy

of Sciences.- — Paul H. Arnaud, Jr., California Academy of Sciences, San Francisco.

The December meeting is traditionally the business meeting of the year and

President Chemsak asked for the reports of the auditing and nominating com-

mittees. The auditing committee report was read by Dr. E. S. Ross, and the state-

ment was accepted as read.

President Chemsak announced his appointments to the publication committee

for 1974: Dr. E. Gorton Linsley and Dr. Marius S. Wasbauer.

Dr. Marius Wasbauer, Chairman of the nominating committee, presented the

slate of candidates for office in the Society for 1974: President, Cornelius B.

Philip; President-elect, Howell V. Daly; Secretary, Franklin Ennik; Treasurer,

Paul H. Arnaud, Jr. There were no nominations from the floor. The candidates

were elected to office in the Society for 1974 by a unanimous vote.

The principal speaker of the evening was the outgoing President of the Society,

Dr. John A. Chemsak. His presidential address was entitled, “The Relationships

of the Cerambycidae of Baja California and Mainland Mexico.”

Coffee and other refreshments were served during a social hour held in the en-

tomology rooms following the meeting. — M. R. Gardner, Secretary.

Dr. Arnaud reports the Treasurer’s office is indebted to Mrs. V. Hawley, Office

Manager of the Academy, for her dedicated attention to the Society’s accounts,

to Mrs. Maria Paz Malkerson, Entomology secretary, for her attention to many

letters and mailings of Society publications, and to Mr. H. Vannoy Davis, member

of the firm J. K. Lasser and Company, not only for the audit of the Treasurer’s

records, but also for the completion of our federal and state income tax forms.

The Pan-Pacific Entomologist

PACIFIC COAST ENTOMOLOGICAL SOCIETY

(A California Corporation)

Statement of Income, Expenditures and

Changes in Fund Balances

Years Ended September 30, 1973 and 1972

1973 1972

Income:

Dues and subscriptions $ 5,526 $ 5,231

Reprints and miscellaneous 2,999 5,174

Sales of memoirs 3,748 1,487

Advertising 333 437

Interest on savings accounts 601 574

Dividends, American Telephone & Telegraph Co. 224 208

Increase or (decrease) in value of capital

stock of American Telephone & Telegraph Co. 410 470

13,841 13,581

Expenditures:

Publication costs — Pan Pacific Entomologist 9,751 8,017

Memoirs 199 4,735

Reprints, postage and miscellaneous 2,075 1,442

12,025 14,194

Increase or (decrease) in fund balances 1,816 (613)

Fund balances October 1, 1972 and 1971 20,219 20,832

Fund balances September 30, 1973 and 1972 $22,035 $20,219

Statement of Assets

September 30, 1973 and 1972

1973 1972

Cash in bank:

Commercial accounts $ 3,366 $ 2,960

Savings accounts:

General fund 8,426 8,069

Memoir fund 2,189 1,928

Fall fund 2,173 1,972

Life membership fund 1,721 1,540

Total cash in bank 17,875 16,469

Investment in 80 share of American Telephone & Telegraph

Co. common stock (Life membership and Fall funds) ,

at market value 4,160 3,750

Total $22,035 $20,219

Vol. 50, No. 1, January 1974

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Vol. 50

APRIL 1974

No. 2

THE

Pan-Pacific Entomologist

CARLSON AND RITCHER — A new genus of Ochodaeinae and a description

of the larva of Pseudochodaeus estriatus (Schaeffer) (Coleoptera:

Scarabaeidae) 99

SLOBODCHIKOFF — Notes on the biology of Therion circumflexum (L.),

with a description of the immature stages (Hymenoptera: Ichneu-

monidae) 111

STEFFAN — Nearctic species of Schwenkfeldina (Diptera: Sciaridae) 118

HARDY AND ANDREWS — Observations on Megasoma with behavioral

notes on some lamellicorn beetles associated with sand dunes (Cole-

optera: Scarabaeidae, Lucanidae) 124

CHEMSAK AND LINSLEY — Descriptions and records of Clytini from Mexico

and Central America (Coleoptera: Cerambycidae) 129

HOVORE AND GIESBERT — Two new species of Cerambycidae from Southern

California (Coleoptera) _ 139

PHILIP AND FLOYD — New North American Tabanidae XXL Another new

Bolbodimyia from Mexico (Diptera) 145

POTTS — Revision of the Scarabaeidae: Anomalinae 1. The Genera occurring

in the United States and Canada (Coleoptera) 148

SLOBODCHIKOFF — Behavioral and Morphological mimicry in a cranefly

and an ichneumonid 155

HARDY — A new species of Cyclocephala Latreille from California sand

dunes (Coleoptera: Scarabaeidae) 160

CHANDLER — A redefinition of the Tyrini with the addition of Anitra Casey

(Coleoptera: Pselaphidae) 162

DOUTT — The genus Dicopus Enock (Hymenoptera: Mymaridae) 165

SIRI AND BOHART — A review of the genus Mellinus (Hymenoptera:

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MILLER — Two new Heliococcus species, a key to the North American species,

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MILLER — Mealy bugs of San Clemente Island, California (Homoptera:

Pseudococcidae) 193

SCIENTIFIC NOTES 203, 204, 205, 207

RECENT LITERATURE 154, 159, 164, 168

OBITUARY 209

SAN FRANCISCO, CALIFORNIA • 1974

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

THE PAN-PACIFIC ENTOMOLOGIST

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The Pan-Pacific Entomologist

Yol. 50

April 1974

No. 2

A New Genus of Ochodaeinae and a Description of

the Larva of Pseudochodaeus estriatus (Schaeffer)

(Coleoptera: Scarabaeidae) 1

David C. Carlson and Paul 0. Ritcher

Department of Entomology

Oregon State University , Corvallis 97331

Schaeffer (1906) described Ochodaeus estriatus based on a single

specimen from California. From his comments it is clear that he con-

sidered the species different enough from other Ochodaeus to warrant

placement in another genus. He did not do so solely because he had

only the one specimen and did not wish to dissect it.

Since 1906, additional specimens of Ochodaeus estriatus have been

taken in California and Oregon. Study of their morphology indicates

that Schaeffer was indeed correct in his original conclusions. We here-

by erect the new genus Pseudochodaeus for Schaeffer’s species.

Pseudochodaeus New Genus

Type Species. — Ochodaeus estriatus Schaffer, 1906, p. 271; monobasic.

Description. — Body elongate, convex; head, pronotum, and elytra densely setose;

ventral surfaces and legs clothed with longer setae. Head transverse; mandibles

and labrum visible from above; labrum transverse, densely setose and very shallowly

emarginate anteriorly; anterior clypeal margin strongly reflexed. Antennae 10-

segmented, first segment triangular, densely setose; club with eighth segment en-

larged and cupped, receiving segments 9 and 10 (Fig. 5) . Eyes not divided by

ocular canthus. Pronotum subquadrate, evenly convex except for shallow, median

longitudinal impression; anterior margin shallowly and evenly indented; posterior

margin sinuate on each side of midline; marginal bead entire. Scutellum setose and

punctate, rounded posteriorly. Elytra long, apex of abdomen not visible from above;

posterior sutural angle simple, not dentate; 1 punctate stria between median suture

and humeral umbone, this stria close to median suture, becoming confluent with

median suture at elytral declivity. Propygidium simple, not modified to accept

apex of elytra. Pygidium and 6 visible abdominal sternites densely setose; sternites

free, not connate, fifth stemite lacking stridulatory peg. Foretibia tridentate,

teeth well developed. Outer apical spur on meso- and metatibia pectinate; meta-

trochanter produced beyond posterior edge of femur into sharp projection. Male

1 The financial support of the National Science Foundation (Grants GB-3586, GB-6194X, and

GB-31129) and of the Oregon Agricultural Experiment Station is gratefully acknowledged. Oregon

Agricultural Experiment Station, Technical Paper No. 3786.

The Pan-Pacific Entomologist 50: 99-110. April 1974

100

The Pan-Pacific Entomologist

genitalia (Figs. 1-3) : proximal end of basal piece tapering to point; parameres

symmetrical, lacking basal elongate process; V-shaped apophysis present; aedeagal

sac well developed, bearing numerous sclerotized structures.

The genus Pseudochodaeus possesses the following combination of characters

which distinguishes it from Ochodaeus Serville: single elytral stria, pectinate

metatibial spur, absence of a stridulatory peg, enlarged and cupped eighth antennal

segment, unmodified propygidium, elongate elytra not modified at the apical

sutural angle, and produced metatrochanters. In Ochodaeus the eighth antennal

segment is not cupped (Fig. 6) and the parameres of the male genitalia each

possess an elongate basal process (Fig. 4) . It should be noted that at least one

species now placed in the genus Ochodaeus lacks a stridulatory peg (Carlson,

1975) .

The genus Pseudochodaeus is of particular interest because it has

characters in common with other Ochodaeinae such as Ochodaeus,

Codocera Eschscholtz, and Chaetocanthus Peringuey, and with Pachy-

plectrus LeConte, now placed in the Hybosorinae. Chaetocanthus is

known only from South Africa and differs from Pseudochodaeus in

having bidentate foretibia, 9-segmented antennae with the eighth seg-

ment not enlarged, and pectinate tibial spurs on all legs. Chaetocanthus,

however, has elytra with only one stria, a tranverse and non-emarginate

labrum, and lacks the stridulatory peg.

VanDyke (1904) and Fall (1904) mentioned that a new species of

Ochodaeus, presumably 0. estriatus, was similar in some ways to Pachy-

plectrus laevis LeConte. We have examined P. laevis and find several

similarities to Pseudochodaeus and other genera in the Ochodaeinae.

The male genital apparatus in Pachyplectrus laevis has a pointed basal

piece, symmetrical parameres, and a well developed aedeagal sac, char-

acters shared by Pseudochodaeus, Ochodaeus, Codocera, and Chaetocan-

thus (Carlson, 1975). The eighth abdominal spiracle of Pachyplectrus

laevis is located in the tergites and the seventh is adj acent to, but not sur-

rounded by the tergite. In the Ochodaeinae both the seventh and eighth

abdominal spiracles are in the tergites, however, in P seudochodaeus

the seventh spiracle is close to the edge of the tergite. In Ochodaeus

Figs. 1-8. Fig. 1. Male genitalia of Pseudochodaeus estriatus (Schaeffer),

right lateral view. AS, aedeagal sac. Fig. 2. Male genitalia of P. estriatus, (Schaef-

fer), left lateral view. AS, aedeagal sac; BP, basal piece; PM, parameres. Fig. 3.

Male genitalia of P. estriatus (Schaeffer), dorsal view. AP, V-shaped apophysis;

BP, basal piece; PM, parameres. Fig. 4. Male genitalia of Ochodaeus praesidii

Bates, dorsal view. Fig. 5. Antennal club of P. estriatus (Schaeffer), dorsal view.

Fig. 6. Antennal club of Ochodaeus pectoralis LeConte, dorsal view. Fig. 7. Fe-

male, hind leg of P. estriatus (Schaeffer), ventral view. SP, setal patch. Fig. 8.

Male, hind leg of P. estriatus (Schaeffer), ventral view.

Vol. 50, No. 2, April 1974

101

102

The Pan-Pacific Entomologist

the seventh spiracle is well within the tergite. Pachyplectrus laevis

lacks pectinate tibial spurs and the stridulatory peg.

Examination of several other genera in the Hybosorinae ( Hybosorus

MacLeay, Phaeochrous Laporte, and Chaetodus Westwood) has revealed

that the male genital apparatus has a truncate basal piece and asymmetri-

cal parameres. Also, according to Ritcher (1969 a, b), only the eighth

abdominal spiracle of these genera is located in the tergites.

On the basis of the characters we have examined, it appears that

Pachyplectrus is an intermediate form and the limits of the Ochodaeinae

and Hybosorinae may need to be reevaluated. A final decision on this

will have to await a more extensive study of the morphology of Pachy-

plectrus and examination of Synochodaeus Kolbe and Enodognathus

Benderitter, two other genera in the Ochodaeinae.

Pseudochodaeus estriatus (Schaeffer) New Combination

(Figures 1-3, 5, 7-9)

Ochodcieus estriatus Schaeffer, 1906, p. 271.

Type Material. — Holotype: Female. In the NMNH. Data on labels: Cal-

ifornia, Fresno County, Millwood; Holotype; estriatus Schaef. ; Property of

Mark Robinson; M. Robinson collection 1959. We have examined the holotype

and find it is missing one hind leg and one antenna, but is otherwise intact. It

is 9.0 mm in length and 4.2 mm in width through the midpoint of the elytra.

Description. Males— Length 5.7 mm to 9.5 mm, width through midpoint of

elytra 2.7 mm to 4.6 mm. Color light brown to dark reddish-brown. Vertex, pronotal

disc, scutellum, and elytra covered with relatively short, yellow, semi-erect setae.

Margins of pronotum and elytra with longer yellow setae. Mandibles distinctly

angulate on outer margins, not evenly convex. Left mandible bidentate with large

prostheca and basal molar areas; secondary tooth broad, not pointed. Right

mandible bidentate with large prostheca and molar area; secondary tooth bluntly

pointed. Labrum densely setose, nearly obscured by setae, dorsal surface simple.

Mentum subquadrate, shallowly emarginate anteriorly, with shallow, median,

longitudinal impression; setose on each side of median impression. Anterior

clypeal margin reflexed, forming tubercle at center; clypeus setose. Frons im-

pressed medially, setose and punctate, setae arising from punctures, anterior edge

of punctures slightly elevated; median area of vertex sparsely punctate and lack-

ing setae; posterior- lateral areas of vertex setose and punctate. Pronotum convex,

with median longitudinal impression; disc evenly setose, each seta arising from

puncture. Scutellum setose and punctate. Surface of each elytron densely setose;

setae arising from punctures. Apical sutural angle of elytra simple, not dentiform.

Foretibia with apical spur large, arcuate, and with apex enlarged. Meso tibia

expanded laterally; outer spur pectinate; first tarsal segment longer than next

three segments together. Metafemur with large tooth on posterior margin at

apical one-third (Fig. 8) . Metatibia expanded laterally (Fig. 8) ; oblique row

of spines running from base to inner margin at distal one-third; subapical oblique

row of spines at outer margin; outer apical angle between subapical row of

Vol. 50, No. 2, April 1974

103

Fig. 9. Distribution of Pseudochodaeus estriatus (Schaeffer) in California

and Oregon, U. S. A.

104

The Pan-Pacific Entomologist

spines and apex sparsely setose; outer spur pectinate; first tarsal segment longer

than next three segments together. Male genitalia (Figs. 1—3) : aedeagal sac

with 1 basal sclerotized structure and 5 to 7 large, sclerotized spines; distal

two-thirds of sac covered with irregularly arranged patches of pigmented spiculi.

V-shaped apophysis with distal margin rounded.

Females. — L ength 5.8 mm to 9.3 mm, width through midpoint of elytra 2.9

mm to 4.3 mm. Other characters same as males except that the outer apical

angle of the metatibia bears a dense patch of setae (Fig. 7) .

Distribution (Fig. 9) and specimens examined (holotype, 41 $ , 49 $).

California: El Dorado County : 2 2, July 1932, F. T. Scott (KU) ; 4 $, 4 2,

Pacific House, July — 3 August 1931, H. Hinton (CAS); 1 $, Snowline Camp,

20 July 1948, P. D. Hurd (UCB) ; 1 $ Riverton, August 1930, H. Hinton (CAS) ;

1 2, Pollock Pines, 25 July 1938, A. T. McClay (ASU) ; 1 d , nr. Whitehall,

21 July 1931, H. Hinton (NMNH) ; 1 2, Garden Valley, 4 July 1965 (ARH).

Fresno County. 1 $ 2 $ , vie. Wishon, 15 August 1971, D. Marqua (PHS) ;

1 2, vie. Wishon, 6 August 1971, F. T. Hovore (ORSU) ; 1 d, Cedar Grove,

Kings River Cyn, 17 July 1952, Cazier, Gertsch, Schrammel (AMNH) ; 1 $,

Huckleberry Meadow, 6500', August 1912 (CAS); 1 $, Camp Greeley, 2800',

25 July (CAS); 1 $, Millwood (NMNH). Madera County: 1 $, Sugar Pine,

21 July 1933, R. P. Allen (CAS); Id, Placer Ranger Station, NE Northfork,

30 July 1931 (CAS) ; 1 2 , Miami Ranger Station, 10 mi N Oakhurst and S of

Yosemite Nat. Park, 24 June 1938 (CAS). Mariposa County: 1 2, Miami Ranger

Station, 4000', 29 June 1946, H. P. Chandler (CAS). Modoc County: 1 2, Hacka-

more, 17 July 1934, 0. H. Schwab (CAS) ; 2 d, Pitt River, May (CAS). Plumas

County: 1 2, Walker Mine, 25 July 1930, E. F. Wohletz (CAS) ; 1 2, S end

of Red Clover Valley, 8 mi N Beckworth, 28 August 1963, H. B. Leech (CAS).

Shasta County: Hat Creek: 1 2, July 1948, A. S. Perry (UCB) ; 1 2 , 16 July

1951, G. Pronin (CAS) ; 1 d, 20 July 1951, G. Pronin (CAS) ; 2 d, 27 July

1951, G. Pronin (CAS); 1 d, 25 August 1951, G. Pronin (CAS); 1 2 , 25

August 1952, G. Pronin (CAS); 1 $ , 12 July 1955, J. W. MacSwain (UCB);

1 2, 19 July 1955, G. Pronin (CAS) ; 1 $ 1 2 , 29 July 1971, D. Carlson (DCC).

Trinity County: 1 d, Carrville, 6 July 1950, D. Guiliani (CAS). Tulare County:

1 d , 2 mi E Posey, 21 August 1971, E. Giesbert (DCC). Tuolomne County:

1 d , trib. to Herring Creek, 1.5 mi N Pinecrest Lake, 30 August 1964, H. B.

Leech (CAS) ; 1 2, Twain-Harte, 4000', July 1937, Blaisdell (CAS) ; 1 2, Straw-

berry Resort, 5243', 8 August 1937, E. Herald (CAS); 1 d, Strawberry, 15

August 1962, C. A. Toschi (UCB). Oregon: Jackson County: Fourbit Ford

Cmpgnd., 11 mi SE Butte Falls: 4 d 5 2 , 18 July 1973, P. O. Ritcher and D. C.

Carlson (ORSU, DCC) ; 8 d 10 2,9 August 1973, D. C. Carlson and W. N.

Mathis (ORSU, DCC, NMNH) ; 3 d 9 2, 29 August 1973, D. C. Carlson and

R. L. Westcott (ORSU, DCC, RLW, HFH) : 1 2, Jet. Rd 3520 and S. Fork

Fourbit Creek, 11 mi SE Butte Falls, 29 August 1973, D. C. Carlson and R. L.

Westcott (ORSU). Lake County: 1 d, Warner Canyon, 31 July 1968, R. L.

Westcott (RLW).

Remarks. — T he color variation of the adults of P. estriatus is ap-

parently a result of age. Specimens which we collected on 18 July 1973

were light brown with the setation intact and legs unabraded. Later

Vol. 50, No. 2, April 1974

105

collections at the same site (9 and 29 August 1973) yielded all dark

reddish-brown specimens which were quite worn.

This species has also been recorded from Deschutes County, Oregon

(Hatch, 1971) , but we have been unable to verify the record. Ochodaeus

simplex LeConte is the only Ochodaeus we and others have collected

in Deschutes County.

We have visited four of the localities where P. estriatus occurs and

have collected specimens at two of them (Fourbit Ford Campground,

Jackson Co., Oregon and Hat Creek, Shasta Co., California). All of

these are forested areas with sparse ground cover of shrubs and grass

and all are close to streams. At Fourbit Ford Campground the follow-

ing plant species were fairly abundant: Pseudotsuga menziesii (Mirb.),

Pinus ponder osa Laws., Abies grandis (Dougl.), Quercus hello ggi

Newb., Holodiscus discolor (Pursh), Ceanothus integerrimus Hook

and Arm., Amelanchier florida Lindl., Symphoricarpos albus (L.),

Mahonia sp., and Corylus sp. Bracken fern and sparse grass were also

present.

Included with the holotype which we borrowed from the NMNH was

another specimen bearing a handwritten locality label identical to that

with the holotype. This specimen is a male and we believe it to be the

misplaced specimen which Schaeffer referred to in the original descrip-

tion of 0. estriatus.

We are depositing specimens of P. estriatus from Oregon, including

males with everted aedeagal sacs, in the NMNH and collection of H. F.

Howden, Carleton University, Ottawa, Ontario, Canada.

Biology and Immature Stages

Information on the biology of the Ochodaeinae is exceedingly scanty,

which is surprising since the adults are often abundant. About all that

has been known of the adults is that they fly at night, are attracted to

light, and burrow in the soil.

By dissection, we have determined that the mid- and hindgut of P.

estriatus adults often contain numerous small spores of a basidiomycete

similar to those of puff balls. The female reproductive system has 6

ovarioles in each ovary but only 1 egg develops at a time on each side.

A typical fully developed egg is white, elongate-oval in shape, 2.78 mm

long, and 1.44 mm wide. This is quite large for such a small beetle.

Like other Ochodaeinae, the flight period of P. estriatus is quite

long. Adults were taken at light in southern Oregon from July 18 to

August 29, 1973. Dissection indicated that egg development proceeds

slowly and that the few eggs are laid over a long period of time.

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While digging for Pleocoma LeConte larvae in southern Oregon in

1961, David Fellin, then a graduate student, obtained a small scarabaeid

larva with 4-segmented antennae, which was unlike any other with

which the junior author was familiar. He suspected it to be an Ochodaeus

since he was familiar with the larvae of all scarab genera occurring in

in Oregon except those of Glaresis Erichson, Bolborhombus Cartwright,

and Ochodaeus. Glaresis was eliminated because of its very small size

and Bolborhombus because larvae of related Geotrupinae all have 3-

segmented antennae.

The main problem with this tentative identification was that adult

Ochodaeus had never been collected in southern Oregon west of the

Cascade Mountains. In recent years, a number of unsuccessful attempts

were made to collect Ochodaeus adults at black-lights in southern

Oregon, in the vicinity of Butte Falls. Finally, on 18 July 1973 we took 9

adults (4 #,5$) of Pseud ochodaeus estriatus (Schaeffer), 11 miles

southeast of Butte Falls. This site is less than a mile from where the

supposed Ochodaeus larva was dug in 1961.

Medvedev (1960) described the larva of Codocera ferruginea Esch-

scholtz, a common Eurasian species of Ochodaeinae, based on 2 un-

associated larvae collected in the soil of a Ukrainian tree plantation.

Unfortunately, characters of the epipharynx, mandibles, and maxillae

were neither described nor figured and we were unable to borrow the

specimens for study.

At first, we assumed our Ochodaeus larva differed from Medvedev’s

Codocerra larva because they belonged in different genera. However,

Medvedev’s larva could well be a Trox Fabricius since it resembles that

genus in the type of head setation, in having 3-segmented antennae,

and in having fleshy anal lobes.

Pseudochodaeus estriatus (Schaeffer), Third-Stage Larva

(Figures 10-20)

Description based on one third-stage larva collected from the soil 10 miles east

of Butte Falls, Jackson County, Oregon, 28 January 1961, by David Fellin.

Figs. 10-18. Larva of Pseudochodaeus estriatus (Schaeffer). Fig. 10. Head.

A, antenna; C, clypeus, DES, dorsoepicranial seta; F, frons; L, labrum; PFS,

posterior frontal setae. Fig. 11. Epipharynx. CPA, chaetoparia; GP, gymno-

paria; NI-3, nesia; PE, pedium; PA, phobae; ZY, zygum. Fig. 12. Left mandible.

PA, preartis; MA, molar area; SA scissorial area (upper surface). Fig. 13. Right

maxilla (lower surface). CAR, cardo; G, galea; GU, uncus of galea; LA,

Vol. 50, No. 2, April 1974

107

lacina; LAU, uncus of lacinia; MST, maxillary stridulatory teeth. Fig. 14, Labium

with hypopharynx. GL, glossa. Fig. 15. Last 2 segments of antenna (ventral

surface) . VSS, ventral sensory spot. Fig. 16. Right mandible (Ventral surface) .

MA, molar area. SA, scissorial area; VP, ventral process. Fig. 17. Claw of

prothoracic leg. Fig. 18. Left lateral view of entire third-stage larva. DL1, DL2,

DL3, dorsal lobes.

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Description. — Larva typically scararbaeiform, C-shaped (Fig. 18). Head light

yellow-brown, thorax and abdomen whitish.

Maximum width of head capsule (Fig. 10) 2.28 mm. Surface of head fairly

smooth. Epicranial stem present, frontal sutures largely absent. With 1 dorso-

epicranial seta on each side. Frontal area with pair of posterior frontal setae

on each side, other frontal setae absent. Frontoclypeal suture mostly absent.

Labrum trilobed. Antenna 4-segmented with distal segment reduced in size, distal

segment less than half as long as third segment (Fig. 10) . Ventral surface of

next to last antennal segment (third) with 1 apical, elliptical sensory spot; last

antennal segment also with similar ventral, apical spot (Fig. 15). Apex of last

antennal segment with about 7 sensory pegs. Mandibles (Figs. 12 and 16) subtri-

angular in shape, each with 2 scissorial teeth and with typical ventral, accessory,

articulating process. Maxilla (Fig. 13) with separate galea and lacinia. Maxillary

stridulatory teeth consisting of a group of about 14 short, conical teeth. Maxillary

palpus 4-segmented. Labium (Fig. 14) with symmetrical oncyli. Epipharynx

(Fig. 11) with complete, symmetrical zygum. Pedium surrounded by phobae and

with inner second group of phobae posterior to anterior phobae. Tormae fused,

symmetrical, lacking either epitorma or pternotormae. Haptolachus with 3 nesia;

1 large sclerotized plate on each side and 1 small median sense cone. Chaetopariae

with a few (10 to 11) strong chaetae on each side. Chaetopariae without

microsensilla.

Abdominal segments 1-7 each with 3 dorsal lobes (Fig. 18) . Anterior 2 dorsal

lobes each with tranverse row of setae, posterior dorsal lobe bare. Spiracles crib-

riform, inconspicuous, occurring on prothorax and first 8 abdominal segments.

Prothoracic spiracle elliptical, with slight caudal emargination. Respiratory

plates of abdominal spiracles all similar in size, crescent shaped, with caudo-

ventral emarginations. Legs 4-segmented, each ending in well developed claw

(Fig. 18) : Claws similar in size, each bearing 2 setae (Figs. 17 and 18). Ventral

surface of last abdominal segment (Fig. 19) with poorly developed raster. Raster

consisting of a few (about 18) irregularly distributed, rather short, slender setae.

Raster extending less than one-half the distance to seventh abdominal segment;

anteriorly, with broad, bilobed raised area. Anal opening Y-shaped surrounded

by narrow, transverse, dorsal anal lobe and 2 larger, subtriangular, lower anal

lobes (Fig. 20) .

Remarks. — The larva of P. estriatus has a number of morphological

characters indicating it belongs to the more primitive Scarabaeidae.

These characters are similar to those of the less specialized scarabaeid

subfamilies Troginae and Aphodiinae. It resembles larvae of Trox (sensu

strictu) in lacking a well-developed fronto-clypeal suture, in having a

haptolachus with 2 sclerotized plates and a median sense cone, and in

the type of anal opening and surrounding lobes. It differs from Trox

and Omorgus Erichson larvae, however, in having 4-segmented antennae

(not 3 as in Troginae), a trilobed labrum (not bilobed), and a different

type and arrangement of maxillary stridulatory teeth.

The larva of P. estriatus will key to Aphodiinae in Ritcher’s 1966

monograph and in his 1967 paper. It can be distinguished readily

Vol. 50, No. 2, April 1974

109

Figs. 19 and 20. Larva of Pseudochodaeus estriatus (Schaeffer). Fig. 19.

Ventral view of last (tenth) abdominal segment. Fig. 20, caudal view of last

abdominal segment. AO, anal opening; DAL, dorsal anal lobe; VAL, ventral

anal lobe.

from larvae of Aphodiinae by such characters as the incomplete

frontoclypeal suture (Fig. 10), the complete zygum of the epipharynx

(Fig. 11), and the absence of an epitorma (Fig. 11). It resembles

larvae of Aphodiinae in that both have ventral, apical sensory areas

on the last 2 antennal segments (Fig. 15).

Although nothing is known of the subterranean habits of the larva

of P. estriatus, some inferences can be drawn from its morphology.

The worn mandibles, well developed legs and claws, and body setation

are all indicative of an active larva, not of a larva whose food was pro-

visioned by the adult.

Acknowledgments

We wish to thank the following for the generous loan of specimens:

American Museum of Natural History (AMNH) ; Arizona State Uni-

versity, Tempe (ASU) ; California Academy of Sciences (CAS) ; Oregon

State University (ORSU) ; National Museum of Natural History

(NMNH) ; University of California, Berkeley (UCB) ; A. R. Hardy

(ARH) ; F. T. Hovore (FTH) ; P. H. Sullivan (PHS) ; and R. L.

Westcott (RLW). The letters in parentheses are the abbreviations used

in the text for the collections from which we borrowed specimens. We

are indebted to W. C. Denison, Oregon State University botanist, for

identifying the basidiomycete spores. We also thank R. B. Roberts for

many helpful suggestions and Mrs. B. Hall for assistance with the larval

illustrations.

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Literature Cited

Carlson, D. C. 1974. Taxonomic characters of the genus Ochodaeus Serville

with descriptions of two new species in the 0. pectoralis LeConte

species complex (Coleoptera: Scarabaeidae) . Bull. S. Calif. Acad.

Sci., (in press) .

Fall, H. C. 1904. In: Proc. Pacific Coast Entomol. Soc., 1:22.

Hatch, M. H. 1971. The beetles of the Pacific Northwest. Part V. Univ.

Washington Press, Seattle, 662 pp.

Medvedev, S. I. 1952. Larvae of scarabaeoid beetles of the fauna of the USSR.

Opredeliteli Faune USSR, Moscow, 47:1-343.

Medvedev, S. I. 1960. Descriptions of the larva of eight species of lamelicorn

beetles from the Ukraine and Central Asia. Zool. Zhurn., 39:381-393.

Ritcher, P. 0. 1966. White grubs and their allies, a study of North Ameri-

can scarabaeoid larvae. Oreg. State Univ. Press, Stud. Entomol.,

4:1-219.

Ritcher, P. 0. 1967. Keys for identifying larvae of Scarabaeoidea to the family

and subfamily (Coleoptera). Calif. Dept. Agr. Occ. Papers, 10:1-8.

Ritcher, P. 0. 1969a. Spiracles of adult Scarabaeoidea (Coleoptera) and their

phylogenetic significance. I. The abdominal spiracles. Ann. Entomol. Soc.

America, 62 (2) -.869-880.

Ritcher, P. O. 1969b. Spiracles of adult Scarabaeoidea (Coleoptera) and their

phylogenetic significance. II. Thoracic spiracles and adjacent sclerites.

Ann. Entomol. Soc. America, 62 (6) : 1388-1398.

Vandyke, E. C. 1904. In: Proc. Pacific Coast Entomol. Soc., 1:22.

Vol. 50, No. 2, April 1974

111

Notes on the Biology of Therion circumflexum (L.),

with a Description of the Immature Stages

(Hymenoptera: Ichneumonidae)

C. N. Slobodchikoff

Department of Biological Sciences

Northern Arizona University, Flagstaff 86001

Wasps of the genus Therion are internal parasitoids of Lepidoptera

larvae. A female wasp injects one or more eggs into the body cavity of

a caterpillar, and the parasitoid larva, upon hatching from the egg, begins

to feed on the non-vital tissues of its host. The parasitoid larva under-

goes three larval molts. When its host pupates, the larval parasitoid

kills the host and pupates within the host’s pupal shell. The biology of

larvae and adults of T. morio (Fabr.) has been described by Tothill

(1922). The purpose of this present paper is to describe the egg, larval

stages, and pupa of T. circumflexum (L.). Behavior of T. morio and

T. circumflexum adults has been described elsewhere (Slobodchikoff,

1973).

Materials and Methods

Six T. circumflexum females were captured on 22 April 1969 at Alpine

Lake, Marin County, California. Each female was immediately in-

troduced into a round cardboard container 10 cm in diameter and 7

cm deep, which was covered with a plastic top. Three containers each

had 5 third instar larvae of Spodoptera exigua (Hbn.), the other

three each had 5 third instar larvae of Pseudaletia unipuncta (Haw.).

Host larvae were obtained from the insect pathology Lepidoptera cul-

tures maintained by the Division of Entomology and Parasitology,

University of California, Berkeley. Female wasps were kept in the con-

tainers for two days, then removed and kept in larger plastic cages

(see Slobodchikoff, 1973, for a description of the cages). All con-

tainers were kept together at an average temperature of 22 °C and an

average relative humidity of 45 percent. A total of 9 Spodoptera and

11 Pseudaletia larvae or pupae were dissected. One Spodoptera larva

was dissected immediately after the adult wasps were transferred to

plastic cages. Six Therion eggs were found. On 28 April, 1969, five

days after initial contact with a parasitoid wasp, 3 larvae of both

Spodoptera and Pseudaletia were dissected. Two Spodoptera larvae

lacked Therion larvae, while the third contained 4 first instar Therion

larvae. Two Pseudaletia larvae were not parasitized, while one con-

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The Pan-Pacific Entomologist

3 4

Figs. 1-4. Therion circumflexum. 1. Egg, lateral view. 2. First instar larva,

enclosed in sac. Dorsal view, head pointing down. 3. First instar larva head

capsule, anterior view; see explanation of fig. 4 for names of labeled structures.

4. Third instar head capsule, anterior view. Labeled structures are the following:

a, antenna; c, clypeus; eps, fused epistoma, pleurostoma, and hypostoma; Ip,

labial palp; Is, labial sclerite; m, mandible; mp, maxillary palp; sp, silk press;

ss, stipital sclerite.

Vol. 50, No. 2, April 1974

113

tained a single first instar Therion larva. Twelve days after initial con-

tact (5 May, 1969), two Spodoptera and six Pseudaletia were dissected.

Each Spodoptera contained a single second instar Therion larva. Three

Pseudaletia larvae were unparasitized, one contained two live and seven

dead larvae, one contained two live and six dead larvae, and one con-

tained three live and 10 dead larvae. Nineteen days after initial con-

tact (12 May, 1969), two Spodoptera larvae and one Pseudaletia larva

were dissected. All three larvae each contained a single third instar

Therion larva. Twenty six days after initial contact (19 May, 1969),

one Spodoptera and one Pseudaletia pupa were dissected. Each con-

tained a Therion pupa. One Spodoptera pupa produced an adult

male Therion on 25 May, 1969. The five remaining Spodoptera larvae

died before pupating. One Pseudaletia pupa produced an adult moth on

27 May, 1969. The remaining Pseudaletia died before pupating. Measure-

ments were made with an ocular micrometer and a Wild M7 binocular

dissecting microscope. Drawings were made with a Wild drawing tube

attachment to the M7 microscope.

Descriptions and discussion

Egg (Fig. 1). Total length, 0.371-0.266 mm (x zz 0.306 mm, N = 6). Total

width, 0.214-0.186 mm (x zz 0.199 mm) ; Color milky- white. Anterior end

elongate — globose; Caudal end prolonged into a tapering petiole; Length of

petiole, 0.221-0.184 mm (x zz 0.199 mm) ; Width of petiole at widest point,

0.020-0.011 mm (x = 0.014 mm); Width of petiole at caudal tip, 0.0020-0.0016

mm (x zz 0.0018 mm) .

All eggs found were attached to the body wall of the host. Two eggs

were found in the thoracic region of the host caterpillar, and 4 eggs

were removed from the abdominal region. Based on this small sample,

it appears that the female wasp does not have a preferential oviposition

site, at least under confined laboratory conditions. Tothill (1922) re-

ported that T. morio females lay an egg into the body wall opposite

to that initially pierced by the ovipositor. This egg-laying behavior is

also present in T. circumflexum (Slobodchikoff, 1971).

First instar (Figs. 2-3). Body enclosed in translucent white sac; length of

larva, 3.1-2.8 mm (x zz: 3.0 mm, N = 11) ; width of larva at midgut, 1.2-1.0 mm

(x zz 1.0 mm) ; body slender; color white; head capsule large, quadrate, strongly

sclerotized; antenna papilliform; mouthparts prognathous; mandibles large, falcate,

strongly sclerotized; maxillary palp present as disc; labial palp papilliform;

labral sclerite present as large, sclerotized hemispherical plate, incomplete laterally;

stipital sclerite large, hemiellipsoid, open laterally; hypostoma, pleurostoma, and

epistoma strongly sclerotized, hypostomal arch complete dorsally, silk press strongly

sclerotized.

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The Pan-Pacific Entomologist

Figs. 5-6. Therion circumjlexum. 5. Third instar larva, lateral view. 6.

pupa of T. circumflexum, lateral view.

Tothill (1922) reports that the sac which encloses the first instar

larva of T. morio is made up of 2 non-cellular layers. The mandibles

of the larva are enclosed in this sac and are not used for lacerating the

tissues of the host. Tothill postulates that the food of T. morio first

instar larvae is the strained liquid of the hemolymph. The first instar

larva of T. circumflexum has its mandibles protruding from the sac,

and the mandibles may be used for lacerating tissue. One first instar

larva was observed lacerating and ingesting its host’s fat body tissue.

Another apparent difference between T. morio and T. circumflexum is

Vol. 50, No. 2, April 1974

115

the location of the first instar larvae in its host. Tothill reports that

T. morio larvae are always found in the caudal half of their host larva,

Hypantria cunea (Drury), and in the extreme caudal portion of a

host pupa. In the present study no pupae were found to contain first

instar larvae, and in both Spodoptera and Pseudaletia first instar

Therion larvae were found in both thoracic and abdominal regions.

Results of the dissections suggest that a T. circumflexum egg develops

into a first instar larva within five days in both Pseudaletia and

Spodoptera. First instar larvae may persist for as long as 12 days in

Pseudaletia. The dead first instar larvae of Therion found in Pseu-

daletia 12 days after oviposition were represented by head capsules and

portions of the enclosing sacs, suggesting that they had been killed and

partially eaten by other first instar larvae. No encapsulated larvae were

found.

Second instar. Body free of any enclosing sac; length 6.5-5. 0 mm (x = 5.75

mm, N = 2) ; width at midlength of body 1. 3-1.0 mm (x 1.15 mm) ; color

white; similar in attributes to first instar.

The second instar differs from the first primarily in the lack of an

enclosing sac and a larger body size. Tothill (1922) reports that the

mandibles of the second instar of T. morio are smaller than those of the

first instar, and are probably used for feeding on the host’s tissues. The

mandibles of the second instar of T. circumflexum were not observed

to be any smaller than those of the first instar. Although feeding was

not observed for the second instar, it seems probable that this instar,

like the first, feeds directly on the tissues of the host. In Spodoptera ,

the time of development from oviposition to second instar is approxi-

mately 12 days. The available data suggest that the time of develop-

ment from oviposition to second instar may be slightly longer in

Pseudaletia.

Third instar (Figs. 4-5). Length, 14.0-9.0 mm (x = 11.2 mm, N = 3) ; width

at midlength, 7.2-4.0 mm (x 5.6 mm) ; color yellow-white; no apparent setae

on head or body; antenna disc-shaped; antennal discs dorsomedially contiguous,

separated by thin extension of dorsal head capsule sclerotization ; maxillary and

labial palps disc-shaped; mandible falcate; strongly scleortized distally, broadly

trapezoidal in shape proximally; clypeus pentagonal, lightly sclerotized; labrum

small, ellipsoid, heavily sclerotized; labial sclerite thin, heavily sclerotized laterally,

expanded into lightly sclerotized hemisphere ventromedially ; sclerotic spur absent;

epistoma, pleurostoma, and hypostoma fused, expanded laterally; hypostoma

strongly sclerotized; stipital sclerite ellipsoid, medially recurved toward base of

mandible, thickening laterally; silk press prominent, strongly sclerotized dorsally,

weakly sclerotized ventrally ; abdominal segments 2-10 each with one pair of

lateral lobes.

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The Pan-Pacific Entomologist

Tothill (1922) reports that the third instar larva of T. morio is

similar to the second instar, and the thoracic segments have buds indicat-

ing the position of the legs and the wings. The third instar larva of

T. circumflexum is very different from the second instar. The head

capsule and mouthparts of the third instar are greatly reduced, the body

is wider, and the abdominal segments have lateral lobes. No leg or

wing buds are present on the thorax. In both Spodoptera and Pseu-

daletia, third instar larvae were found after the hosts had pupated.

In each case, the larva was found occupying the abdominal part of

the pupa. One Spodoptera pupa and one Pseudaletia pupa had no

tissue left inside the pupal integument. The other Spodoptera pupa

had no tissue remaining in the abdominal region, but the thoracic re-

gion was still filled with tissue. All three instar larvae removed from

Spodoptera were oriented with their heads pointing toward the thorax

and the head of the host. Development time from oviposition to third

instar is approximately 23 days in both Spodoptera and Pseudaletia.

Pupa (Fig. 6). Length, 14.0-12.0 mm (x = 13.0 mm, N = 2) ; width at widest

point 3.5-3.0 mm (x 3.25 mm) ; color white-pink, eyes light purple; antennal

sockets not prominent; labrum and clypeus elevated, projecting outward from head;

clypeus pentagonal, height 0.75 its width; labrum reduced; mandibles bidentate,

dorsal tooth 1.3 length of ventral tooth, maxilla well differentiated; postmentum

conspicuous, quadrate; fore and hind wings lobiform, white, forewing equal in

length to hind femur, hind wing 0.75 length of hind femur; front wing lobe covers

hind wing lobe; second abdominal segment of abdomen appressed to propodeum,

mesonotum, and scutellum, shortening total body length; first tarsal segment

of hind leg equal in length to segments 2-3 combined; fifth tarsal segment of hind

leg equal in length to segments 3-4 combined; antennae, legs appressed to body.

One Therion pupa was found in a Spodoptera pupa, the other was

found in a Pseudaletia pupa. In each species of host, the Therion

pupa was oriented with its head in the same direction as that of the

host. The pupa of each host was an empty shell, with no tissue remain-

ing inside. No evidence of a cocoon spun by the Therion was found.

Development time from oviposition to pupa is approximately 26 days.

Adults. One male T. circumflexum emerged from a Spodoptera pupa

33 days after oviposition. As an adult, this individual lived in the

laboratory on a diet of honey and water for 28 days. This indicates

that, at least under the laboratory conditions to which the hosts and

parasitoids were exposed, development time from egg to adult is ap-

proximately one month, and the entire life cycle takes approximately

two months. Field studies with marked adults of T. circumflexum

(Slobodchikoff, unpublished) show that individuals may be recaptured

up to 30 days after they are marked. This suggests that the life-span of

Vol. 50, No. 2, April 1974

117

the male reared under laboratory conditions is comparable to, or even

less than, the life span of adults in the field.

Literature Cited

Slobodchikoff, C. N. 1971. Patterns of differentiation and evolution in the

genus Therion. Ph.D. Dissertation, University of California, Berkeley.

Slobodchikoff, C. N. 1973. Behavorial studies of three morphotypes of

Therion circumflexum (L.) . (Hymenoptera: Ichneumonidae) . Pan-

Pacific Entomol., 49: 197-206.

Tothill, J. D. 1922. The natural control of the fall webworm ( Hyphantria

cnnea Drury) in Canada together with an account of its several para-

sites. Canada Dept. Agric. Tech. Bull. No. 3, 107 pp.

118

The Pan-Pacific Entomologist

Nearctic Species of Schwenkfeldina

(Diptera: Sciaridae)

Wallace A. Steffan

Bishop Museum, Honolulu, Hawaii 96818

Schwenkfeldina was known only from the Palaearctic and Oriental

Regions except for the extension of the Holarctic species, S. tridentata

(Riibsaamen) into Canada and Greenland (Tuomikoski, 1967). Some

time ago Tuomikoski suggested that the North American species Brady -

sia dux (Johannsen) and B. imitans (Johannsen) belong in Schwenk-

feldina (pers. commun.). While revising the key to genera of North

American Sciaridae, I examined specimens of B. imitans and com-

pared the drawings of B. dux with those of B. imitans and agree that

they are Schivenkfeldina species.

Initially, Schwenkfeldina was proposed as a subgenus of Neosciara

(Frey, 1942) and contained the Palaearctic species, N. (S.) carbonaria

(Meigen) and the Oriental species, TV. (S.) ponderosa (Walker). In

his revision of northern European Sciaridae, Frey (1948) included

both Schwenkfeldina and Neosciara as subgenera of Bradysia. Tuomiko-

ski (1960) elevated Schwenkfeldina to generic rank. Steffan (1973)

listed S. breviantenna Steffan, S. carbonaria (Meigen), and S. Pon-

derosa (Walker) from the Oriental Region.

Schwenkfeldina species are characterized by the relatively short and

stout antennae, wide eyebridge, setae on the posterior pronotum, the

long Ri which extends beyond the base of the medial fork and frequently

has ventral macrotrichia near the apex.

The biology of the Nearctic species is unknown. There is con-

siderable literature on the Palaearctic species and type, S. carbonaria

(Meigen). Winnertz (1867) reported adults on willow blossoms dur-

ing spring and summer. Beling (1886) noted that larvae live on the

forest floor, preferably in spruce stands ( Abies excelsa D. C.), under

the leaves or litter from dead needles.

The use of the late Dr. Jean L. Laffoon’s sciarid species file is

acknowledged and greatly appreciated. I also wish to thank Ms. Suzanne

Keenan and Mrs. Mary Keeler for preparing the illustrations.

Schwenkfeldina Frey

Neosciara ( Schwenkfeldina ) Frey, 1942, p. 32. Type species Sciara carbonaria

Meigen (original designation) .

Bradysia {Schwenkfeldina) , Frey, 1948, p. 51.

Schwenkfeldina, Tuomikoski, 1960, p. 29; Steffan, 1973, p. 469.

The Pan-Pacific Entomologist 50: 118-123. April 1974

Vol. 50, No. 2, April 1974

119

Schwenkfeldina DUX (Johannsen), new combination

Sciara dux Johannsen, 1912, p. 127-128 [Type: Cornell #2084] ; Leonard, 1928,

p. 747; Johnson, 1930, p. 126; Fisher, 1937, p. 413.

Neosciara dux, Pettey, 1918, p. 322; Johnson, 1925, p. 89.

Lycoria dux, Shaw and Fisher, 1952, p. 212.

Brady sia ( Brady sia ) dux. Stone and Laffoon, 1965, p. 232.

Bradysia dux, Steffan, 1966, p. 35.

Type locality: Wisconsin.

Distribution: Michigan (Stone & Laffoon, 1965). New York (Johannsen, 1912

and Leonard, 1928). Massachusetts (Johnson, 1930). Rhode Island (Shaw and

Fisher, 1952). Wisconsin (Johannsen, 1912).

S. dux is very similar to S. imitans and may be conspecific.

Schwenkfeldina imitans (Johannsen) new combination

Sciara imitans Johannsen, 1912, p. 128 [Type: Cornell #2085]; Garret, 1925,

p. 16; Fisher, 1937, p. 413.

Neosciara imitans, Pettey, 1918, p. 322.

Bradysia ( Bradysia ) imitans, Stone and Laffoon, 1965, p. 233.

Bradysia imitans, Steffan, 1966, p. 36.

Head. Interfacetal hairs abundant, extending well beyond outer curvature of

facets; eyebridge 4-5 facets wide. Anterior vertex with strong setae. Antenna:

flagellomeres with short necks, about % length of flagellomeres, hairs short and

dense about % width; hyaline sensilla sparse; flagellomere 4 (fig. lb) about

1.5 times longer than wide. Prefrons with 7 very long and stout median setae.

Clypeus bare. Palpus 3 segmented, segment 1 with 8 dorsolateral setae and

broad patch of numerous dorsal hyaline sensilla, sensory pit absent. Thorax:

acrostichals and dorsocentrals strong. Posterior pronotum with setae; anterior

pronotum with 7 strong setae. Proepisternum with 8 strong setae. Posterior

mesepimerite rectangular in shape. Legs. Forelegs: length of coxa, 0.51 mm;

femur, 0.71 mm; tibia, 0.86 mm; basitarsomere 0.45 mm; fore tibia with preapical

triangular patch of dense setae (fig. lc) . Tibial spurs, 1: 2: 2; apex of hind

tibia with 11 large lateral setae; hind tibial spurs distinctly longer than width of

tibial apex. Pretarsal claws simple. Wing: length 3.71 mm, width 1.53 mm. Vena-

tion as in fig. la. R-M index 0.8; C-M index 0.8 Ri with ventral macrotrichia near

apex. Abdomen: tergal and sternal setae strong and dark. Terminalia as in fig.

Id; distimere with large median protuberance, dense patch of strong spines slightly

distad of protuberance, and 3-4' short preapical spines. Basimere with dense patch

of mesoapical setae on distinct lobe.

2 . Essentially as male with following differences: wing length, 4.29 mm; width,

1.63 mm. Vaginal furca as in fig. le.

Type locality: Friday Harbor, Washington.

Distribution: S. imitans was previously known only from the type locality, Friday

Harbor, Washington. I have examined specimens from Washington and Cal-

ifornia as listed below.

California: San Bernadino Co., Mt. Home Canyon, 8-VI-1924, F. R. Cole

(C.I.S.), 2$ $, 2 2 2- Washington: Mason Co., Lake Cushman, 27-VI-1917,

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The Pan-Pacific Entomologist

Fig. 1. Schwenkfeldina imitcins. A, wing, male; B, flagellomere 4, male; C,

apex of foretibia, male; D, genitalia, ventral view, male; E, vaginal furca, dorsal

view, female.

H. G. Dyar (USNM), 1 $; San Juan Co., Friday Harbor, 29-V-1906 (Cornell,

Paratype #2085.2), 1$.

S. imitans is very similar to S. dux and may be conspecific; how-

ever, additional specimens of S. dux need to be examined before any

decision can be made.

Schwenkfeldina tridentata (Riibsaamen)

Sciara variabilis Boheman (not Sciara variabilis Zetterstedt, 1838) , 1865, p. 575.

Sciara atrata Holmgren (not Sciara atrata Say, 1824), 1869, p. 366.

Sciara tridentata Riibsaamen, 1898, p. 107 ; Coquillett, 1900, p. 392 ; ,

1904, p. 6; Lundbeck, 1901, p. 312; Nielsen, 1907, p. 371; de Meijere, 1910,

p. 23; Johannsen, 1912, p. 127; Sherman, 1920, p. 15; Edwards, 1922, p. 194;

Vol. 50, No. 2, April 1974

121

, 1923, p. 235; , 1935, p. 532; , 1937, p. 360; Fisher,

1937, p. 413; Bertram and Lack, 1938, p. 50; McClure, 1943, p. 16; Stone and

Laffoon, 1965, p. 236; Steffan, 1966, p. 51.

Sciara validicornis Lundbeck, 1898, p. 243; , 1901, p. 312.

Sciara holmgreni Jacobson (not Sciara holmgreni Riibsaamen, 1894), 1898, p. 34.

Lycoria ( Neosciara ) meigeni, Lengersdorf (not Sciara meigeni Riibsaamen,

1894), 1930, p. 40; Edwards, 1935, p. 532.

Rhynchosciara laguncularis Lengersdorf, 1930, p. 52; Tuomikoski, 1967, p. 46.

Neosciara tridentata, Pettey, 1918, p. 321; Cole, 1921, p. 170; Malloch, 1923,

p. 180.

Lycoriella (Hemineurina) atratci, Frey, 1942, p. 36.

Brady sia ( Neosciara ) incisiforceps Frey, 1948, p. 53.

Schwenkfeldina tridentata, Tuomikoski, 1966, p. 137 ; , 1967, p. 45.

Type locality: Greenland.

Distribution: Alaska, Pribilof Islands (Cole, 1921, “specimen .... near tri-

dentata Riibs.”). Arctic: Bear Island (Edwards, 1935); Jan Mayen Island

(Edwards, 1923); Spitzbergen (Edwards, 1922; Tuomikoski, 1967). British

Columbia (Coquillett, 1900) ; Greenland (Riibsaamen, 1898; Lundbeck, 1898,

as Sciara validicornus) . Manitoba (McClure, 1943).

S. tridentata is apparently a Holarctic species primarily found in the

Nearctic region with extension of its range to arctic islands east and

north of Greenland.

Literature Cited

Beling, T. 1886. Beitrag zur Metamorphose der Zweifliigler in Gattung Sciara

Meig. Wiener Entomol. Z., 5: 93-96.

Bertram, G. C. L. and D. Lack. 1938. Notes on the animal ecology of Bear

Island. J. Anim. Ecol., 7: 27-52.

Boheman, C. H. 1895. Spetsbergens Insekt-fauna. Ofver. K. Svenska. Vetensk.

Akad. Forhandl., 22: 563-577.

Cole, F. R. 1921. Diptera from the Pribilof Islands, Alaska. Proc. Calif.

Acad. Sci., 11: 169-177.

Coquillett, D. W. 1900. Papers from the Harriman Alaska Expedition. IX.

Entomological Results (3) : Diptera. Proc. Wash. Acad. Sci., 2: 389-

464. (Also published separately as Harriman Alaska Expedition, Vol.

9, P. 2, 78 pp., in New York, 1904.)

Edwards, F. W. 1922. Results of the Oxford University Expedition to Spits-

bergen, 1921. No. 14. Diptera Nematocera. Ann. Mag. Natur. Hist.,

10: 193-215.

Edwards, F. W. 1923. On the nematocerous Diptera of Jan Mayen Island.

Ann. Mag. Natur. Hist., 11: 235-240.

Edwards, F. W. 1935. Diptera from Bear Island. Ann. Mag. Natur. Hist.,

15: 531-543.

Edwards, F. W. 1937. Diptera, Oxford Expedition to North-East Land. Ann.

Mag. Natur. Hist., 20: 360-362.

122

The Pan-Pacific Entomologist

Fisher, E. G. 1937. A comparative study of the male terminalia of the

Mycetophilidae of Nearctic America. Unpublished Ph.D. Thesis in Cor-

nell University Library. 430 pp.

Frey, R. 1942. Entwurf einer neuen Klassification der Miickenfamilie Sciaridae

(Lycoriidae). Notul. Entomol., 22: 5^14.

Frey, R. 1948. Entwurf einer neuen Klassification der Miickenfamilie Sciaridae

(Lycoriidae) . II. Die nordeuroparschen Arten. Notul. Entomol.,

27: 33-92.

Garrett, C. B. D. 1925. Seventy new Diptera. Cranbrook Courier Print. 16 pp.

Holmgren, A. E. 1869. Bidrag til kannedomen om Beeren Eilands och Spits-

bergen Insekt-fauna. K. Svenska. Vetensk. Akad. HandL, (n.s.)

8: 3-55.

Jacorson, G. 1898. Compte rendu de l’expedition, envoyee par l’Academie Im-

periale des Sciences a Novaia Zemliaen ete 1896. IV. [Zoological ex-

plorations on Novaya Zemlya. Insects of Novaya Zemlya]. Imp. Akad.

Nauk St. Petersburg, Zap., Fiz.-Mat. Otd., 8: 171-244.

Johannsen, 0. A. 1912. The fungus gnats of North America. The Myceto-

philidae of North America. Part IV. Bull. Maine Agr. Exper. Sta.,

196: 249-328.

Johnson, C. W. 1925. Fauna of New England. 15. List of the Diptera or

two-winged flies. Occas. Pap. Boston Soc. Natur. Hist., 7: 1-326.

Johnson, C. W. 1930. A list of the insect fauna of Nantucket, Massachusetts.

Publ. Nantucket Maria Mitchell Ass., 3: 174.

Lengersdorf, F. 1930. Lycoriidae (Sciaridae). In E. Lindner Die Fliegen der

palaearktischen Region. 43: 33-71.

Leonard, M. D. 1928. A list of the insects of New York with a list of the

spiders and certain allied groups. Mem. N. Y. Agr. Exper. Sta., 101:

1 - 1121 .

Lundbeck, W. 1898. Diptera groenlandica. Vidensk. Medd. Naturhist. Foren.

Kj0benhavn 1898: 236-314.

Lundbeck, W. 1901. Diptera groenlandica. II. Vidensk. Medd. Naturhist.

Foren. Kj0benhavn 1900: 281-316.

McClure, H. E. 1943. Aspection in the biotic communities of the Churchill

area, Manitoba. Ecol. Monogr., 13: 1-35.

Malloch, J. R. 1923. Insects, arachnids, and chilopods of the Pribilof Islands,

Alaska. Diptera (except Tipulidae, Rhyphidae and Calliphoridae) .

N. Amer. Fauna, 46: 170-227.

Meijere, J. C. H. de 1910. Die Dipteren der arktischen Inseln. Fauna Arctica,

5: 13-72.

Nielsen, J. C. 1907. The insects of East-Greenland. Medd. Gr0nland, 29: 363-

414.

Pettey, F. W. 1918. A revision of the genus Sciara of the family Myceto-

philidae (Diptera). Ann. Entomol. Soc. Amer., 11: 319-343.

Rubsaamen, E. H. 1898. Zoologische Ergebnisse der von der Gesellschaft fur

Erdkunde zu Berlin unter Leitung Dr. von Drygalski’s ansgesandten

Gronlands-expedition. VIII. Gronlandische Mycetophiliden, Sciariden,

Cecidomyiden, Psylliden, Aphiden und Galium. In R. Leuckart & C.

Chun Eds. Bibl. Zook, 20: 101-119.

Vol. 50, No. 2, April 1974

123

Shaw, F. R. and E. G. Fisher. 1952. Guide to the insects of Connecticut.

Part VI. The Diptera or true flies of Connecticut. Fifth Fascicle:

Midges and gnats [part]. Family Fungivoridae (= Mycetophilidae) .

Bull. Conn. State Geol. Natur. Hist. Surv., 80: 177-250.

Sherman, R. S. 1920. Notes on the Mycetophilidae of British Columbia. Proc.

Entomol. Soc. Brit. Columbia, 14: 12-15.

Steffan, W. A. 1966. A generic revision of the family Sciaridae (Diptera)

of America North of Mexico. Univ. Calif. Publ. Entomol., 44: 1-77.

Steffan, W. A. 1973. Family Scaridae. In M. Delfinado and D. E. Hardy A

Catalog of the Diptera of the Oriental Region. Vol. 1: 464-476. Univ.

Hawaii Press. Honolulu.

Stone, A. and J. J. Laffoon. 1965. Family Sciaridae (Lycoriidae) . In Stone,

et al. A Catalog of the Diptera of America North of Mexico. Agric. Hand-

book No. 271. pp. 229-236.

Tuomikoski, R. 1960. Zur Kenntis der Sciariden (Dipt.) Finnlands. Ann.

Zool. Soc. ‘Vanamo,’ 21: 1-164.

Tuomikoski, R. 1966. Die Stellung von Sciara tridentata Rubs. (Dipt., Sciari-

dae). Ann. Entomol. Fenn., 32: 137.

Tuomikoski, R. 1967. 6. Mycetophilidae and Sciaridae from Spitsbergen, col-

lected by Dr. J. Kaisila in 1965. Ann. Entomol. Fenn., 33: 43-51.

Winnertz, J. 1867. Beitrag zu einer monographic der Sciarinen. Zool. Bot.

Gessell. Wien, 1867: 1-187.

124

The Pan-Pacific Entomologist

Observations On Megasoma with

Behavioral Notes on Some Lamellicorn Beetles

Associated With Sand Dunes

(Coleoptera: Scarabaeidae, Lucanidae)

Alan R. Hardy and Fred G. Andrews

Insect Taxonomy Laboratory

California Department of Food and Agriculture

Sacramento, California 95814

The description of Megasoma sleeperi Hardy (1972) was based upon

four female specimens collected in Joshua Tree National Monument,

California. Since the original description, the senior author has examined

24 additional specimens of both sexes of this species, as well as the

fragments of numerous additional individuals. The male of Megasoma

sleeperi is described below, additional distribution records and a syn-

onymy in Megasoma are presented, as well as observations of unusual

behavior of M. sleeperi and Pseudolucanus mazama (LeC).

Megasoma punctulatus Cartwright

The following specimens have been examined: ARIZONA:

Pima Co.; Madera Canyon VIII-19-1949 (1 2, Los Angeles Co. Museum).

Santa Cruz Co., Pena Blanca, VII-29-1972 (2 2 2 ) ; Madera Canyon, VIII-

22-1971 (1 $,1 2) (Kane and Langston, Colins.)

Megasoma elephas occidentalis Bolivar y Pieltain et al.

Megasoma elephas occidentalis Bolivar y Pieltain et al. 1963:187

Megasoma mexicanus Fischer 1968:139, new synonymy

From an examination of the illustration accompanying the original

description of M. mexicanus , and from the geographical distributional

data (type of M. mexicanus from Colima, paratypes of M. elephas oc-

cidentalis from Colima), the above synonymy is apparent.

Megasoma sleeperi Hardy

(Fig. 6.)

Male: Length 24.8 to 30.5 mm; width 12.5 to 15 mm. HEAD: Clypeal

plane with close, medium punctures, becoming larger pits posteriorly. Vertex

polished, smooth with few scattered medium punctures. Cephalic horn short,

bifurcate, curved posteriorly apically. Prothorax coarsely punctured, punctures

becoming laterally confluent, appearing rugose ; prothorax occasionally lack-

ing posterior marginal bead medially, marginal bead present laterally and an-

teriorly; lateral margin occasionally poorly defined anteriorly, prothorax laterally

smoothly rounded from dorsal to ventral surface, without lateral carina; anterior

The Pan-Pacific Entomologist 50: 124-128. April 1974

Vol. 50, No. 2, April 1974

125

Figs. 1-5. Fig. 1; Area 4.8 km NW Glarnis, Imperial Co., California, showing

advancing dunes (looking S., March). Fig. 2; Coral Pink Sand Dunes area, Kane

Co., Utah, showing advancing dunes (looking S., July). Fig. 3; Tracks of adult

Pseudolucanus mazama on sand. Figs. 4-5; Adult P. mazama burrowing into sand

at base of dead Juniper.

prothoracic angles acute, not spinaform, not produced anteriorly; near eyes.

Elytra with scattered fine punctures interspersed with close, very fine punctures,

polished, shiny; lateral marginal bead, fading apically. Pygidium evenly convex,

closely punctured with uniform, fine punctures, which are seldom confluent;

punctures separated by less than their diameter; with recumbent, pale hair;

ventral marginal bead present. Six visible abdominal sternites; membranous

area visible between ultimate and penultimate segments; ultimate segment

emarginate apically. Body ventrally clothed with long, white hairs; appendages

with long, reddish hairs.

The male of M. sleeperi most closely resembles the male of M. cedrosa Hardy,

from which it can be easily distinguished by the absence of long dorsal pubescence.

M. sleeperi is glabrous dorsally.

The following specimens of Megasoma sleeperi have been examined: CAL-

IFORNIA: Imperial Co.; 4.8 km (3 mi.) NW Glarnis, IX-16-1972, A. R.

Hardy, M. S. Wasbauer, Collrs. (10 2 2, 3 $ $ ) ; same locale, various dates,

numerous dead examples and fragments. Imperial Valley, XI-29-1937 (1 $,

U.C. Berkeley Colin.). Palm Desert-Deep Canyon, VII-30-1970, E. Giesbert

Collr. (1 2, Giesbert Colin.).

The authors have recently made several investigations of insects in

the sand dune habitats in the Southwest. On a number of occasions,

insects not usually associated with sand dunes have been observed to

utilize vegetation killed by the drifting sand.

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The Pan-Pacific Entomologist

Figs. 6-11. Fig. 6; Adult male Megasoma sleeperi as collected on sand. Fig.

7; Dead M. sleeperi being uncovered by wind action in trough of dune (see

text). Figs. 8-11; Pseudolucanus mazama mating activities (see text).

In March and April 1972, a number of dead, mummified adult Mega-

soma sleeperi (Fig. 7), were collected in the troughs of the dunes, near

the leading edge. Many of the specimens were fragmentary and badly

broken, but a few were intact. Some individuals, which were being

uncovered as the sand moved, were in a vertical position, as if they

had died while digging toward the surface.

The area where the Megasoma were observed (about 4.8 km NW

Vol. 50, No. 2, April 1974

127

Glamis, Imperial Co., California) is an interface area, where the sand

meets an area of hardpan. The hardpan is laced with washes that

drain the Chocolate Mountains to the Northeast. During flash flood-

ing, water moves southwest towards the shores of fossil Lake Cahuilla,

(now the Salton Sea), where it encounters the sand hills. The runoff

is retained, creating local pockets of dense desert vegetation (Fig. 1),

which support large populations of otherwise scarce organisms. These

vegetated areas are slowly buried and the plants killed as the sand moves

eastward at from three to five or more feet per year. Apparently,

Megasoma oviposts in the dead or dying trees, probably utilizing Palo

Verde ( Cercidium floridum Benth.), Mesquite, ( Prosopis sp.), or both,

and develops and pupates in the buried logs. Evidently, after emergence,

the adults attempt to burrow to the surface. Many are unsuccessful in

reaching the surface (which may be through as much as 30 feet of sand) ,

die, and are mummified.

On July 6, 1972, the junior author observed similar events involving

Pseudolucanus mazama (LeC.) (Coleoptera, Lucanidae) at the Coral

Pink Sand dunes near Kanab, Kane County, Utah. The area is a Pinyon-

Juniper association in a valley bordered by red sandstone mountains.

First indications of beetle activity were noted at the north end of the

dunes, about 30 minutes before dark. In this area, Juniper Trees

( Juniperus sp.) were being covered by drifting dunes (Fig. 2), which

had killed the smaller trees. Near the bases of the trees and stumps,

which projected from the sand, were found parts of adult beetles

which were still exhibiting signs of life after a recent attack by an

unidentified predator, probably vertebrate.

Several beetles were seen emerging from the fine unconsolidated

sand in the vicinity of projecting stumps and the bases of live trees.

By nightfall, more than 100 beetles were observed on the sand in an

area about 30 meters by 65 meters. First observations were of apparent

random wandering, followed by copulation when males and females

met. Mated pairs were observed struggling and rolling on the sand

(Fig. 8), or with the male mounted while the female continued to move

across the dune (Fig. 9). Males outnumbered females and often con-

gregations of six to eight beetles were found, with three to four males

clustered about a single female (Figs. 10, 11). It appeared that only

the first male mated with the female. A preponderance of males also

occurs in Pseudolucanus placidus Say (Mathieu, 1969:1059).

After approximately one hour, the beetles began to burrow into the

sand, and within another 15 minutes all except a few wandering males

were gone. The beetles usually burrowed into the dunes within 25 cm of

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The Pan-Pacific Entomologist

a stump or tree (Figs. 4, 5), but several entered the sand at distances

of from four to five meters from the nearest exposed juniper. No at-

tempt was made to determine whether there was dead vegetation be-

neath the sand at these remote points of entry.

At the peak of evening activity, living Junipers within 100 meters of

the breeding congregations, but away from the dunes, were inspected for

beetles with negative results. This may indicate that either dead or

dying junipers or sand or both are important to the life history of

P. mazama. Mathieu (1969:1059) indicates that females of P. placidus

use “a typical sandy-soil habitat” in breeding.

The following morning no evidence of beetles was observed except

the tracks of entering and leaving adults (Fig. 3). Several excavations

at the bases of stumps where Fucanids were seen to burrow the previous

night resulted in the location of a few adults, all at least 45 cm. below

the surface.

Fiterature Cited

Bolivar y Pieltain, C., L. Jiminez-Asua and A. Martinez. 1963. Notas sobre

Dynastinae neotropicales con especial referenda a especies Mexicanas

(Col., Scarab.). Ciencia 22(6) : 181-190, 9 figs, 1 pi.

Fischer, Heinz. 1968. Zwei neue Megasoma-arten aus Amerika. Natur-forsch.

Ges. Augsburg, 22:137-142.

Hardy, Alan R. 1972. A brief revision of the North and Central American

species of Megasoma (Coleoptera: Scarabaeidae) . Can. Entomol. 104:

765-777.

Mathieu, Juan M. 1969. Mating behavior of five species of Lucanidae ( Coleop-

tera: Insecta) . Can. Entomol. 101:1054-1062.

Vol. 50, No. 2, April 1974

129

Descriptions and Records of

Clytini from Mexico and Central America

(Coleoptera: Cerambycidae)

John A. Chemsak and E. G. Linsley

University of California, Berkeley

There have been relatively few Clytini described from Mexico and

Central America since the publications of Bates in 1885 and 1892.

Casey (1912), Linsley (1935), and Chemsak and Linsley (1965) added

a few new species and genera, and Fuchs (1963) described a species

of Neoclytus from Colombia subsequently found in Panama. Although

there are additional undescribed species at hand, only a few are de-

scribed at this time in order to make the names available for other

projects. The remainder will be made known when the generic limits

of the Neotropical Clytini are better defined.

Since many species have previously been known only from single

or a few localities, we are presenting additional distribution records for

a number of these.

We wish to thank the National Science Foundation for support

through Grant No. GB-4944X and the following persons and institu-

tions for the loan of material: The American Museum of Natural His-

tory; Canadian National Collection; Cornell University; H. F. Howden;

Kansas State University; Kansas University; and Ohio State University.

Special thanks are due Celeste Green for preparing the illustrations.

Placoclytus, new genus

Head with front short, not carinate, median line deep; palpi with apical seg-

ments dilated; antennae stout, not spinose, scape robust, cylindrical, slightly

shorter than third segment, third longer than fourth, fifth longer than third.

Pronotum rounded, as broad as or broader than long; disk with a well defined

flattened area which is distinctly delimited; prosternum deeply to shallowly

impressed, intercoxal process narrow, not expanded at apex, coxal cavities wide

open behind; mesosternum with intercoxal process gradually declivous in front,

not protuberant; episternum of metathorax very gradually narrowed behind. Elytra

slightly tapering posteriorly; pubescence depressed, forming bands of condensed

yellowish hairs; apices rounded to truncate, unarmed. Legs slender, rather elongate;

hind tibiae rather short, not compressed; hind tarsi with first segment much

longer than two following together.

Type species — Neoclytus championi Bates.

This genus differs from others presently known in the Clytini by the

flattened, dish-like disk of the pronotum. In addition to the type of the

The Pan-Pacific Entomologist 50 : 129 - 138 . April 1974

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genus, championi, two other species may be assigned to the genus:

distortus Chevrolat, and virgulatus, new species.

Placoclytus championi (Bates), new combination

(Fig. 2)

Neoclytus ? championi Bates, 1885, Biologia Centrali- Americana, Coleoptera,

5:302.

Color black, appendages reddish. Pronotum densely clothed with short, yellow-

ish, appressed pubescence; discal flat area moderately coarsely, separately punctate.

Elytra with undulating fasciae yellowish; apices rounded. Body beneath with

patches of dense appressed, yellowish pubescence at sides. Length, 13 mm.

Type locality: Totonicapam, Guatemala, 10,000 feet.

In addition to the type, we have seen one other specimen of this species

from 6 miles E. San Cristobal de las Casas, Chiapas, Mexico, 2 June,

1969, on Pine (H. F. Howden).

Placoclytus distortus (Chevrolat), new combination

(Fig. 3)

Clytus ( Plagithmisus ?) distortus, Chevrolat, 1860, Ann. Soc. Entomol. France,

(3) 8:500.

Neoclytus? distortus, Bates, 1880, Biologia Centrali-Americana, Coleoptera,

5:58; ibid., 1885:302.

This is a large (25 mm.) robust species having the flattened pronotal

disk, stout antennae, and non-costate, unarmed elytra. The ground

color is black, legs reddish, and the fine transverse fasciae of the elytra

are yellow. Bates (1885) noted that this species should not remain in

the genus Neoclytus and characterized it with championi.

Placoclytus virgulatus, new species

(Fig. 1)

Male: Form moderate sized, tapering slightly; integument black, parts of

legs and apices of antennae reddish; pubescence rather sparse, yellowish, black

on elytra except for yellow fasciae. Head with front finely punctate except for

glabrous median triangle; vertex irregularly punctate; pubescence sparse, ap-

pressed; antennae extending about 2 segments beyond elytra, segments from fifth

densely clothed with dark appressed pubescence. Pronotum as long as broad; disk

with flattened area converging toward apex, punctures dense, irregular; pubescence

moderately dense, golden, appressed; prosternum rather deeply impressed, sparsely

Figs. 1-6. Typical elytral patterns of:

1, Placoclytus virgulatus Chemsak and Linsley;

2, P. championi (Bates) ;

3, P. distortus (Chevrolat) ;

Vol. 50, No. 2, April 1974

131

4, Neoclytus personatus Chemsak and Linsley;

5, N. columbianus Fuchs;

6, Xylotrechus durangoensis Chemsak and Linsley.

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pubescent; meso- and metasternum minutely punctate, moderately densely

pubescent, metepisternum with a patch of appressed pubescence at posterior end.

Elytra more than twice as long as broad; subbasal fasciae transverse, post basal

w-shaped, median arcuate, preapical rounded at suture, angulate away from

suture, sutural line pubescent for most of its length; pubescence on dark areas

black, short, appressed; punctures vague, minute; apices subtruncate. Legs

slender; posterior femora not surpassing elytral apices; pubescence pale, sparse,

appressed. Abdomen minutely punctate, rather sparsely pubescent; apex of last

sternite shallowly emarginate. Length, 13 mm.

Holotype male (California Academy of Sciences) from 34 miles E. Morelia,

Michoacan, Mexico, 2 July, 1963 (W. A. Foster).

The pattern of the fasciae of the elytra, shape of the pronotum, and

longer antennae will separate this species from the two others in the

genus.

Neoclytus personatus, new species

(Fig. 4)

Male: Form moderate sized, cylindrical; integument dark brown, antennae

and tibiae paler; pubescence thick, appressed. Head minutely punctate on front,

a little more coarsely on vertex; pubescence fine, yellowish, appressed; antennae

extending to about basal % of elytra, segments from fifth slightly thickened,

scape slender, longer than third segment, third longer than fourth. Pronotum as

long as broad, sides broadly rounded; median line prominently elevated, rather

finely asperate, each side with slightly elevated, sparsely asperate lines; punctures

vague on middle, dense and deep at sides; pubescence yellowish, appressed,

dense, sparser on disk ; prosternum shallowly impressed, moderately clothed

with yellowish recumbent pubescence and long white erect hairs; meso- and

metasternum finely, shallowly punctate, moderately pubescent, episterna with

bands of condensed yellowish pubescence at posterior halves. Elytra more than

2% times as long as broad, very slightly tapering posteriorly; base with two

large obtuse gibbosities behind scutellum; punctures very fine, dense; pubescence

very short, appressed on dark areas, yellowish pubescent fasciae consisting of two

oblique postbasal bands extending away from near suture and vaguely curving

up toward lateral margin, a sutural premedian pair expanding posteriorly

away from suture, apex very broadly clothed with very thick appressed pubescence

which is darker at posterior end, each side directed obliquely back to lateral

margin ; apices subtruncate, outer angles dentate. Legs elongate ; middle and

hind femora bidentate at apices, front femora with a row of small acute teeth along

inside margins, hind pair exceeding elytral apices by almost % of their length;

hind tibiae elongate, compressed, strongly concave along inner side. Abdomen

minutely, densely punctate, densely clothed with appressed pubescence which

becomes sparser toward apex; fifth sternite rounded at apex, lightly emarginate

at middle. Length 11 mm.

Female: Form more robust; antennae with outer segments more thickened;

pronotum with disk inflated, middle with a broad row of transverse asperites,

small asperites dispersed over sides. Abdomen with last sternite narrowly rounded

at apex. Length, 14 mm.

Vol. 50, No. 2, April 1974

133

Holotype male (Kansas State University) from Monteverde, Costa Rica,

10°29' N., 84°50' W., 1400 M., 23 February 1963, in malaise trap (C. W. and

M. E. Rettenmeyer) . One female paratype, same data, 16 February 1963; 3 male

paratypes, same data, 17-21 February 1963 (R. D. Akre) .

The elytral pattern of this species somewhat resembles that of N.

justini Chevrolat from Colombia. The overall aspect of the pattern

of persojiatus is that of a primitive mask especially enhanced by the

darker pubescence at the extreme apex.

The strong basal gibbosities of the elytra, the elytral pattern, and

toothed front femora make this species distinctive.

Neoclytus columbianus Fuchs

(Fig. 5)

Neoclytus columbianus Fuchs, 1963, Senckenberg Biol., 44: 451, pi. 1.

This species was described from Colombia with no other indication

of locality. We have a male and female from Santa Fe, Darien

Province, Panama, 6 June, 1967 (D. M. DeLong and C. A. Triplehorn)

and one male from Barro Colorado Island, Gatun Lake, Canal Zone, 2

April, 1924 (J. C. Bradley).

In addition to the distinctive elytral pattern, N. columbianus is

characterized by the rather prominent spines at the apices of the elytra.

Neoclytus irroratus (LeConte)

Clytus irroratus LeConte, 1858, Jour. Acad. Nat. Sci., Philadelphia, (2) 4:26.

Neoclytus irroratus, Leng, 1887, Ent. Amer., 3:6; Hopping, 1932, Ann. Entomol.

Soc. Amer., 25: 550, pi. 4, fig. 12; Linsley, 1964, Univ. Calif. Pubis. Entomol.,

22: 139, fig. 42.

Rhopalopachys irroratus, Horn, 1894, Proc. Calif. Acad. Sci., (2) 4: 339.

Clytus ( Rhopalopachys ) morosus Chevrolat, 1860, Ann. Soc. Entomol. France,

(3)8: 501.

Rhopalopachys morosus, Bates, 1880, Biologia Centrali-Americana, Coleoptera,

5: 58.

The following are additions to the previously listed Mexican records for this

species: 1 female, Samalayuca, Chihuahua, 24 June, 1947 (D. Rockefeller Exp.,

Michener) ; 1 male, Sta. Barbara, Chihuahua, 6 May, 1947 (C. M. Bradt) ; 1

female, Monterrey, Nuevo Leon, 6 May, 1950 (Hernandez) ; 1 male, V. de

Santiago, Nuevo Leon (A. Salazar) ; 3 females, Jalapa, Veracruz (W. Schauss) ;

1 male, 3 miles NW Cuernavaca, Morelos, 24 May, 1959 (H. E. Evans).

Neoclytus smithi Bates

Neoclytus smithi Bates, 1892, Trans. Entomol. Soc., London, 1892: 164, pi. 6,

fig. 9.

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The Pan-Pacific Entomologist

Known previously from a unique specimen from Rincon, Guerrero, Mexico.

We have 1 female from Jesus Maria, Nayarit, 27 July, 1955 (B. Malkin).

Neoclytus justini Chevrolat

Neoclytus justini Chevrolat, 1861, Ann. Soc. Entomol. France, (4) 1: 382.

Originally described from Colombia, 1 female is at hand from Santa Fe, Darien

Province, Panama, 29 May, 1967 (D. M. DeLong and C. A. Triplehorn).

Ocliraethes nigrescens, new species

Female: Form moderately robust; ground color black, elytra vaguely tinted

with brown; pubescence fine, pale, moderately long, appressed, with scattered

long erect pale setae on the head, pronotum, base of elytra and ventral surface.

Head coarsely confluently punctured above, more finely on front, lower face

and mouthparts shining, punctation fine; antennae not attaining middle of elytra,

second segment longer than broad, fourth segment shorter than third or fifth,

fifth segment longest. Pronotum broader than long; sides rounded, dorsal sur-

face coarsely, contiguously and somewhat confluently punctate, sides finely densely

punctate; pubescence thin, not obscuring surface; pro-, meso-, and metasterna

finely punctate; scutellum very finely punctate, thinly pubescent. Elytra more

than twice as long as basal width ; contiguously and somewhat confluently punctate

basally, very finely punctate apically; apices obliquely truncate. Legs slender,

finely punctate, thinly pubescent, posterior tarsi more than half as long as tibiae.

Abdomen finely densely punctate, thinly pubescent except for a band along pos-

terior margins of first four sternites. Length 13 mm.

Holotype female (American Museum of Natural History) from Palos

Colorados, Durango, Mexico, 8000 ft., 5 August, 1947, David Rockefeller

Expedition (Gertsch) and 15 paratypes, all female, same locality and date, four

collected by Michener, four by Cazier, three each by Schrammel and Gertsch

and one by Spieth. Two additional paratypes, also female, are from near Otinapa,

Durango, Mexico, 13 August, 1953, collected by C. and P. Vaurie.

This species, judging from the females in the type series, is extremely

variable. The pubescence varies from grayish white to bright golden

yellow and the ground color of the elytra from black tinted lightly with

brown to uniformly reddish-brown to reddish-brown broadly clouded

with dark brown basally in the past-scutellar area along the suture, trans-

versely behind the middle, and more narrowly and obliquely subapically

and apically. The antennae may extend over the basal one-fourth of the

elytra or reach nearly to the middle. However, the body length ranges

only from 11 to 13 mm. in the 18 examples at hand.

The species does not appear to be closely related to other described

Ochraethes, and its general appearance, although variable, is quite

distinctive.

Vol. 50, No. 2, April 1974

135

Trichoxys longipes, new species

(Fig. 7)

Male: Form moderately robust, tapering; ground color black, antennal flagella,

tibiae, tarsi and elytra centrally and posteriorly rufotestaceous. Head moderately

finely, densely somewhat confluently punctate on vertex, frons and mandibles,

pubescence white and depressed on lower face, mandibles and underside with

long erect pale cilia intermixed, eyes encircled with dense yellow pubescence

which extends between the upper lobes and between the lower lobes as a yellow

band, pubescence of vertex between eyes fine, appressed, brown with an inter-

mixture of long erect setae; antennae slender, extending a little beyond middle

of elytra, segments finely punctate, thinly pubescent, ciliate at apices, fourth

segment barely shorter than third, distinctly shorter than fifth, eleventh segment

a little longer than tenth. Pronotum wider than long, sides rounded, base some-

what tubular, surface finely, densely, contiguously punctate, clothed with fine

appressed brown pubescence, with scattered long erect pale setae, anterior and

basal margins lined with dense pale yellow pubescence; scutellum margined

with yellow pubescence; prosternum clothed with white pubescence, thinner

at sides; mesosternum clothed with white pubescence, a patch of dense yellow

pubescence on each side, intercoxal process prominent, polished, elevated above

the coxae, vertical in front ; metasternum thinly clothed with white pubescence,

a patch of dense yellow pubescence on each side on anterior margin behind

mesocoxae and a patch along posterior margin on each side in front of metacoxae;

metepisternum densely yellow pubescent ; metacoxae with a dense patch of white

pubescence laterally. Elytra two and one half times as long as basal width,

narrowing posteriorly, suture depressed, apices shallowly emarginate; surface very

finely punctate, ground color black over basal one-third except suture, rufotestaceous

basally along suture and most of apical two-thirds except for an elongate-oval

oblique black area near middle and a brownish-black cloud over apices; pubescence

brown except for a narrow U-shaped, yellow-pubescent band running from suture

near base to lateral margin, a yellow patch in humeral cavity, a dense yellow

band surrounding oblique median dark areas, a thin yellow band along suture

from basal one-fourth to apical one-third and a dense oblique white patch of

pubescence extending from the suture toward the lateral margins narrowing

laterally and posteriorly. Legs slender; posterior femora extending well beyond

apices of elytra; posterior tarsi slightly more than half as long as tibiae. Abdomen

thinly clothed with white pubescence, with a dense yellow triangular patch of

yellow pubescence at sides of first two stemites; fifth sternite emarginate. Length

12 mm.

Female: Antennae extending over basal one-fourth of elytra, outer segments

abbreviated, thickened; fifth abdominal sternite entire. Length 13.5 mm.

Holotype male, allotype (California Academy of Sciences) and two male para-

types from Guelatao, Oaxaca, Mexico, elev. 1800 m., 15 November, 1964.

This species differs from all others presently assigned to Trichoxys

in the long hind legs with the femora surpassing the elytral apices and

the unique pattern of elytral coloration and pubescence (Fig. 7). It is

not closely related to any other described form.

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The Pan-Pacific Entomologist

Fig. 7. Trichoxys longipes Chemsak and Linsley,

Vol. 50, No. 2, April 1974

137

Xylotrechus durangoensis, new species

(Fig. 6)

Female: Form moderate sized, cylindrical; integument dark reddish brown;

pubescence rather sparse, white, condensed into patches on pronotum and under-

side, elytra with sparsely pubescent white fasciae. Head with indistinct narrow,

flattened frontal carinae on each side of median line; vertex confluently, rugosely

punctate, concave behind antennal tubercles; palpi with apical segments not

dilated; pubescence rather sparse, short and appressed and long and erect, long

hairs more numerous below eyes and under mouth ; antennae short, attaining

about basal % of elytra, basal segments with a few white appressed hairs toward

base. Pronotum wider than long, sides subangulate; disk impressed on each side

of middle, elevated, covered with short, irregular, transverse carinae, remainder

of surface asperate; white appressed pubescence forming indistinct lines along

apex, base and obliquely up from base around disk to middle, apex with 2

indistinct short patches on each side of middle and middle broadly suffused;

prosternum shining, impressed at apex, transversely rugulose, sparsely pubescent;

meso- and metasternum subopaque at sides, sparsely punctate at middle, metepi-

sternum with a condensed patch of white pubescence posteriorly, appressed

pubescence suffused over middle. Elytra less than 2% times as long as broad;

pale areas connecting and encircling round and oblique areas near suture; punctures

very fine, dense, subopaque; apices obliquely truncate, outer angles obtusely

dentate; scutellum densely white pubscent. Legs rather short; femora finely

rugulose, lacking white appressed pubescence on clavate portions, posterior pair

not attaining elytral apices. Abdomen minutely, sparsely punctate, moderately

densely clothed with white appressed pubescence; last sternite rounded at apex.

Length, 17 mm.

Holotype female (Canadian National Collection) from 24 miles W. la Ciudad,

Durango, Mexico, 2 July 1964 (H. F. Howden).

The pubescent pattern of this species is unlike that of any other North

American Xylotrechus. The indistinct narrow frontal carinae of the head

also make it very distinctive.

Xylotrechus quercus Schaeffer

Xylotrechus quercus Schaeffer, 1905, Bull. Brooklyn Inst. Arts Sci., 1 (7): 163;

Hopping, 1932, Ann. Entomol. Soc. Amer., 25: 544, pi. 4, fig. 1; Linsley, Knull,

and Statham, 1961, Amer. Mus, Nov., 2050: 19; Linsley, 1964, Univ. Calif.

Pubis. Entomol., 22: 108.

One female, Arroyo del Nido, 35 miles SW Gallego, Chihuahua, Mexico, 23

June, 1957 (W. C. Russell) ; 1 male, 1 female, 8 miles W. El Palmito, Sinaloa,

Mexico, 29 July, 1964, on Quercus (J. A. Powell, H. F. Howden) ; 1 female, 8

miles W. El Palmito, July 1972, reared ex. Quercus (J. A. Chemsak) .

Xylotrechus sagittatus subsp.

Three specimens (1 male, 2 females) are at hand from 6 miles NE El Salto,

Durango, Mexico, 10 August, 1947 (D. Rockefeller Exp., Spieth, Cazier, Gertsch) .

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The Pan-Pacific Entomologist

More material is necessary before this population can be linked to X. sagittatus

chiricahuae Chemsak from southeastern Arizona.

Xylotrechus SARTORII Chevrolat

Clytus ( Xylotrechus ) Sartorii Chevrolat, 1860, Ann. Soc. Entomol. France,

(3)8:492, pi. 9, fig. 7; Bates, 1880, Biologia Centrali-Americana, Coleoptera,

5:58.

This species was originally described from “Mirador,” Veracruz and subsequently

recorded from Paso del Macho, Veracruz by Bates. We have a series of 7

specimens (3 males, 4 females) from Chipinque Mesa, 5400 ft., nr. Monterrey,

Nuevo Leon, Mexico, 22 and 30 July, 1963, on Quercus (H. and A. Howden) .

Mecometopus hogei Bates

Mecometopus hogei Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5:56.

Originally described from Paso del Macho, Veracruz. New records from Mexico:

2 males, 2 females, Tamazunchale, San Luis Potosi, 13 July, 1956 (D. H. Janzen) ;

1 male, 20 miles E. Xicotepec, Puebla, 23 June, 1964 (C. D. Johnson).

Clytopsis dimidiaticornis (Chevrolat)

Clytus ( Clytus ) dimidiaticornis Chevrolat, 1860, Ann. Soc. Entomol. France,

(3)8:487.

Clytus dimidiaticornis, Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5:54,

ibid., 1885:299.

Clytopsis dimidiaticornis, Casey, 1912, Memoirs on the Coleoptera, 3:73.

2 females, Pedregal, Distrito Federal, 30 September, 1947 (J. Hendricks) ; 1 male,

4 miles E. Toluca, Mexico, 17 August, 1954 (Univ. Kansas Mex. Expedition) :

1 male, Tlalpam, Mexico, 4 October, 1942 (W. F. Foshag) ; 1 male, 1 female, 11

miles SE San Juan del Rio, Queretero, 21 August, 1965 (C. Slobodchikoff) ; 1 male,

4 miles SW Cuidad Mendoza, Veracruz, 16 August, 1962 (H. E. Milliron) ; 1

male, nr. Espiritu Santo, Rio Verde, Chihuahua, 8 August, 1952 (J. D. Lattin) ;

1 male, Tzintzuntzan, Michoacan, 8 August, 1954 (Ryckman, Spencer, Christian-

son) ; 1 female, 10 miles NW Tamazulapan, Oaxaca, 22 August, 1959 (Menke,

Stange) ; 1 male, 2 females, Oaxaca, August, 1920.

Literature Cited

Bates, H. W. 1879-1886. Biologia Centrali-Americana, Coleoptera, Longicornia,

5: 1-436.

Bates, H. W. 1892. Additions to the Longicornia of Mexico and Central

America, with remarks on some of the previously recorded species.

Trans. Entomol. Soc. London, 1892: 143-183.

Casey, T. L. 1912. Memoirs on the Coleoptera, 3: 1-286.

Chemsak, J. A. and E. G. Linsley. 1965. New genera and species of North

American Cerambycidae. Pan-Pacific Entomol., 41: 141-153.

Fuchs, E. 1963. Neue neotropische Clytini aus der Sammlung des Sencken-

berg-Museums. Senck. Biol., 44: 451-456.

Linsley, E. G. 1935. Studies in the Longicornia of Mexico. Trans. Amer.

Entomol. Soc., 61: 67-102.

Vol. 50, No. 2, April 1974

139

Two New Species of Cerambycidae from Southern California

(Coleoptera)

Frank T. Hovore

Placenta Canyon Nature Study Center

19152 W . Placenta Canyon Road,

Newhall, CA 91321

Edmund Giesbert

9780 Drake Lane

Beverly Hills, CA 90210

The Cerambycid genera Clytus Laicharting and Atimia Haldeman

are each represented in California by four species or subspecies. In

each case, three occur predominantly in the northern and central por-

tions of the state. Two species, Clytus planifrons LeConte and Atimia

confusa (subspecies dorsalis LeConte) have been recorded from

throughout the Pacific coast region, and are the only species in either

genus which are known to occur in California south of the Tehachapi

Mountains. Intensive collecting in recent years in the San Gabriel

and San Bernardino mountain ranges has produced new species in

both genera, one of which, Atimia gannoni, described below, also oc-

curs in the Sierra Nevada as far north as Tuolumne County.

Atimia gannoni, new species

(Figs. 1 & 3)

Male. — Form moderately elongate, robust, only slightly tapering posteriorly; color

black; vestiture coarse, appressed, luteus, with scattered long, erect, pale hairs

on head, pronotum, elytra and undersurface of body. Head densely clothed with

appressed hairs; vertex and frons densely, coarsely punctate, with glabrous, im-

punctate median line; antennae attaining apical fourth of elytra, scape moderately

robust, subconical, more than twice as long as broad, third segment twice as long

as second segment, fourth segment distinctly longer than third, fifth segment longest,

outer segments subequal in length to third segment, cylindrical, not flattened

nor expanded. Pronotum. slightly wider than long, sides very feebly, obtusely

rounded, widest before apex, subapical angles obtusely rounded; surface coarsely,

regularly punctate medially, lateral punctures smaller, confluent to rugose, disk

densely pubescent, pubescence obscuring surface, without impunctate or glabrous

vittae; scutellum longer than broad, rounded posteriorly, densely pubescent. Elytra

finely, irregularly punctate, with scattered large punctures, each with a single long

erect hair, surface densely pubescent except for irregular denuded areas; denuded

areas small, impunctate, pattern variable, usually consisting of an irregular, inter-

rupted subsutural vitta, a postmedian spot and a vague, subapical oblique line;

apices broadly, separately rounded or feebly subtruncate. Legs black, finely punctate,

pubescent; posterior tarsi moderately broad, first segment shorter than follow-

The Pan-Pacific Entomologist 50: 139-144. April 1974

140

The Pan-Pacific Entomologist

Fig. 1 . Atimia gannoni Hovore and Giesbert, male.

Vol. 50, No. 2, April 1974

141

ing two together, second segment about one and one-half times as long as broad.

Abdomen with sternites finely, closely punctate, densely pubescent, fifth tergite

narrowly rounded at apex, fifth sternite feebly truncate or emarginate at apex

(Fig. 3). Length: 7-9.5 mm.

Female.- — Form larger and more robust than male; antennae slightly shorter,

attaining apical one-third of elytra; pronotum with sides slightly more obtuse

than in male; fifth abdominal tergite and sternite broadly rounded or subtruncate

at apex. Length: 8-11 mm.

Holotype male, allotype (California Academy of Sciences) and 54 paratypes

(37 males, 17 females), from Little Mt. Gleason, approximately 5 miles W

Mill Creek Summit, Los Angeles County, California, ex. pupal cells in

CaU.ocedrus decurrens (Torr.) , 21 February 1971 to 27 March 1971 (F. T. Hovore,

M. T. Gannon, E. Giesbert). Additional paratypes: 13 males, 4 females, same

locality as holotype, 27 February 1971 to 6 March 1971, 25 February 1972 to 6

March 1972 (D. G. Marqua, A. E. Lewis) ; 1 male, 2 females, Fallsvale, 6000 ft,

San Bernardino County, California, 1 February 1970 (F. T. Hovore). Also represent-

ing this species but not designated as paratypes: 1 male, Stevenson Creek,

Sierra N. F., Fresno Co., Calif., 27 May 1915, “ Libocedrus decurrens Lot 173”

(R. Hopping Collection, California Academy of Sciences) ; 1 male, Long Barn,

Tuolumne Co., Calif., 5 June 1931 (Van Dyke Collection, California Academy

of Sciences) . Paratypes are deposited in the collections of the following institu-

tions and individuals: California Academy of Sciences; California Insect Survey

Collection, Berkeley; Los Angeles County Museum of Natural History; California

State Department of Agriculture, Sacramento; F. T. Hovore; E. Giesbert; D. G.

Marqua; A. E. Lewis; J. N. Knull; W. T. Tyson; J. Cope.

Discussion. — The short third antennal segment, separately rounded

or feebly truncate elytral apices and lack of pronotal polished areas

readily separate Atirnia gannoni from the described species of North

American Atimia , with the exception of Atimia hoppingi Linsley.

From A. hoppingi , to which it seems most closely related, A. gannoni

may be distinguished by the coarser, denser, luteus, appressed pubes-

cence (grayish-white in A. hoppingi ), glabrous, impunctate median

line on the vertex and frons, and the very feebly truncate or emarginate

apex of the fifth abdominal sternite in the male (broadly, deeply emargin-

ate in male A. hoppingi) . (Fig. 2).

Atimia chinensis Linsley is known to us only from the original de-

scription of the female holotype (Linsley, 1939: p. 76, pi. 14, fig. 3),

which seems to bear a strong superficial resemblance to A. gannoni.

In addition to the considerable geographic separation (the type locality

of A. chinensis is in Chekiang Province, China), females of A. gannoni

differ from the description of A. chinensis in the black instead of dark

brown integument, long erect hairs on the discal surfaces of the pro-

notum and elytra, denser appressed pubescence, and more elongate

scutellum.

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Figs. 2-3: Fig. 2. Atimia hoppingi Linsley. Ventral view, fifth abdominal

sternite and tergite (pubescence and punctation omitted) ; Fig. 3. Atimia

gannoni Hovore and Giesbert. Ventral view, fifth abdominal sternite and tergite

(pubescence and punctation omitted).

Biology . — Atimia gannoni infests the thin outer bark of living or

very recently dead Incense Cedar, most commonly in young trees, rang-

ing from two to ten inches in basal diameter. Infested trees show no

external signs of damage due to the presence of the Atimia. The larvae

construct flat, meandering galleries, filled with fine reddish frass, in

the recently dead layers of bark adjacent to the living inner bark tissues.

The shallow, oval pupal cell is placed vertically, with the exit hole

cut by the larva to the loose outer bark and plugged with a thin wad

of stringy frass. Pupation and transformation probably occur in the

late summer or fall since mature adults have been collected from their

pupal chambers in February and March. The life cycle may require

two or more years, as adults and several sizes of larvae have been found

together in a single infestation. Atimia helenae Linsley has also been

observed overwintering in both larval and adult stages (Frankie and

Jensen, 1971), and from this evidence it has been suggested that A.

helenae also has a two-year life cycle. It is also possible that A. gan-

noni , like Atimia confusa dorsalis (Linsley, 1936, 1939), is double-

brooded in the southern portions of its range.

This species is named for the late Michael T. Gannon, with whose

valuable assistance much of the above material and observations were

gathered.

Clytus chemsaki, new species

(Fig. 4)

Male. — Form elongate, slender; integument piceous to black, legs and antennae

dark brown ; entire surface of body sparsely clothed with erect pale hairs ; pronotum

margined with a band of coarse, appressed, pale yellowish- white hair; pale pubes-

cent pattern of elytra consisting of small, tranverse subhumeral bars, narrow,

deeply arcuate, lunate antemedian fasciae, extending forward to sutural and

Vol. 50, No. 2, April 1974

143

Figs. 4-5: Fig. 4. Clytus chemsaki Hovore and Giesbert, male elytral pattern.

Fig. 5. Clytus planifrons LeConte, male elytral pattern.

lateral margins, transverse postmedian fasciae at apical third and oblique apical

fasciae which extend anteriorly along suture to postmedian fasciae. Head

coarsely, densely punctate, eyes and vertex margined with coarse yellowish-white

hair; antennae extending over basal third of elytra. Pronotum slightly longer

than broad, sides somewhat narrowly rounded, surface closely, coarsely punctate;

mesepisterna and metepisterna clothed posteriorly with a dense patch of appressed

pale hairs; pro-, meso-, and metesterna shining, sparsely clothed with pale ap-

pressed hairs; scutellum densely clothed with pale appressed hairs. Elytra slightly

more than 2 1 /j times as long as broad, parallel-sided, surface shining, closely,

finely punctate, clothed with fine, brownish, appressed pubescence except on

fasciae; apices feebly truncate. Legs slender; femora feebly clavate. Abdomen

with sternites shining, sparsely punctate, clothed with pale appressed pubescence;

fifth sternite subequal in length to fourth, apex narrowly, evenly rounded. Length:

8-10 mm.

Female. — Form more robust than male, antennae slightly shorter; legs and basal

antennal segments testaceus; pale appressed pubescence yellow; pubescent bands

on pronotum wider than in male; undersurface of body more densely pubescent

than in male; pronotum more broadly rounded at sides; abdomen with fifth

sternite slightly longer than fourth, apex broadly, evenly rounded. Length: 7-11.5

mm.

Holotype male, allotype (California Academy of Sciences) and 12 paratypes

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(2 males, 10 females), from Mt. Wilson Road, 5000 ft, Los Angeles County,

California, 16 to 18 June 1971 (F. T. Hovore, E. Giesbert) . Paratypes are de-

posited in the collections of the following institutions and individuals: California

Insect Survey, Berkeley; Los Angeles County Museum of Natural History; F. T.

Hovore; E. Giesbert.

Discussion. — Clytus chemsaki will key to C. planifrons LeConte

(Fig. 5) in Linsley ’s (1964) key to North American Clytus. From

C. planifrons it may be immediately distinguished by the narrower

elytral fasciae, the apical elytral fasciae which are directed anteriorly

along the suture, typically forming a “T” with the postmedian fasciae,

the darker appendages, and the smaller, less robust form. From

Clytus pacificus (Van Dyke), to which it seems morphologically

closest, and the Californian species, C. blaisdelli Van Dyke and C.

clitellarius (Van Dyke), C. chemsaki may be separated by the narrow,

strongly arcuate antemedian elytral fasciae, which are directed anteriorly

at the sutural and lateral margins, the configuration of the apical mark-

ings, the dense bands of appressed pubescence on the fourth and fifth

abdominal sternites in both sexes, and the pale yellowish-white appressed

pubescence in the male.

Biology. — The type series of Clytus chemsaki was collected in the

late afternoon from the blossoms of Ceanothus sp. on the steep north

slope of Mt. Wilson in the San Gabriel Mountains. The larval host for

this species is not yet known, but it probably infests dead branches of

coniferous trees.

Acknowledgments

We wish to extend our gratitude to Dr. John A. Chemsak and Dr. E.

Gorton Linsley, University of California, Berkeley, for reviewing the

manuscript; to Dr. Paul Arnaud, California Academy of Sciences, for

the loan of comparative material; and to D. G. Marqua and A. E. Lewis

for the use of specimens from their personal collections.

Literature Cited

Frankie, G. W & G. L. Jensen. 1971. Studies on the distribution and biology

of Atimia helenae Linsley on two California Cupressus species. Pan-

Pac. Entomol., 47 : 287-293.

Linsley, E. G. 1936. Observations on the habits of some western longhorn

beetles. Pan-Pac. Entomol., 12: 199.

Linsley, E. G. 1939. The longicorn tribe Atimiini. Bull. S. Calif. Acad. Sci.,

38: 63-80.

Linsley, E. G. 1964. The Cerambycidae of North America, Pt. V ; Taxonomy

and classification of the subfamily Cerambycinae, tribes Callichromini

through Ancylocerini. Univ. Calif. Publ. Entomol., 22: 1-197.

Vol. 50, No. 2, April 1974

145

New North American Tabanidae XXI.

Another New Bolbodimyia from Mexico

(Dipt era)

Cornelius B. Philip

California Academy of Sciences,

San Francisco 94118

AND

Linda Floyd

Californa State University,

San Francisco 94132

Bolbodimyia comprises a New World genus of strikingly marked

species mostly distributed in the northern Neotropical Region. There

are rare Nearctic records of B. atrata (Hine) in Arizona as far north

as the type locality in famed Oak Creek Canyon. This is the only

species that has been reared (Burger in Goodwin and Murdock, 1974).

Philip (1954) described the male of B. atrata from Jalisco, Mexico.

Species of the genus, revised by Stone (1954), are never taken in

large numbers as adults, which suggests peculiar habits. Since the

unique type male of the brightly marked new species described below

was taken not far below the arbitrarily established northern neotropical

boundary in Chihuahua, it is possible that a more northern occurrence

of this species will eventually be discovered.

The present specimen was among miscellaneous Mexican tabanids

kindly submitted for study by Dr. R. 0. Schuster of the University of

California, Davis, where the type will be on deposit.

Bolbodimyia lampros, new species

(Fig. 1)

A robust, contrastingly orange and black species with orange pollinose cheeks

and face; pleura orange with concolorous pile; legs blackish brown, tibiae swollen;

wings evenly fumose with reduced apical hyaline crescents.

Holotype male. — length 15.0 mm; wing length 11 mm. Head subhemispherical,

eyes bare, facets in upper two-thirds enlarged, dull tan, small facets black, extended

up occipital margin to vertex. Tubercle in occipital notch small and moderately

depressed. Frontal triangle bare, dull grayish pollinose in apex, mostly shining,

dark brownish black and swollen around antennal fossae (Fig. la) . Cheeks slightly

swollen and face only moderately depressed, entirely orange pollinose with con-

colorous hairs below. Scapes greatly enlarged, a little more so below, shiny and

dark mahogany to black with sparse black hairs; flagella reddish, the styles about

one-half lengths of plates; latter rather slender with obtuse dorso-basal angles.

Palpi porrect, the basal segments and hairs orange; apical segments thicker,

ovoid, brown with orange and black hairs (Fig. lb).

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146

The Pan-Pacific Entomologist

Fig. 1. Bolbodimyia lampros, male. a. Profile of head and antenna show-

ing protuberant frontal triangle around antennal fossae and swollen scape, b.

Palpus, c. Outer wing with reduced apical hyaline crescent.

Unlined notum and scutellum subshiny black with sparse concolorous hairs;

chest and vestiture similar. Pleura and prealar tubercles bright orange with

concolorous pile. Legs dark mahogany to black, mostly black-haired, some yellow

hairs basally on coxae. Wings evenly fumose with hyaline crescent restricted to

apex between veins R+ and R 5 (Fig. lc) .

Abdomen almost entirely bright orange, with short coarse black hairs dorsally;

venter similarly colored but with orange hairs. Edges of dorsum with a narrow

black line and tufts of black hairs.

Type Locality. — Mexico: Chihuahua, Cuiteco, 30 August 1969. T. A. Sears, R. C.

Gardner, and E. C. Glasser.

This new species is one of the more robust representatives of the

genus. It is likely that the unknown female will show sexual dichroma-

tism, familiar to that found in some other species in the group. Though

B. lampros is structurally somewhat similar to B. atrata and B. celeroides

Stone, it differs from both in having the scape less produced below and

tibiae a little less inflated. Both sexes of the former are entirely black,

but only the females of celeroides from Peru are available for com-

parison; the wide geographic separation as well as the tinctorial dif-

ferences in the abdomens and flagella reinforce the doubt that B. lampros

could be the dichromatic male, unless unexpected intervening intergrada-

tion is eventually discovered. Presumably, the still unknown male of

B. dampfi Philip from Mexico will be distinguished at least by shining

black face and cheeks. It may be noted that Fairchild (1964) reported

the male of B. philipi Stone as taken “at light” in Panama.

The name lampros (Gr., lamp or lantern) is assigned for the bright,

almost luminous, contrasting orange abdomen.

Vol. 50, No. 2, April 1974

147

References

Fairchild, G. B. 1964. Notes on Neotropical Tabanidae (Diptera) IV. Further

new species and new records for Panama. J. Med. Entomol., 1: 169-

185.

Goodwin, J. T. and W. P. Murdoch. 1974. A study of some immature Neo-

tropical Tabanidae (Diptera). Ann. Entomol. Soc. Amer., 67: 85-133.

Philip, C. B. 1954. New North American Tabanidae (Diptera) Part V. A

striking new Bolbodimyia from Mexico. Wasmann J. Biol., 12: 29-33.

Stone, A. 1954. The genus Bolbodimyia Bigot (Tabanidae, Diptera) . Ann.

Entomol. Soc. Amer., 47 : 248-254.

148

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Revision of the Scarabaeidae : Anomalinae

1. The Genera Occurring in the United States and Canada

( Coleoptera )

Robert W. L. Potts

California Academy of Sciences, San Francisco, California

It appears necessary in a consideration of the generic arrangement

of this subfamily that we remember that taxonomic divisions above the

species are necessarily artificial, with no objective existence, and are

formed largely as a matter of convenience. Nonetheless, an organized

taxonomy demands a certain stability based on the work of previous

authors, and while their arrangements cannot be ruled sacrosanct

merely on a basis of age or previous acceptance, such factors should not

be lightly dismissed.

In the Anomalinae we find an exceptionally large and remarkably

interrelated group of species. Except for a relatively few genera, with

some that may be intermediate to adjacent subfamilies, the Melo-

lonthinae and Rutelinae, it is often difficult or impossible to separate

genera in a manner consistent with the usual practice of assembling

species only when they share several characters reasonably disparate to

the species assembled in other genera. Despite a number of variable

and distinctive characters, intermediates are the rule, not the exception.

A veritable wonderland of generic and subgeneric names have been

proposed in the subfamily. A strict application of what most systematists

would consider necessary to a valid generic grouping would synonymize

a majority of these into the single genus Anomala, with thousands of

species, worldwide, and dozens, if not even hundreds of secondary

homonyms.

This, most certainly, would not be more convenient, and it would

scarcely contribute to the ease of identification. Compromise, with a

minimum of violence to a few questionable, but nonetheless long-estab-

lished and widely-used genera probably becomes necessary. However,

it is also obvious that a number of the proposed groupings were without

true value from their inception, merely following in the mould of an

early-day taxonomy that regarded any large genus as unmanageable

until infinitely divided, no matter how precariously.

We believe the following best represents the status of the Anomalinae

of our area:

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Vol. 50, No. 2, April 1974

149

Tribe Anomalini

AnOMALA Samouelle 1819:191. Type species: Scarabaeus aeneus DeGeer,

1774, Arrow designation, 1917:126.

syn. Anomalepta Casey 1915:8. Type species: Anomala semilivida LeConte,

1878, Machatschke designation, 1957:30.

syn. Anomalopus Casey 1915:40. Type species: Anomala rhizotrogoides

Blanchard, 1850, Casey designation, 1915:40.

syn. Blitopertha Reitter 1903:85. Type species: Melolontha lineata Fabricius,

1798, Arrow designation, 1917:127.

syn. Callistethus Blanchard 1850:198. Type species: Mimela auronitens

Hope, 1835, Arrow designation, 1917:127.

syn. Hemispilota Casey 1915:45. Type species: Melolontha lucicola Fabri-

cius, 1798, Machatschke designation, 1957:30.

syn. Oliganomala Casey 1915:38. Type species: Strigoderma exigua Schwarz,

1878, monobasic.

syn. Pachystethus Blanchard 1850:201. Type species: Popillia vidua Newman,

1838, monobasic.

syn. Paranomala Casey 1915:12. Type species: Melolontha binotata Gyllen-

hal, 1817, Machatschke designation, 1957:31.

syn. Phyllopertha Stephens 1830:223. Type species: Scarabaeus horticola Lin-

naeus, 1758, Arrow designation, 1917:126.

syn. Rhombonalia Casey 1915:5. Type species: Anomala cavifrons LeConte,

1867, Machatschke designation, 1957:31.

syn. Rhombonyx auctorum (not Rhombonyx Hope 1837:106). Type species:

Melolontha holosericea Fabricius, 1787, Arrow designation, 1917:126.

syn. Spilota Burmeister 1844:266. Type species: Melolontha marginata Fab-

ricius, 1792, Arrow designation, 1917:126.

Only the synonyms with application in our area are included; many

others are necessary in other sections of the world. All have been

synonymized previously, however several are still accepted and in present

use by some systematists.

Blitopertha Reitter and the much older Phyllopertha Stephens must

be included because of our introduced species, Anomala orientalis

Waterhouse, which is variously placed in one or the other. Other

species included in the genera are Asiatic and European, though

species from Mexico and Central America have been included. Both

genera were established on a combination of characters that have been

subject to a confusing emendation and change by almost every author

that has used them. No single character is common to all species

usually included, and all are found duplicated in species not included,

where they are sometimes carried to an even greater extreme. To com-

pound this confusion, most authorities have failed to agree on their

lists of species properly belonging in the two genera.

Our introduced species, orientalis , does not differ in any material

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way from our native species, and can be separated on characters of no

more than species-level significance.

Spilota Burmeister, Callistethus Blanchard and Pachystethus Blanchard

are all largely based on the development of a prominent intermesocoxal

process. This might have a certain validity for the species with the

most extreme development, except that there is no place where an arbi-

trary limit might be placed logically in a perfectly intergraded series.

Moreover, the character also appears in some Strigoderma and in species

placed in Epectinaspis. The three genera have also been subject to

varying application and have never had universal acceptance.

The type species of Callistethus is of Himalayan origin, and Ameri-

can species probably should never have been assigned to the genus,

however Machatschke, 1957:93, somehow, and quite mistakenly gave

Melolontha marginata Fabricius as the type, crediting the designa-

tion to Arrow 1917:127, whereas Arrow actually designated Mimela

auronitens Hope as the type, and designated Melolontha marginata

Fabricius as the type of Spilota , which immediately preceded Callistethus

in his list. But Machatschke, 1957 :92, also gives marginata as the type

of Spilota , thus incorrectly indicating the same species as the type of

both genera.

Among our own species sometimes variously placed in one or another

of these three genera, such as marginata Fabricius, the intermesocoxal

character is quite variable, from a prominent boss to a large porrect

process, and selected specimens might be fitted into each of the three

genera.

Unfortunately, this forces the change of the name of a distinctive

Florida species described by Mark Robinson, his name becoming a

secondary homonym of floridana Casey 1915:33. We propose:

Anomala robinsoni, new name, for Pachystethus floridana

Robinson 1941:133.

A majority of the Casey names were proposed as subgenera, almost

entirely on characters of merely species-level importance, and admit-

tedly in an attempt to fraction a large genus. But whether as genera

or subgenera, merely adding species from no greater distance than

Mexico and Central America immediately demonstrates the lack of

validity of his groupings.

Inexplicably Oliganomala Casey was misplaced as a synonym of

Strigoderma by Machatschke 1957:139 and 1972:216, possibly on the

basis of the fact that the unique species, exigua, equally inexplicably was

described as a Strigoderma by Schwarz, although it met none of the key

Vol. 50, No. 2, April 1974

151

criteria. Casey, 1915:38, made it clear it definitely was not a Strigo-

derma, but the characters cited are not of sufficient importance to

justify its separation from Anomala.

The genus Rhombonalia Casey has had a certain acceptance, however,

and for some time we struggled to maintain the concept, since it is based

on a notable character, the simple protarsal claw, and does include an

obviously closely-related group of species. But, it would necessarily

need enlarging to include other species with the same character and

eventually breaks down in a way similar to the case of genera pre-

viously discussed. The simple protarsal claw is highly variable and there

is no point in an intergrading series where it may be established. More-

over, it was found that even cavifrons LeConte, the type, may have

distinctly cleft claws not too uncommonly, and that the simple claw

occurs occasionally in many of our species. It is moderately com-

mon in one species that rather obviously should not be assigned to the

genus. Finally, a species that is very closely related to cavifrons in

its other major characters almost always has cleft claws. Other char-

acters that have been cited for the genus proved valueless, and it clearly

becomes necessary to accept the judgment of those who had already

reduced the name to the synonymy.

STRIGODERMA Burmeister 1844:310. Type species: Strigoderma sulcipennis

Burmeister, 1844, Casey designation, 1915:48.

syn. Alamona Casey 1915:61. Type species: Alamona parviceps Casey, 1915,

Casey designation, 1915:61.

syn. Strigodermella Casey 15:59. Type species: Melolontha pygmaea Fab-

ricius, 1798, monobasic.

Considering the entire genus and its separation from worldwide

Anomalinae, the definition is far from satisfactory, with no single

character that is not duplicated in other genera, both in the Americas

and in the Old World. Placement in the genus depends on general

habitus almost as much as on any distinct character, while the assign-

ment of a number of intermediate species is entirely arbitrary. Also,

several Asiatic species would have to be included in the genus by any

definition except distribution. Nonetheless, in our area the genus is

easily distinguished on the characters noted in the key. It has had a

very long-standing acceptance in the literature, with many species de-

scribed in the genus, and it would only create greater confusion to

return it to Anomala , regardless of its weakness.

However, the Casey attempt to split the genus with two additional

genera is indefensible at any level. The small justification the genera

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had when applied to our few species disappears entirely with the addi-

tion of species from south of our borders.

Alamona has been misplaced by Blackwelder, 1944:242, as a sub-

genus and synonym of Anomala , but does not belong with that series

at all, being a typical Strigodermid.

Tribe Anisopliini

AnOMALACRA Casey 1915:10. Type species Anomalacra cuneata Casey,

1915, monobasic.

While this genus has been synonymized with Anomala by

Machatschke, 1957:30 and 1972:83, and there has been confusion as

to the species belonging in the group, in the present state of our

knowledge it is a very distinct and valid genus. It is closely related

to the tropical Callirhinus, with the clypeus remarkably thinned and

the labrum reduced and withdrawn posteriorly. These characters are

also common to the Old World Anisopliini, to which Tribe it would

necessarily be assigned.

Charles Schaeffer described our first North American species as

Anomala clypealis , and despite its remarkable differentiation from

other Anomala the species is still found assigned to that genus in all

published lists, but it must be reassigned to Anomalacra.

Tribe Popilliini

POPILLIA Serville 1825:367. Type species: Trichilis bipunctata Fabricius,

1787, Arrow designation, 1917:58.

Represented in our area by the single introduced species, Popillia

japonica Newman, although the genus is large and diverse. It is

primarily an Asiatic and African development, and has elements of be-

ing parallel to the Strigoderma development in the Americas. It is

generally given a separate Tribal status, however, this may be question-

able as strong intergradation to the Anomalini certainly exists.

Tribe Lepothopliini, new Tribe

LEPTOHOPLIA Saylor 1935:132. Type species: Leptohoplia testaceipennis

Saylor, 1935, monobasic.

This genus is apparently intermediate to the Melolonthinae where it

was originally placed as belonging with the Hopliini. Disagreement

as to its relationship still exists, however we believe it belongs in the

Anomalinae, where it was reassigned by Howden and Hardy, 1971:337.

Vol. 50, No. 2, April 1974

153

The very distinctive tumid and penicillate ligula; the exceptionally

thick anterior face of the clypeus, some two or more times as thick

as normal in the Anomalinae; the obsolescent epipleurae; and the

reduction of the minor protarsal claw to half or less the length of the

the major claw indicate a disparity sufficient for the erection of a

separate Tribe. Withal the major differences, the genitalia are of the

generalized anomaline type, closely similar to Anomala binotata and

flavipennis.

Key to the Genera of Anomalinae of the

United States and Canada

1. Ligula and undermouth parts essentially glabrous, with no more than a

a few distant hairs, or, rarely, with some close-set flexuous hairs pos-

teriorly 2

Ligula penicillate, with thick erect bristles; California (Imperial Val-

ley) Leptohoplia

2. Clypeus anteriorly thickened, with apical face more or less vertical, round-

ing under or sloping forward to distinct labrum beneath 3

Clypeus, as viewed from side, apically reflexed and thinned, without thick-

ened anterior face; labrum withdrawn and not visible; Arizona Anomalacra

3. Basal pronotal margin posteriorly rounded or sinuate, always projecting

backward over scutellum 4

Scutellum projecting forward into an arcuate emargination of basal pronotal

margin ; Eastern States, 1 introduced species Popillia

4. Mesosternal epimera antero-dorsally raised and swollen, narrowly projecting

before elytral humeri ; pronotum subquadrate, distinctly narrower than

elytra; elytra more or less tapering posteriorly from near humeri .... Strigoderma

Epimera reduced, obliquely rounding under covering humeri; pronotum

obviously broader than long, as broad as elytra or nearly; elytra widest near

middle or toward apex Anomala

Literature Cited

Arrow, G. J. 1917. Fauna of British India, Coleoptera Lamellicornia, Vol. 2.

Blackwelder, Richard E. 1944. Checklist of the coleopterous insects of Mexico,

Central America, The West Indies and South America, Part 2, U. S.

Nat. Mus. Bulletin 185, pp. 189-341.

Blanchard, Emile. 1850. Catalogue de la collection entomologique Coleopteres,

Musee d’histoire naturelle de Paris.

Burmeister, Hermann C. C. 1844. Handbuch der Entomologie, Berlin, Vol. 4,

Pt. 1.

Casey, Thos. L. 1915. Memoirs on the Coleoptera, Vol. 6, A Review of the Ameri-

can species of Rutelinae, Dynastinae and Cetoniinae.

Hope, Frederick W. 1837. The Coleopterist’s Manual, Vol. 1.

Howden, H. F. and A. R. Hardy. 1971. Generic placement and adult behavior

of the genus Leptohoplia Saylor. Proc. Entomol. Soc. Wash., 73: 337-

341.

154

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Machatschke, Johann W. 1957. In Wytsman, Genera Insectorum, Fasc. 199,

Coleoptera Lamellicornia, Family Scarabaeidae, Subfamily Rutelinae.

Machatschke, Johann W. 1972. Coleopterorum Catalogus Supplementa, Part

66, Fasc. 1, Scarabaeoidea: Melolonthidae, Rutelinae.

Reitter, Edmond. 1903. Yerh. Nat. Ver. Brunn, 41: 85.

Robinson, Mark. 1941. Studies in the Scarabaeidae of North America, Trans.

Amer. Entomol. Soc., 67: 127-136.

Samouelle, George. 1819. The entomologists useful compendium, etc., London.

Stephens, James F. 1830. Illustrations of British Entomology, Mandibulata,

Vol. 3.

RECENT LITERATURE

Frederick Valentine Melsheimer, Parent of American Entomology. Robert

Snetsinger, 86pp. (Melsheimer Entomological Series, No. 12.), 1973. Available

from the Entomological Society of Pennsylvania, 106 Patterson Building, Uni-

versity Park, Penna. 16802. $3.75 plus $.25 for mailing.

Although most Americans prefer to consider Thomas Say as the male parent of

entomology in the United States, F. V. Melsheimer certainly played a prominent

role in the early history of the science. This volume contains a brief biography,

followed by a facsimile reproduction of Melsheimer’s catalogue of the insects

of Pennsylvania.- — Editor.

Vol. 50, No. 2, April 1974

155

Behavioral and Morphological Mimicry in a

Cranefly and an Xchneumonid

C. N. Slobodchikoff

Department of Biological Sciences

Northern Arizona University, Flagstaff 86001

Field observations in Costa Rica (Slobodchikoff, unpublished),

Mexico (P. Arnaud, personal communication), and Arizona (J. Wick,

personal communication) suggest that mimetic relationships between

craneflies and hymenopterans are fairly common. However, apparently

such relationships have not been described (C. P. Alexander, personal

communication) nor have such relationships been subjected to experi-

mentation to determine whether Batesian or Mullerian mimicry is in-

volved. The purpose of this paper is to describe an apparent mimetic

situation between a crane fly, Ctenophora vittata angustipennis Loew,

and an ichneumonid wasp, Therion circumflexum (L) .

The observations reported here were made at Alpine Lake, Marin

County, California, during May of 1971-1973. The vegetation sur-

rounding Alpine Lake may be described as mixed coniferous forest

interspersed with small patches of oak-grassland. Occasional streams

are present, draining into Alpine Lake. The oaks in the oak-grassland

are primarily live oaks, Quercus agrifolia. Both the cranefly and the

ichneumonid are found mainly around the oak trees. Mimicry in these

two species apparently involves two factors : similarity in flight behavior

and similarity in coloration.

Flight behavior in Therion circumflexum has been described by

Slobodchikoff (1973). Female T. circumflexum fly comparatively

slowly, at a speed of approximately 10 cm/sec, about one meter above

the top of grass stems in shady areas underneath oak trees. Males, on

the other hand, fly twice as fast as females, approximately 20 cm/sec,

around the perimeter of the upper three-quarters of the oak canopy.

When the male and female ichneumonids fly, their third pair of legs

are stretched out 20 degrees below the horizontal plane of the body, their

abdomen is held up 20 degrees above the plane of the body, the femora

and tibiae of the first pair of legs are pressed close to the thorax, and

the femora of the middle legs are pressed to the thorax while the

tibiae of the middle legs are held out away from the body. Both the

male and female fly with their antennae extended forward.

Male and female craneflies exhibit the same flight patterns as the

ichneumonid males. The craneflies fly around the perimeter of oak

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canopies, at a speed equivalent to that of male ichneumonids. Males

and females both fly with their hind legs stretched out downward and

back, and with the abdomen elevated 15 to 20 degrees above the horizon-

tal plane of the body. The middle femora are pressed to the thorax,

while the middle tibiae are held away from the body. The front femora

are kept pressed to the thorax, while the front tibiae are extended for-

ward, beyond the head, apparently simulating the ichneumonid’s anten-

nae. In flight, the craneflies are very difficult to separate from the

ichneumonids. Apparently, neither male nor female craneflies mimic

the flight behavior of female ichneumonids. No Ctenophora have been

observed flying at the speed or the level of the female Therion.

The similarity in coloration of the ichneumonid and cranefly is very

marked. Both the Therion and the Ctenophora have a yellow, orange,

and black overall color pattern. The ichneumonid (fig. 1) has a black

thorax, with yellow scutellum dorsally and irregular orange splotches

laterally. The head is black, with long orange antennae. The abdomen

is orange, with an incomplete black stripe dorsally, and the three termi-

nal abdominal segments are entirely or mostly black. The hind and mid-

dle coxae are black, the front coxae are yellow. The hind femora

and tibiae are yellow in their proximal 0.7, black in their distal 0.3.

The middle and front femora and tibiae are yellow, as are all tarsal

segments.

The cranefly (fig. 2) has a black thorax, with faint yellow in the

scutellar region and irregular yellow splotches laterally. The head is

black, the short antennae are orange. The abdomen is orange, with

a longitudinal black stripe dorsally and a reddish-black longitudinal

stripe laterally. The basal third of the first abdominal segment is black,

creating the illusion of a constriction or petiole when the abdomen is

seen in side view against a dark background. The terminal three visible

segments of the abdomen are reddish black. All coxae are black, and

all trochanters are yellow. The distal ends of the trochanters of the

first pair of legs are pointed dorsally, assuming the same position in

flight as the yellow coxae of the first pair of legs of Therion. All

femora and tibiae are yellow-orange in their proximal 0.8, and black

in their distal 0.2. When seen against a dark background, such as the

dark green of oak leaves, the black on the legs creates the illusion that

the legs are shorter than they are in actuality. Even though all the

cranefly ’s legs are longer than the ichneumonid’s legs, the cranefly ’s

legs in flight appear to be no longer than those of the ichneumonid.

The first tarsal segment of each leg is orange, the remaining tarsal

segments are blackish-orange. This also contributes to the illusion of

Vol. 50, No. 2, April 1974

157

Fig. 1. Male T. circumflexum, lateral view, shown in position of flight.

Fig. 2. Female C. vittata angustipennis, lateral view, shown in position

adopted during flight.

shorter legs in flight. Since the orange front tibiae are held out in

front of the cranefly in flight, an illusion of orange ichneumonid an-

tennae is created.

The geographical distribution of the cranefly coincides with the

distribution of the ichneumonid. Ctenophora vittata angustipennis has

been recorded from British Columbia, Washington, Oregon, and Cal-

ifornia (Stone et al., 1965). Although T. circumflexum occurs through-

158

The Pan-Pacific Entomologist

out Canada and the United States in transition life zones, 19 percent (31

localities) of the total localities (163) from which T. circumflexum

has been recorded occur in the Pacific Northwest (Slobodchikoff,

1971).

Neither the Therion males nor the Ctenophora are present in very

large numbers. During May 1971 and May 1972 males of Therion

and both sexes of the Ctenophora were captured, marked with enamel

paint, and released. Marking and subsequent attempts at recapture were

carried out for 30 days in May 1971, and 21 days in May 1972. A

total of 32 male Therion were captured, marked, and released in 1971,

and 27 were captured, marked, and released in 1972. Seven Ctenophora

were captured, marked, and released in 1971, and 15 were captured,

marked, and released in 1972. In this two year sampling period,

Therion represented 73 percent of the total number of ichneumonids

and craneflies caught, while Ctenophora represented 27 percent of the

total captures. Each year, after the first 5 days of sampling, more

than 50 percent of the Therion caught on a given day were recaptures

of previously marked individuals. No marked Ctenophora have been

recaptured.

The mimicry may be either Batesian or Mullerian. The lower per-

centage of craneflies present suggests that mimicry may be of the

Batesian type, with the ichneumonid serving as the model. Protection

from predation is probably provided by some distasteful quality. Sup-

porting this conclusion are: 1) male ichneumonids lack an ovipositor

that could be used for defensive purposes; 2) both the ichneumonid

and cranefly have orange, yellow, and black coloration, colors that

frequently have an aposematic function in insects (Rettenmeyer, 1970).

Acknowledgments

I would like to thank Drs. R. Baida and C. Holm, Northern Arizona

University, for their helpful criticisms of this manuscript.

Literature Cited

Rettenmeyer, C. W. 1970. Insect mimicry. Ann. Rev. Entomol., 15: 43-74.

Slobodchikoff, C. N. 1971. Patterns of differentiation and evolution in the

genus Therion (Hymenoptera: Iclineumonidae) . Ph.D. dissertation,

University of California, Berkeley.

Slobodchikoff, C. N. 1973. Behavioral studies of three morphotypes of

Therion circumflexum (L). (Hymenoptera: Ichneumonidae) . Pan-Pa-

cific Entomol., 49: 197-206.

Vol. 50, No. 2, April 1974

159

Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R.

Coulson. 1965. A catalog of the Diptera of North America. U.S.D.A.

Agriculture Handbook No. 276, 1696 pp.

RECENT LITERATURE

The Anisodactylines (Insecta: Coleoptera: Carabidae: Harpalini) :

Classification, Evolution, and Zoogeography. Gerald R. Noonan. Questiones

entomologicae, 9: 266-480. 1973.

This revision contains keys to genera and subgenera, with complete taxonomic

treatment of Anisotarus, Notobia, and Gynandrotarsus. Nearly 50 pages are de-

voted to discussion of the zoogeography and phylogeny of the Anisodactylina. The

application of Hennig’s methods of analysis to a group of insects with no fossil

record and few characters which can intrinsically be considered plesiomorphic

or apomorphic should be of interest to many systematists. — Editor.

160

The Pan-Pacific Entomologist

A New Species of Cyclocephala Latreille

from California Sand Dunes

(Coleoptera: Scarabaeidae)

Alan R. Hardy

Insect Taxonomy Laboratory

Department of Food and Agriculture

Sacramento, CA 95814

In the course of investigations of the sand dune habitat in the western

United States, the following undescribed species of Cyclocephala was

collected from black light in series with Leptohoplia testaceipennis

Saylor, a strikingly similar species.

Cyclocephala wandae, new species

Holotype Male: length 7.3 mm; width (elytral humeri) 3.0 mm. Head rufo-

testaceous to testaceous, rest of body pale flavotestaceous; clypeus with angles

acute, upturned, sides divergent to base, disc glabrous, with scattered medium

punctures, posterior clypeal margin raised above bisinuate suture; front and vertex

with scattered coarse punctures, those of vertex with short erect setae; eye large,

anterior edge visible in front of canthus from dorsal aspect; antennae 9 segmented,

club three segmented, approximately twice as long as basal segments. Pronotum

widest just behind anterior angles; sides straight behind and in front of angle; an-

terior and posterior angles broadly rounded ; marginal line complete except medially

posteriorly; margin broad anteriorly, narrowing laterally and posteriorly; disc

with scattered fine to medium punctures. Scutellum with large shallow scattered

punctures; punctures anteriorly and laterally each with a single posteriorly in-

clined seta. Elytra without easily discernable striae or intervals; surface with

very short setae, setae barely visible (best seen in side view). Pygidium closely,

shallowly rugosely punctate; provided with long, fine, scattered hairs, these hairs

longer and denser basally. Mentum with a dense, anteriorly directed brush made

up of many closely set erect bristles. Anterior tibiae with three, acute, widely

separated teeth on outer margin, basal tooth median; anterior tarsal claws with

larger claw not split at tip, smaller claw reduced to a short stub. Hind tibia with

median carina poorly defined, evident as a dark pigmented, setose, poorly elevated

line, apical marginal carina missing; posterior tarsal claws fine, straight, acute. Male

genitalia: figure 1.

Variation in paratypical series: length: 6.6 mm to 7.5 mm; width 2.7 mm

to 3.0 mm. Generally agreeing with type.

Holotype male (CAS # 11941) and 2 male paratypes, California, Imperial

Co., 3 mi. NW Glamis, IX-15/16-1972, M. Wasbauer and A. Hardy (USNM,

HFH) ; 1 male paratype, California, Imperial Co., Regina, approx. 20 mi. SE

Niland, VII-4-1971, Sand Dunes, Hardy et al. Collrs. (ARH). Paratypes are de-

posited in the following collections: USNM, Howden and that of the author.

I take pleasure in naming this species for my wife, Wanda, who participated

in the first collection of this species.

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Vol. 50, No. 2, April 1974

161

This species is similar to Cyclocephala arenosa Howden and Endrodi

in the form of the apex of clypeus and general body shape and color,

although smaller. C. wandae keys to C. puberula LeConte or C. crepuscu-

laris Martinez in Endrodi’s key (1966:63) (however, not fitting either

couplet), but may be readily distinguished by the male genitalia (fig.

1 ) . The male genitalia closely resembles that of C. seditiosa LeConte,

however seditiosa has a 10-segmented antennae vs. 9-segmented for

wandae , and is larger (10-12 mm vs. 6. 6-7. 5 mm).

This species exhibits a number of characteristics which appear to be

adaptations to the Sand Dune habitat. The well developed brush of

hairs on the mentum is present in Leptohoplia (Howden and Hardy,

1971 ) , and an undescribed species in Anomalini. The light color is

present in Pseudocotalpa andrewsi Hardy; an undescribed species of

Anomala ; and Cyclocephala arenose H. & E., in addition to the above.

Literature Cited

Endrodi, Sebo. 1966. Monographic der Dynastinae. Teil I. [Cyclocephalini] .

Entomol. Abhandl., 33: 1-460.

Howden, H. F. and Endrodi, S. 1966. Five new species of Cyclocephala

Latreille from North and Central America (Coleoptera: Scarabaeidae) .

Can. Entomol., 98(1) : 295-302.

Howden, H. F. and Hardy, A. R. 1971. Generic placement and adult behavior

of the genus Leptohoplia Saylor. Proc. Entomol. Soc. Washington,

73(3): 337-341.

162

The Pan-Pacific Entomologist

A Redefinition of the Tyrini with the Addition of

Anitra Casey

(Coleoptera: Pselaphidae)

Donald S. Chandler

Ohio State University, Columbus, 43210

A small series of a pselaphid was recently collected at Molino Basin

in the Santa Catalina Mountains of Arizona by Karl Stephan on March

7, 1970. The beetles were found under a large, flat rock and were as-

sociated with a colony of ants warming brood. The ants were determined

as Paratrechina melanderi (Wheeler).

The pselaphids were identified as Anitra glaberula Casey upon com-

parison with the male holotype (USNM #38748). Anitra was initially

placed in the Ctenistini by Casey (1893) and has been retained there by

succeeding workers (Raffray 1908, Park 1953). Detailed examination

of the specimens convinced me that the genus was misplaced and should

be removed to the Tyrini.

Casey placed the genus in the Ctenistini when the differences between

that tribe and the Tyrini were not very obvious to him (p. 499). His

detailed description gives no indication as to why the genus was placed

in the Ctenistini rather than the Tyrini. Raffray (p. 331) questioned

Casey’s placement and suggested that the genus might best be placed

in the Hybocephalini. However, Park (p. 316) still retained Anitra in

the Ctenistini long after Raffray ’s suggestion.

The Ctenistini are placed in the same subfamily, the Pselaphinae, and

the same division, the Macroscelia, as the Tyrini, but the two tribes can

be separated easily by two characters. The Ctenistini are characterized

by the presence of squamiform setae, particularly in the foveae and sulci

of head, thorax and the articular surfaces of the abdominal segments.

Also, the epistome is large, expanded and usually prominent laterally.

The Tyrini are characterized by moniliform setae and the epistome may

he large, but is neither expanded nor prominent. The Hybocephalini,

which was referred to by Raffray, also possess squamiform setae.

Anitra possesses no squamiform setae. On the basal prontal projec-

tion there are some short, thick setae which were clearly not squamiform

when a specimen mounted on a slide was examined. A similar grouping

of short, thick setae occurs also in Juxtahamotopsis Park of the Hamotini

(Park 1945), a tribe very closely related to the Tyrini. The epistome is

smoothly fused to the frons and is neither expanded nor prominent. On

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Vol. 50, No. 2, April 1974

163

Fig. 1. Right maxillary palpus.

Fig. 2. Right mandible.

the basis of these characters, it is apparent that Anitra does not be-

long in the Ctenistini or the Hybocephalini and should be placed in the

Tyrini.

Anitra does not fit within the Tyrini as it is currently defined, but

the differences can be explained as reduction of characters due to the

myrmecophilous habits of the genus. The Tyrini as currently defined

have four-segmented maxillary palpi and one to three basal pronotal

foveae. Anitra has two-segmented maxillary palpi (Fig. 1) (Casey

said three segments) and has no basal pronotal foveae. The genus is

certainly myrmecophilous because of the circumstances of its collection

and probably functions as a synoekete. Reduction of the number of

palpal segments is common to myrmecophilous genera, for example the

two-segmented maxillary palpi of Biotus Casey (Ctenistini) and the

one- or two-segmented palpi of the Clavagerinae. The reduced teeth on

the mandibles (Fig. 2) are a characteristic common to myrmecophilous

beetles. The lack of basal pronotal foveae is not too surprising in a tribe

where the number varies from one to three

Because of the absence of squamiform setae and the lack of a promi-

nent epistome, I conclude that Anitra should be placed in the Tyrini

rather than the Ctenistini or Hybocephalini. I am also expanding the

limits of the Tyrini so that the number of the segments of the maxillary

palpus vary from two to four and the number of basal pronotal foveae

vary from zero to three.

I would like to thank Dr. Charles Gaspar of the Institut Agronomique

de Tetat Gembloux, Belgium, for the identification of the ants, Mr.

T. J. Spilman of the United States National Museum for permission

to examine the Casey collection, and Dr. C. A. Triplehorn of the Ohio

State University for checking the manuscript.

164

The Pan-Pacific Entomologist

Literature Cited

Casey, T. L. 1893. Coleopterological notices, V. Ann. New York Acad. Sci.,

7: 281-606.

Park, 0. 1942. A study in neotropical Pselaphidae. Northwestern Univ. Studies

Bio. Sci. Med., no. 1, 403 pp.

Park, 0. 1953. Discrimination of genera of pselaphid beetles of the United

States. Bull. Chicago Acad. Sci., 9: 299-331.

Raffray, A. 1908. Pselaphidae. Genera insectorum, 64th fascicule, P. Wytsman,

ed. Bruxelles. 487 pp.

RECENT LITERATURE

The Insects of Virginia: No. 6. Horseflies and deer flies of Virginia

(Diptera: Tabanidae). L. L. Pechuman. 92pp. (Research Division Bulletin

81, Virginia Polytechnic Institute and State University), 1973. Available on an

exchange basis from the Research Division Mailing Room, Hutcheson Hall,

Virginia Polytechnic Institute and State University, Blacksburg, Virginia

24061.

This faunal treatment ostensibly covers only a small geographic area, but will

probably be useful as a guide to the horseflies of much of the central Atlantic

coast. Included are short sections on the biology and ecology of tabanids, their

taxonomic characters, distribution, and economic significance, as well as lengthier

taxonomic treatment of the 95 species known to occur in Virginia. Keys are pro-

vided for separating the genera and the species within each genus (separate keys

for males and females). For each species a concise summary is given of the

distribution in Virginia, and the seasonal occurrence, along with biological notes. —

Editor.

Vol. 50, No. 2, April 1974

165

The Genus Dicopus Enock

(Hymenoptera: Mymaridae)

Richard L. Doutt

University of California, Berkeley

A century has passed since Enock in England became interested in the

Mymaridae. He quaintly described his initial fascination with these

parasitic wasps as follows (1909) : “ in 1872 I commenced my

own observations from a simple exhibit by Mr. Frederick Fitch at the

Quekett Microscopical Club. Under the microscope was shown ‘A Fairy

Fly in a Spider’s Web.’ It was illuminated on a black background,

which gave it the most wonderful appearance, every limb, each tiny

hair and the long cilia resembling brilliant silver. It was a most fasci-

nating object to any one, and speaking for myself, I had never before

seen anything to compare with its delicate fairy-like structure. I then

and there determined to know more about its relatives. Next day I

searched the spiders’ webs in my garden at Holloway and found quite

a number of several species, which I mounted in Canada balsam.”

“On removal in 1882 to Woking I very soon found I was in the land

of plenty for Mymaridae, which simply swarmed on the windows of the

house and in a very small conservatory — where I caught seven of the

new genera which I am introducing for the first time tonight.” One

of the new genera in the paper which Enock read to the Entomological

Society of London was Dicopus. The type species is Dicopus minutis-

sirnus Enock which he described from a female, the male being unknown.

In 1911 A. A. Girault received a single male mymarid from Quebec

“so minute as almost to be lost in the medium in which it was mounted.”

At first glance he thought it was an Alaptus but quickly recognized that

it belonged to another genus and he made a very astute analysis (1911) :

“There is an English genus, however, recently described by Enock for

Dicopus minutissima Enock, which closely resembles Alaptus Haliday

in form, but which differs in bearing two more antennal segments; the

male of this genus is unknown. Now, this Canadian species precluded

from being an Alaptus must belong to Dicopus, at least until we know

to the contrary. It is one of the smallest North American Mymaridae

and because of this and also because of its characteristic appearance,

I believe it is incumbent on me to describe it rather than risk its being

lost.” This single male individual, the type of Dicopus halitus Girault,

was the sole record for the genus on the North American continent until

the specimens described herein were collected in California.

The Pan-Pacific Entomologist 50 : 165 - 168 . April 1974

166

The Pan-Pacific Entomologist

In September 1911, while en route to Australia, Girault stopped for

a few hours at Suva, Fiji and he “managed to collect a few parasitic

Hymenoptera.” One of these was captured on a window pane in a

woodworker’s shop. Girault recognized it as a male Dicopus and named

it D. psyche. “This Fijian species, like the American form, is very

minute, also extremely delicate, and was extraordinarily difficult to

capture; it was moving slowly over the pane of a window, but I was

not able to keep sight of it for more than the fraction of a second at a

time. This fact, taken in conjunction with its fragility, made it necessary

to spend three-quarters of an hour in effecting its capture.” (Girault,

1912).

In 1912 Ricardo Mercet described Dicopus citri from Spain, and

Girault later added 3 Australian species by describing D. victoria and

D. maximus in 1920, and D. hidentiscapus in 1931. Also in 1931

Claude Morley in England described D. cervus from a male hymenop-

teran which “ flew on to the book I happened to be reading at

5 p.m. on July 17th, 1921, while sitting upon the lawn beside the moat

which surrounds Monks’ Soham House in Suffolk.”

This extraordinary detailed collection datum for a mymarid was

rendered ineffectual in 1955 when Alejandro Ogloblin indicated that

D. cervus Morley is not a mymarid. In the same paper Ogloblin pro-

posed four new genera from Argentina all closely related to Dicopus.

These are Chromo dicopus, Callodicopus, Dicopulus and Dicopomorpha.

Ogloblin suggested that some of the species referred to Dicopus by

Mercet and Girault might possibly be transferred to other genera. From

my examination of Girault ’s types at the Queensland Museum I believe

that D. victoria Girault and D. maximus Girault are incorrectly placed

in Dicopus and instead probably belong to one of Ogloblin’s genera

or to a closely related but undescribed genus.

The genus Dicopus is therefore presently constituted of D. minutis-

simus Enock, D. halitus Girault, D. psyche Girault, D. citri Mercet,

D. hidentiscapus Girault and the California species described as follows:

Dicopus pygmaeus, new species

Female. — Body length 0.22 mm; forewing length 0.44 mm; wing blade 0.30 mm,

apical marginal cilia 0.14 mm; head, thorax, abdomen brown, appendages pale

testaceous, eyes and ocelli deep maroon. Toruli high on face near frontal carina

of vertex; mandibles elongate. Vertex finely granulate, lateral carinae well de-

veloped, thick, interrupted, in 3 sections. Face with several prominent bristles.

Antennae 1.5 X body length. Funicle segments 6, 7 distinctly wider than preced-

ing funicle segments, club elongate with 2 terminal projections (Fig. 2) . Abdomen

sessile, mesophragma penetrates % its length, ovipositor short, not exserted. Legs

Vol. 50, No. 2, April 1974

167

Figs. 1-3. Antennae. 1. Dicopus pygmaeus male. 2. D. pygmaeus female.

3. D. enocki female.

longer than body, fore femora and tibiae slightly swollen, fore coxae largest.

Foretarsus with basal segment barely exceeding segment 2, apical segment (5)

longest.

Male. — Similar to female in size and color. Antenna (Fig. 1) 12 segmented,

funicle segments 6, 7, 8, 9 largest, widest ; club with terminal spine. Hind

trochanters elongate.

Holotype female, Mecca, Imperial County, California, on sugar beet, 19

June 1964, H. R. Moffitt. Allotype, same data as holotype except collected 21

May 1964. Types at Division of Biological Control, University of California,

Albany.

The male antenna of D. pygmaeus is similar to D. halitus , but the

terminal spine of the club and the elongate hind trochanters are dis-

tinctive. There are also color differences as D. halitus is sooty black.

The female of D. halitus is unknown.

Dicopus enocki, new species

Female. — Body length 0.35 mm; forewing length 0.73 mm; wing blade 0.46

mm, apical marginal cilia 0.27 mm. Head, thorax, abdomen brown; legs and an-

tennae light brown; eyes red. Scutum, postscutellum lightly reticulate; scutellum,

propodeum smooth. Head large, as wide as thorax. Vertex with gross carinae.

Mandibles elongate, single toothed. Toruli high on face, near frontal carina of

vertex. Scape long with two distinct tooth-like projections bearing spines on

ventral aspect (Fig. 3). Funicle segments 1-6 slender, elongate; segment 7

shortest, widest. Club elongate with 2 terminal projections. Legs slender; fore

coxae largest; foretarsus with basal segment distinctly longer than segment 2.

Mesophragma penetrates Vs abdomen; ovipositor short, not exserted.

Male. — Unknown.

Holotype female, Kingsburg, Fresno County, California, suction trap in

vineyard, 23-24 April 1965, J. Nakata; 2 paratype females same data as holotype

except collected 15-16 May 1965 and 25-26 April 1965; 1 2 paratype; Caswell

State Park, Stanislaus County, California, on Rubus, 29 April 1963, R. L. Doutt.

Type series at Division of Biological Control, University of California, Albany.

168

The Pan-Pacific Entomologist

D. psyche Girault is similar to D. enocki but differs by the lack of

tooth-like projections on the scape and a wider funicle segment 6. D.

pygjnaeus differs in its antennal structure (compare Figs. 2 and 3),

in its shorter basal segment of the foretarsus, and in the larger develop-

ment of the mesophragma. The name D. enocki is in centennial recogni-

tion of the contributions of Enock to the study of Mymaridae.

Literature Cited

Enock, F. 1909. XI New genera of British Mymaridae. Trans. Entomol. Soc.

London, 1909, pt IV, 449-459.

Girault, A. A. 1911. The probable occurrence of the Mymarid genus Dicopus

Enock in North America (Hymen.). Entomol. News, 22: 347-349.

Girault, A. A. 1912. On the probable occurrence of the Mymarid genus Dicopus

Enock in Fiji. Proc. Entomol. Soc. Wash., 14: 22-23.

Girault, A. A. 1920. New genera and species of Australian Mymaridae. Insec.

Inscit. Menstr., 8: 96-100.

Girault, A. A. 1931. A new habit in an old insect. Homo pudicus and new

Eurytomidae. Privately published, Brisbane, 4 pp.

Mercet, Ricardo. 1912. Mimaridos nuevos de Espaha. Boll. R. Soc. Esp.

Hist. Nat., Madrid, 12: 331-337.

Morley, Claude. 1931. New Oxyura from Britain. The Entomologist, 64(812) :

14-16.

Ogloblin, Alejandro. 1955. Los nuevos representantes de la famn. Mymaridae

(Hym.) de la Republica Argentina. Boll. Lab. Zool. Gen. Agr. Filippo

Silvestri, Portici, 33: 377-397.

RECENT LITERATURE

Cerambycidae of North America. Part VI, No. 1. Taxonomy and classifi-

cation of the subfamily Lepturinae. E. G. Linsley and J. A. Chemsak, Uni-

versity of California Publications in Entomology, 69: 1-138, 1973.

The excellence of the first five volumes is continued in this work which treats

the tribes Desmocerini and Necydalini, along with a portion of the Lepturini. As

in the preceding parts, the text is well illustrated with habitus drawings and

distributional maps. — Editor.

Vol. 50, No. 2, April 1974

169

A review of the genus Mellinus

(Hymenoptera: Sphecidae)

M. L. Siri and R. M. Bohart

Department of Entomology, University of California, Davis

Mellinus Fabricius is a small genus occurring in the Palearctic Re-

gion and in North America as far south as Guatemala. These wasps

form an apparently relict group best placed in the Nyssoninae. Gen-

eralized features are the short tongue, evident notauli, simple pro-

podeum, no omaulus, two midtibial spurs, recurrent veins widely sepa-

rated at their forward ends, and a long triangular marginal cell. The

principal specialization is the petiolate gaster which results in a body

form somewhat like that of the philanthine genus Trachypus.

Morphological details useful at the specific level are few except

for size and color. Helpful are clypeal shape, male antennal form,

petiole shape, tibial setation, and male genitalia.

Studies of male genitalia have revealed dramatic differences among

several species (figs. 13-16) and strong similarities among others. In

abdominalis, a peculiarity is a dorsal arm of the digitus which opposes

a fingerlike extension from the enlarged gonostyle (fig. 15). The three

species of the rufinodus group, rufinodus , bimaculatus and imperialis ,

have nearly identical genitalia. In these, the two branches of the

aedeagus are loosely connected by membrane which seems capable of

great expansion (fig. 16). Also unusual in the rufinodus group are

the overlapping distal lobes of the gonostyle.

In palearctic species, ground nests are stocked with flies, especially

Muscoidea. Since arvensis is rather common, it has been observed

frequently in the act of pouncing like a cat on flies attracted to fresh

cow droppings. Huber (1961) has presented a detailed exposition of

known biology.

We are treating Mellinus as including 10 species of which three are

palearctic. One of the seven New World forms is described as new.

Several of the species are rare in collections. We have seen no ma-

terial of obscurus , and females only of alpestris and satanicus. Since

markings are diagnostic for known species, males of the last two should

be readily recognized when found.

Material has been borrowed from several institutions, but we are

especially grateful to curators at the U.S. National Museum, Cornell

University, American Museum of Natural History, California Academy

of Sciences, and Canadian National Collection.

The Pan-Pacific Entomologist 50: 169-176. April 1974

170

The Pan-Pacific Entomologist

Key to species of Mellinus

1. Propodeum generally rugose posteriorly, enclosure medially ridged and

sculptured but with a smooth outer area (fig. 4), palearctic 2

Propodeum weakly sculptured posteriorly except toward abdominal in-

sertion, enclosure evenly shagreened (fig. 5), New World 4

2. Tergal markings yellow arvensis (L.)

Tergal markings whitish, female legs reddish 3

3. Tergum V banded, scutellum with a whitish spot crabronea Thunb.

Tergum V all dark, scutellum all dark obscurus Handl.

4. Abdomen all dark _ 5

Abdomen with pale markings 6

5. Female clypeus without a median discal projection, scutellum lightly sha-

greened and somewhat polished between punctures, pronotal ridge yellow, fe-

male flagellomere I about 2.5 times as long as broad (fig. 8) _ pygmaeus Handl.

Female clypeus with median discal projection (figs. 3, 6), scutellum closely

shagreened and dull between punctures, pronotum all dark, female flagel-

lomere I about 2.8 times as long as broad (fig. 7) satanicus Siri and Bohart

6. Tergum III with a complete pale band or with closely approximated

pale spots 7

Tergum III at most with widely separated pale spots 8

7. Petiole and legs largely red, both scutellum and metanotum with pale spots,

terminal flagellomeres about half yellow and half black, tergum IV usually

with pale spots rufinodus Cr.

Petiole and legs largely brown, both scutellum and metanotum dark, ter-

minal flagellomeres dark, tergum IV banded imperialis Bohart

8. One or more abdominal segments beyond petiole red, male flagellomeres

XXI distinctly enlarged and black (fig. 9), median lobe of male clypeus

protruding and truncate (fig. 1) abdominalis Cr.

No abdominal segments beyond petiole red, no asymmetry or enlarge-

ment of male flagellomeres, male clypeus flat and with a tridentate apex

or at least one distinct median tooth 9

9. Flagellomeres dark, femora and tibiae black, tarsi brown with some yel-

low, petiole black with yellow markings alpestris Cam.

Flagellomeres dark dorsally but yellow ventrally, femora and tibiae light

red or yellow, petiole dark red to light red bimaculatus Pack.

Mellinus abdominalis Cresson

(Figs. 1, 9, 15)

Mellinus abdominalis Cresson, 1882. Trans. Amer. Entomol. Soc., Proc. 9:xxxix.

Lectotype $, Montana (Acad. Nat. Sci. Philadelphia).

Mellinus abdominalis personatus W. Fox, 1894. Entomol. News 5:202. Holotype

2 , Montana (Acad. Nat. Sci. Philadelphia) .

This is the most distinctive species of Mellinus. It is the only one

in which the abdomen is mostly or all red with pale markings reduced

Vol. 50, No. 2, April 1974

171

Figs. 1-12. Fig. 1. Mellinus abdominalis, male face. Fig. 2. M. rufinodus

female face, showing color pattern. Fig. 3. M. satanicus , female face. Figs. 4-5.

Propodeum and enlarged detail. Fig. 4, arvensis. Fig. 5, satanicus. Fig. 6. M.

satanicus head, lateral. Figs. 7 -8. Pedicel and flagellomeres I-II. Fig. 7, satanicus

female. Fig. 8, pygmaeus female. Figs. 9-11, crabronea. Fig. 12, M. satanicus,

petiole and base of abdominal segment II.

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or absent. The male is especially odd with respect to the slender and

gradually expanded abdomen, sculptured clypeal form (fig. 1), special-

ized antenna (fig. 7), prominent hair tuft on last two visible sterna,

and complex genitalia (fig. 15).

We have seen 19 specimens all from localities west of the 100th meridian as

follows: Montana: Platte Co.; Wyoming: Carbon Co.; Nebraska: Valentine,

Thedford, Alliance, Halsey: Colorado: Mesa Verde, Great Sand Dunes.

Mellinus alpestris Cameron

Mellinus alpestris Cameron, 1890. Biol. Centr.-Amer. Hymen. 2:85. Holotype

2, Xucumanatlan, Guerrero, Mexico (Brit. Mus. Nat. Hist. London).

We have seen females only, 11 specimens in all, including the holo-

type. Morphologically, alpestris and bimaculatus are quite similar

and must be separated by color characters as given in the key. All of

our specimens have a pair of yellow spots on the propodeal enclosure,

contrary to the case in bimaculatus. Also, our alpestris have the clypeus

all yellow but this character is known to be variable in other species.

Collections are from Mexican States as follows: Michoacan: El Pueblito;

Guerrero: Xucumanatlan; Mexico: Texcoco; Morelos: Cuernavaca, Amecameca.

Mellinus arvensis (Linnaeus)

(Figs. 4, 13)

V espa arvensis Linnaeus, 1758. Syst. Nat. Ed. 10:573. Holotype 2, “Europa,”

Linnaean Coll., London.

The complicated synonymy is not given here; see Bohart and Menke (in press).

We have studied 52 specimens of arvensis. In determining species

limits, reliance has been placed on the discussions given by Handlirsch

(1888) and Beaumont (1964). This is the largest species of Mellinus ,

surpassing rufinodus on the average. Some females attain a length

of 15 mm. The propodeal sculpture is characteristic of European

Mellinus as shown in fig. 4. Genitalia are depicted in fig. 15. The

species is widespread in Europe and is sometimes abundant in moun-

tainous areas. Specimens from the Alps tend to have the legs some-

what reddish, especially in females.

Mellinus bimaculatus Packard

Mellinus bimaculatus Packard, 1867. Proc. Entomol. Soc. Philad. 6:419. Syntype

2, Dublin, New Hampshire (Mus. Comp. Zool. Harvard); syntype 2, Bruns-

wick, Maine (location of type unknown).

Mellinus ivolcotti H. S. Smith, 1908. Entomol. News 19:299. Holotype 2, Beaver

Island, Michigan (Univ. Nebraska) .

Vol. 50, No. 2, April 1974

173

Figs. 13-16. Male genitalia, ventral, and enlargement from aedeagal area.

Fig. 13, arvensis. Fig 14, pygmaeus. Fig. 15, abdominalis. Fig. 16, imperialis.

We have seen 18 specimens. The face may be nearly all black as

in Packard’s syntypes, variously mottled, or mostly yellow as in Smith’s

wolcotti. The species is similar to alpestris but differs in markings as

indicated in the key. The petiole is ordinarily black but occasionally

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may be dull red and rarely is yellow blotched. M. bimaculatus

average a little smaller than rufinodus, 8-9 mm long rather than 9-12

mm. The male genitalia are similar to those of rufinodus but the distal

lobes of the gonostyle are somewhat more slender and tapering.

This is the only species known to occur in eastern United States. However,

it ranges into Mexico. Localities are: Maine: Brunswick; New Hampshire:

Dublin, Franconia; Massachusetts; New York: Huntington, New Jersey; Ohio:

Stark Co.; Michigan: Clare Co., Midland Co., Beaver Island, Douglas Lake.

In Mexico it has been collected in Jalisco: La Primavera; and Oaxaca: Macuilxo-

chitl.

Mellinus crabronea (Thunberg)

(Fig. 11)

Crabo sabulosus Fabricius, 1787, nec Sphex sabulosus Linnaeus, 1758, now in

Ammophila. Lectotype 2, Copenhagen, Denmark (Univ. Copenhagen Zool.

Mus.) .

Sphex crabronea Thunberg, 1791, new name for Crabo sabulosus Fabricius (Art.

59c, ICZN) .

We have seen only 10 specimens of crabronea and key characters

have been based on discussions (under sabulosus ) by Handlirsch

(1888) and Beaumont (1964). The subserrate form of the male an-

tenna is shown in fig. 11. The serrations seem to be a little more pro-

nounced than in arvensis but it must be said that the two species are

structurally almost identical. Separation on the basis of the white

markings in crabronea is not entirely satisfactory, particularly in males.

Recorded localities are in Europe and central Asia.

Mellinus imperalis R. Bohart

(Fig. 16)

Mellinus imperialis Bohart, 1968. Pan-Pac. Entomol. 44:235. Holotype £, Bard,

California (Univ. California, Davis).

The brown and extensively deep yellow color pattern distinguishes

this species from rufinodus. However, in the absence of structural dif-

ferences there remains the possibility that it is an early season (Feb-

ruary to April) Sonoran desert subspecies of rufinodus (June to Sep-

tember). We have seen a total of 11 specimens: California: Bard;

Sonora (Mexico) : Guaymas, Cocorit, Alamos.

Mellinus obscurus Handlirsch

Mellinus obscurus Handlirsch, 1888. Sitz. Akad. Wiss. Wien 96:288. Holotype

$, Korea (Radoszkowsky Coll., Warsaw).

Mellinus tristis Perez, 1905. Bull. Mus. Paris 1905:156. Holotype 2, Japan,

“Japan central” (Nat. Hist. Mus. Paris).

Vol. 50, No. 2, April 1974

175

Since we have seen no specimens of obscurus , reliance has been

placed on the original descriptions referred to above. The possibility

exists that obscurus is a dark variety or subspecies of crabronea from

eastern Asia.

Mellinus rufinodus Cresson

(Fig. 2)

Mellinus rufinodus Cresson, 1865. Proc. Entomol. Soc. Philad. 4:475. Leetotype

$, Colorado (Acad. Nat. Sci., Philadelphia).

The mostly red legs and petiole are a fairly constant color character

but the femora may be all dark. As in other Mellinus , facial markings

are variable. Some rufinodus have the face mostly yellow (fig. 2),

others mostly black. We have seen two specimens with a yellow band on

tergum IV as in imperialism but the more extensively pale underside of

the flagellum in the former species is distinctive.

We have seen 83 specimens. All United States material has been from west

of the 100th meridian. Nebraska: Thedford; Texas: Valentine; Colorado: Den-

ver, Dumont, Poudre Canyon; New Mexico: Pecos, Organ Mts., Raton; “Mon-

tana”; Utah: Box Elder Co., Farmington, Provo; Arizona: Chiricahua Mts.,

Huachuca Mts., Santa Rita Mts. Mexican localities by States are: Chihuahua:

Arroyo Mestino, 7600 ft.; Durango: 10 mi. w. Durango; Guerrero: 40 mi. n.

Queretaro; Mexico: Teotihuacan; Aguascalientes: Penvelos.

Mellinus pygmaeus Cameron

(Figs. 8, 10, 14)

Mellinus pygmaeus Handlirsch, 1888. Sitz. Akad. Wiss. Wien 96:289. Syntypes,

1 $, 2 2, Orizaba, Mexico (Nat. Hist. Mus. Vienna).

This small dark species is similar to satanicus but smaller, about

7 mm long instead of 12.5, and more polished, especially on terga I-II.

The female antenna is perceptibly more clubbed in pygmaeus and a

pair of yellow spots are nearly always present on the pronotal ridge.

The clypeus is flat and may be black or with considerable yellow.

Genitalia (fig. 14) are distinctive among known forms. The gonostyle

is relatively simple and the aedeagus bears stout spines.

We have seen 20 specimens, one from Panajachel, Guatemala and the others

from Mexico: Michoacan: Puerta Garnica; Veracruz: Coscomatepec, Orizaba

(holotype) ; Mexico: Texcoco; Puebla; Cuetzalan; Morelos: Amecameca.

Cuernavaca; Chiapas: San Cristobal. Handlirsch (1888) reported a pair of

pygmaeus in the Dresden Museum (now destroyed) from “Brasilien.” The oc-

currence of the species in South America needs confirmation.

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Mellinus satanicus, new species

(Figs. 3, 5, 6, 7, 12)

Female: Length 12.5 mm. Black with yellow in narrow spots along inner orbit,

scape in front, faint lateral dots on clypeus, and most of foretibia anteriorly;

wings lightly stained. Pubescence pale fulvous, moderate, short but erect and

thick beneath thorax as well as under trochanters and femora. Body closely

microsculptured in a reticulate pattern, scutum posteriorly with short and fine

longitudinal ridging, mesopleuron and scutellum with minute punctures scattered

over microsculpture. Face (figs. 3, 6) with a sharp, compressed, hornlike, median,

projection at lower one third of disc; antenna slender, flagellomere I about 2.8

times as long as broad (fig. 7) ; propodeum finely sculptured (fig. 5) ; petiole

4.0 times longer than broad, one third as broad as tergum II (fig. 12). Outer

margin of hindtibia with numerous relatively long spines set on low protuberances.

Holotype female, Zontehuitz near San Cristobal, Chiapas, Mexico, 9600

ft., June 25, 1969 (W. R. M. Mason, Canadian National Collection, Ottawa).

The peculiar horned clypeus (figs. 3, 6) separates the species im-

mediately. In addition, the extreme reduction of yellow is remarkable.

This should lead to recognition of the male when it is discovered.

References Cited

Beaumont, J. de. 1964. Insecta Helvetica fauna 3. Hymenoptera: Sphecidae.

169 pp. Soc. Entomol. Suisse, Lausanne.

Bohart, R. M. and A. S. Menke (in press) . Sphecid wasps, a generic revision,

U. C. Press, Berkeley.

Dalla Torre, C. G. de. 1897. Catalogus Hymenopterorum, etc., vol. 8. Fossores.

viii + 749 pp. G. Engelmann, Lipsiae.

Fox, W. J. 1894. Studies among the fossorial Hymenoptera. III. Synopsis of

the N. Am. species of the genus Mellinus Fab. Entomol. News, 5: 201-

203.

Handlirsch, A. 1888. Monographie der mit Nysson und Bembex verwandten

Grabwespen. Sitz. Akad. Wiss. Wien. Math. -Nat. Classe, 96: 219-311.

Huber, A. 1961. Zur biologie von Mellinus arvensis. Zool. Jahrb. Abt. Syst.

Oekol. Geol. Tiere, 89: 43-118.

Vol. 50, No. 2, April 1974

177

Two New Heliococcus Species,

a Key to the North American Species,

and a List of World Species.

(Homoptera: Coccoidea: Pseudococcidae)

Douglass R. Miller

Systematic Entomology Laboratory, IIBIII, USDA, ARS,

Beltsville, Maryland 20705

In this paper two new species of Heliococcus are described for in-

clusion in a study on the mealybugs of San Clemente Island. Because

no comprehensive work has been published on Heliococcus, a diagnosis

of generic characters and a list of known species are given.

Heliococcus Sulc, 1912.

Type species.— Heliococcus bohemicus Sulc, 1912, by monotypy.

Diagnosis. — Adult female: anal lobes large, strongly protruding, normally

sclerotized; ventral-lobe sclerotization normally elongate and conspicuous; 2-18,

normally 17 or 18, pairs of cerarii; 2 or 3 conical setae in each cerarius; multi-

locular pores, when present, ventral only (except on H. phaseoli) , normally re-

stricted to areas near vulva, sometimes absent; quinquelocular pores restricted

to venter (except on H. dorsiporosus) , confined to medial and mediolateral areas;

trilocular pores present in all areas except medial and mediolateral areas of

venter; small discoidal pores sparce on both surfaces; crateriform tubular ducts

present; oral-collar tubular ducts normally present, restricted to venter; anal ring

with 3 pairs of setae and 3-5 rows of pores; dorsal and ventrolateral setae normally

conical; circulus normally present; hind legs with or without translucent pores;

tarsal digitules not apically capitate; claws each with large denticle; claw digitules

capitate; antennae normally 9-segmented; with 2 pairs of ostioles.

Notes. — Heliococcus is most closely related to Saliococcus Kanda and

T akahashicoccus Kanda in that all three genera possess the characteristic

crateriform tubular ducts. Saliococcus and T akahashicoccus differ from

Heliococcus in possessing cerarii which each contain more than 3

conical setae. In Heliococcus all but the ocular cerarii contain less than

4 conical setae.

List of World Species of Heliococcus

Except for one questionably placed Ethiopian species, H. phaseoli,

Heliococcus is exclusively Holarctic. The following list has been com-

piled from the literature and includes 39 species and subspecies, eight

from the New World and 31 from the Old World.

The Pan-Pacific Entomologist 50: 177-192. April 1974

178

The Pan-Pacific Entomologist

1. adenostomae McKenzie, 1960: 707 (California)

2. artemisiae Ter-Gigorian, 1967: 134 (Armenian SSR)

3. atraphaxidis Bazarov, 1963: 38 (Tadzhik SSR)

4. atriplicis McKenzie, 1964: 235 (California)

5. bambusae (Takahashi) (Taiwan)

Phenacoccus bambusae Takahashi, 1930: 6

Heliococcus bambusae (Takahashi), Goux, 1934: 171

6. bohemicus Sulc, 1912: 39 (“Bohemia”; France; Germany; USSR)

7. caucasicus Borchsenius, 1949: 279 (Armenian SSR)

8. cinereus Goux, 1934: 164 (France)

9. clemente Miller, new species (California)

10. cydoniae Borchsenius, 1949: 272 (USSR)

11. deserticola Miller, new species (Mexico; Arizona, California, Nevada, New

Mexico, Texas)

12. destructor Borchsenius, 1941: 6 (Kirgiz SSR, Kazakh SSR, Tadzhik SSR,

Turkmen SSR, Uzbek SSR)

13. dorsiporosus Danzig, 1971: 382 (Eastern USSR)

(It is questionable that this species belongs in Heliococcus because it has

dorsal quinquelocular pores.)

14. glycinicola Borchsenius, 1956: 678 (Korea)

15. halocnemi Borchsenius, 1949: 277 (Uzbek SSR)

16. herbaceus Borchsenius, 1956: 678 (Korea)

17. insignis (Lobdell) (Kansas, Louisiana, Mississippi)

Phenacoccus insignis Lobdell, 1930: 210

Heliococcus insignis (Lobdell), Ferris, 1950: 97

18. kehejanae Ter-Grigorian, 1967: 136 (Armenian SSR)

19. kurilensis Danzig, 1971: 386 (Eastern USSR)

20. marginalis Goux, 1953: 104 (France)

21. maritimus Danzig, 1971: 388 (Eastern USSR)

22. minutus (Green) (Channel Island of Guernsey, England)

Phenacoccus minutus Green, 1925: 519

Heliococcus minutus (Green), Williams, 1962: 29

23. montanus Borchsenius, 1949: 274 (Tadzhik SSR, Uzbek SSR)

24. nivearum Balachowsky, 1953: 238 (France; USSR)

25. nivearum austriacus Balachowsky, 1953 : 240 (Austria)

26. oligadenatus Danzig, 1972: 333 (Mongolian Republic)

27. osborni (Sanders) (Canada; Colorado (?), Indiana, Iowa, Louisiana,

Missouri, New York, Ohio, Virginia; Egypt (?) )

Phenacoccus ( Paroudablis ) osborni Sanders, 1902 : 284

Phenacoccus pettiti Hollinger, 1917 : 281

Heliococcus osborni (Sanders), Ferris, 1950: 99

28. pavlovskii Borchsenius and Tereznikova, 1959: 491 (Maritime Territory,

USSR)

29. phaseoli (Laing) (Sierra Leone)

Phenacoccus phaseoli Laing, 1929: 475

Heliococcus phaseoli (Laing), Goux, 1934: 171

(Based on the large number of dorsal multilocular pores, it is possible that

phaseoli does not belong in Heliococcus.)

30. radicicola Goux, 1931: 113 (France; Germany; Poland; Sweden; Cwmea)

Vol. 50, No. 2, April 1974

179

31. salviae Borchsenius, 1949: 282 (Tadzhik SSR)

32. saxatilis Borchsenius, 1949: 276 (Armenian SSR)

33. Slavonicus Borchsenius and Tereznikova, 1959: 492 (Ukrainian SSR)

34. stachyos (Ehrhorn) (California)

Phenacoccus stachyos Ehrhorn, 1900: 313

Heliococcus stachyos (Ehrhorn), Goux, 1934: 171

35. sulcii Goux, 1934: 167 (France; Germany; Ukrainian SSR)

36. szetshuanensis Borchsenius, 1962: 232 (China)

37. tes quorum Borchsenius, 1949: 284 (Kazakh SSR)

38. wheeleri (King) (Texas)

Dactylopius wheeleri King, 1902: 285

Pseudococcus wheeleri (King), MacGillivray, 1921: 133

Heliococcus wheeleri (King), Ferris, 1953: 363

(Described from immatures. The identity of this species is uncertain.

Ferris (1953) believed that it might be a senior synonym of insignis.)

39. zizyphi Borchsenius 1958: 161 (China)

List of Species Erroneously Placed in Heliococcus

1. hystrix (Baerensprung) = Phenacoccus hystrix (Baerensprung)

Coccus hystrix Baerensprung, 1849: 174

Phenacoccus hystrix (Baerensprung), Lindinger, 1912: 293

Heliococcus hystrix (Baerensprung), Thiem, 1930: 140

2. malvastrus McDaniel = Ferrisia virgata (Cockerell)

Heliococcus malvastrus McDaniel, 1962: 323; McKenzie, 1967: 181

3. multipori Takahashi

Heliococcus multipori Takahashi, 1951: 18

(This species does not belong in Heliococcus because it lacks crateriform

tubular ducts. It probably should be placed in Phenacoccus, but without

specimens I cannot be positive.)

4. sakai Takahashi

Heliococcus sakai Takahashi, 1951:16

(This species does not belong in Heliococcus because it lacks crateri-

form tubular ducts. It probably should be placed in Phenacoccus, but

without specimens I cannot be positive.)

5. takae (Kuwana) = Saliococcus takae (Kuwana)

Dactylopius takae Kuwana, 1907 : 184

Phenacoccus takae (Kuwana) , Kuwana, 1917 : 6

Saliococcus takae (Kuwana), Kanda, 1934: 309

Heliococcus takae (Kuwana), Kanda, 1935: 70

(In 1934 Kanda chose takae to be the type species of Saliococcus Kanda.

In 1935 he stated that Saliococcus was a junior, subjective synonym

of Heliococcus and placed takae in that genus. The description of Salio-

coccus tokyoensis (1959a) indicates that he changed his mind about

the status of Saliococcus. The presence of cerarii with more than 3

conical setae easily distinguishes Saliococcus from Hel.iococcus.)

6. takahashii Kanda — Takahashicoccus takahashii (Kanda)

Heliococcus takahashii Kanda, 1935: 73

Takahashicoccus takahashii (Kanda), Kanda 1959b: 239

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The Pan-Pacific Entomologist

7. ivilmattae (Cockerell) — Phenacoccus wilmattae Cockerell

Phenacoccus wilmattae Cockerell, 1901: 57

Heliococcus wilmattae (Cockerell), Goux, 1934:171

(This species was placed in Heliococcus because of a misidentification

of wilmattae by Bueker (1930).)

Key to Adult Females of North American

Species of Heliococcus 1

1. With 15 or more pairs of cerarii 3

With less than 10 pairs of cerarii 2

2(1). Large-sized crateriform tubular ducts surrounded by cluster of small-

sized ducts; multilocular disk pores absent clemente Miller, n. sp.

Large-sized crateriform tubular ducts without cluster of small-sized ducts;

multilocular disk pores present near vulva atriplicis McKenzie

3(1). Crateriform tubular ducts of at least 2 sizes 5

Crateriform tubular ducts generally of 1 size, occasionally with 1 or 2

smaller ducts in dorsomedial areas of abdomen 4

4(3). Venter with multilocular disk pores and oral-collar tubular ducts; with

at least 10 trilocular pores associated with each cerarius insignis (Lobdell)

Venter without multilocular disk pores and oral-collar tubular ducts;

with 5 or fewer trilocular pores associated with each cerarius

adenostomae McKenzie

5(3). Large-sized crateriform tubular ducts without associated smaller ducts;

multilocular disk pores normally present 6

Large-sized crateriform tubular ducts along body margin with as-

sociated clusters of smaller ducts; mutilocular disk pores ab-

sent deserticola Miller, n. sp.

6(5). Small-sized crateriform tubular ducts much more abundant than large-

sized ducts; normally with a cluster of oral-collar tubular ducts near

anterolateral margins of abdominal segment IX; oral-collar tubular ducts

normally present on venter of abdominal segments VIII-V or

IV osborni (Sanders)

Small-sized crateriform tubular ducts less abundant or equal in number

to large-sized ducts; without a cluster of oral-collar tubular ducts on

abdominal segment IX; oral-collar tubular ducts normally restricted

to abdominal segments IX or VIII, rarely on VII stachyos (Ehrhorn)

Heliococcus clemente, new species

(Fig. 1)

Suggested common name. — San Clemente mealybug.

Type material. — Holotype adult female, Pyramid Head, San Clemente Island,

Los Angeles County, California, 12 May 1973, collected under rock, D. R.

Miller and A. S. Menlte (USNM).

Adult female. — Holotype, mounted, 2.9 mm long, 1.8 mm wide. Body oval;

anal lobes strongly protruding and sclerotized, with 3 large-sized crateriform

ducts on each lobe.

1 H. ivheeleri is not included because adult females are not available.

Vol. 50, No. 2, April 1974 181

Fig. 1. Heliococcus clemente, adult female

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The Pan-Pacific Entomologist

Dorsum with 3 pairs of cerarii on abdomen, 1 pair on head. Anal-lobe cerarii

each with 2 elongate, conical setae, 2 or 3 trilocular pores, and large area of

basal sclerotization. Remaining abdominal cerarii with progressively smaller conical

setae, with or without trilocular pores, without basal sclerotization. Ocular cerarii

each with 1 large and 1 small conical seta, 1 or 2 associated trilocular pores, no

basal sclerotization. Multilocular disk pores and quinquelocular pores absent.

Trilocular pores scattered over surface. Discoidal pores present, few. Crateri-

form tubular ducts of 2 sizes: larger size scattered over surface, with 1-4 setae

at base of each sclerotized crater, surrounded by 13-35 (av. 25) small-sized ducts;

smaller size present in clusters around larger ducts and singly in medial, medio-

lateral, and rarely lateral areas; small-sized crateriform ducts without associated

setae. Body setae conical.

Anal ring bent over apex of abdomen, with 4 or 5 rows of pores; each of 6

anal-ring setae noticeably longer than greatest diameter of ring.

Venter with anal-lobe sclerotization elongate. Multilocular disk pores absent.

Quinquelocular pores abundant in medial and mediolateral areas, absent else-

where. Trilocular pores present in mediolateral and lateral areas. Discoidal pores

present in small numbers. Crateriform tubular ducts of same sizes as on dorsum,

clusters composed of 1 large- and many small-sized ducts, clusters restricted to

near body margins; small-sized ducts lightly scattered over surface, least abundant

on medial areas of abdomen. Oral-collar tubular ducts with conspicuous, sclero-

tized dermal opening, restricted to mediolateral areas of abdominal segments

VIII and VII. Body setae of 2 kinds: bristle-shaped setae present medially and

laterally; conical setae present mediolaterally and laterally.

Circulus present. Legs with small translucent pores on dorsal surface of hind

tibiae; hind tibia/tarsus ratio 1:3.1 and 1:3.2; hind tibia -f- tarsus length 604

and 616 /*; claws with large denticle. Antennae 9-segmented, 823 and 854 g

long.

Notes. — This is the most unusual known species of Heliococcus in

North America because of the large clusters of crateriform tubular

ducts. It is most closely related to H. deserticola, herein described as

new. H. clemente is also similar to H. atriplicis McKenzie in that both

species have less than 15 pairs of cerarii. H. atriplicis differs in lacking

crateriform tubular-duct clusters and in having a few multilocular disk

pores around the vulva. H. clemente has many crateriform tubular-

duct clusters and lacks multilocular disk pores.

H. clemente shows a remarkable resemblance to the Russian species

H. cydoniae. Both species possess large clusters of crateriform tubular

ducts and have small-sized crateriform ducts that lack associated setae.

They can be readily separated as follows. H. cydoniae has multilocular

disk pores surrounding the vulva, translucent pores on the hind coxae,

no pores on the hind tibiae, and 18 pairs of cerarii. H. clemente has

no multilocular disk pores, no translucent pores on the hind coxae, pores

present on the hind tibiae, and no more than 4 pairs of cerarii.

Vol. 50, No. 2, April 1974

183

Heliococcus cleserticola, new species

(Fig. 2-5)

Suggested common name. — Desert crateriform mealybug.

Type material. — Holotype adult female, 10 mi. N. Alamo, Lincoln County,

Nevada, 5 July 1970, on Ambrosia dumosa (Compositae) , D. R. Miller, (USNM).

Paratypes are as follows: Arizona: Welton, Yuma County, 27 March 1940, on

Encelia farinosa (Compositae), R. C. Dickson. California: Niland, Imperial

County, 19 March 1971, on Ambrosia dumosa, R. D. Goeden and D. W. Ricker;

Picacho Peak, Imperial County, 14 December 1970 and 13 April 1971, on Ambrosia

ilicifolia, R. D. Goeden and D. W. Ricker; 3 mi. N. Shoshone, Inyo County, 12

June 1963, on Encelia frutescens and Eriogonum foliolosum (Polygonaceae) ,

D. R. Miller; Palm Desert, Riverside County, 28 April 1971, beaten from Am-

brosia dumosa, R. D. Goeden and D. W. Ricker; Palm Springs, Riverside County,

29 April 1971, beaten from Ambrosia dumosa, R. D. Goeden and D. W. Ricker;

Morongo Valley, San Bernardino County, 9 June 1963, on Baccharis sp. (Com-

positae), D. R. Miller; Twenty -nine Palms, San Bernardino County, 25 March

1970, 20 May 1970, 20 May 1971, Ambrosia dumosa, R. D. Goeden and D. W.

Ricker; Yucca Valley, San Bernardino County, 20 May 1970, Ambrosia dumosa, R.

D. Goeden and D. W. Ricker. Nevada: same data as for holotype. Texas: Presidio,

Presidio County, 17 May 1951, on Viguiera stenoloba (Compositae), J. H. Russell.

Mexico: 10 mi. W. Zacapu, Michoacan, 6 March 1972, under rock, D. R. Miller

and F. D. Parker.

The 33 paratypes will be placed in the following collections: British Museum

(Natural History), London; California Department of Agriculture, Sacramento;

Florida State Collection of Arthropods, Gainesville; Museum National D’His-

toire Naturelle, Paris, France; Mexican National Collection, Mexico City; Uni-

versity of California, Davis; University of California, Riverside; University of

Hawaii, Honolulu; U. S. National Museum (Natural History), Beltsville, Mary-

land; Virginia Polytechnic Institute and State University, Blacksburg; Zoological

Institute, Academy of Sciences, Leningrad, USSR.

Adult female (Fig. 2). — Holotype, mounted, 2.6 mm long (paratypes 2. 1-3.8),

1.8 mm wide (paratypes 1. 2-2.7) . Body oval, anal lobes strongly protruding and

heavily sclerotized, with 3 large-sized crateriform tubular ducts on each lobe

(paratypes with 3-5, av. 3.5).

Dorsum with 17 pairs of cerarii. Anal-lobe cerarii each with 2 elongate, conical

setae, 4 or 5 trilocular pores, and large area of sclerotization. Remaining cerarii

each with 2 somewhat smaller conical setae, 2-4 trilocular pores, and little or no

basal sclerotization; ocular cerarii each with 3 conical setae. Multilocular disk

pores and quinquelocular pores absent. Trilocular pores abundant. Discoidal pores

sparce. Crateriform tubular ducts of 3 sizes: large-sized ducts scattered over

surface, most abundant near body margin, with 2-5 associated setae, marginal

ducts sometimes surrounded by cluster of 4-11 (av. 7) small-sized crateriform

tubular ducts; medium-sized ducts present in medial and mediolateral areas, with

1 or 2 associated setae; small-sized ducts in clusters associated with larger ducts

only, normally without associated setae. Body setae conical.

Anal-ring apical, bent over apex of abdomen, with 3 or 4 rows of pores; each

of 6 anal-ring setae noticeably longer than greatest diameter of ring.

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Fig

Heliococcus deserticola, adult female

Vol. 50, No. 2, April 1974

185

Venter with conspicuous, elongate anal-lobe sclerotization. Multilocular disk

pores absent. Quinquelocular pores present on medial and mediolateral areas.

Trilocular pores present in mediolateral and lateral areas, and in medial areas

of abdominal segments X-VI. Discoidal pores sparce. Crateriform tubular ducts of 2

sizes, same as larger and smaller sizes on dorsum: larger size always with associated

cluster of small-sized ducts, restricted to lateral areas, with 2-6 associated setae;

smaller size present in mediolateral and lateral areas, sometimes in clusters

around larger ducts, normally without associated setae. Oral-collar tubular ducts

with small dermal rim, restricted to mediolateral areas of abdominal segments

VIII and VII. Body setae of 2 kinds: bristle-shaped setae present medially and

laterally; conical setae present mediolaterally and laterally.

Circulus present. Legs with very small translucent pores restricted to dorsal

surface of hind tibiae; hind tibia/tarsus ratio 1:3.2 and 1:3.3 (para types 1:3.3-

1:3.6, av. 1:3.5) ; hind tibia tarsus length 573 and 567 /x (paratypes 573-763 /x,

av. 671) ; claws each with denticle. Antennae 9-segmented, 713 and 732 /x long

(paratypes 683-946 /x, av. 836).

Variation . — Paratypes differ from holotype as follows: with 15 to 18 pairs of

cerarii, normally with 18, with 2-4' setae, normally 2, in each ocular cerarius, some-

times with crateriform tubular-duct clusters more numerous along margin of dor-

sum, sometimes with small-sized crateriform ducts sparcely scattered over dorsal

surface, occasionally with 1 associated seta, with 2-16 (av. 7) smaller ducts in

clusters around larger ducts, with variable numbers of crateriform tubular-duct

clusters along body margin, sometimes with ventromedial trilocular pores restricted

to posterior 3 abdominal segments, without oral-collar tubular ducts on 1

specimen.

Notes . — The above description is based on 16 specimens from 8

localities.

H. deserticola is most closely related to H. clemente but differs in pos-

sessing 15-18 pairs of cerarii, crateriform tubular-duct clusters which

are restricted to the body margin, crateriform clusters which are com-

posed of 2-16 (av. 7) small-sized ducts, setae associated with some

small-sized crateriform ducts, 3 sizes of crateriform ducts. H. clemente

has 3 pairs of cerarii, crateriform tubular-duct clusters which are

scattered over the dorsum, crateriform clusters which are composed of

13-35 (av. 25) small-sized ducts, no setae associated with the small-

sized ducts, and 2 sizes of crateriform ducts.

H. deserticola has been confused with H. stachyos. These species

can be separated as follows: H. stachyos normally has multilocular

disk pores near the vulva and lacks clusters of crateriform tubular

ducts ; H. deserticola lacks multilocular pores and has clusters of crateri-

form tubular ducts.

H. deserticola shows a remarkable resemblance to the Russian species

H. kurilensis in that both have clusters of small-sized crateriform tubular

ducts surrounding a single larger duct. Unlike H. clemente and H.

186 The Pan-Pacific Entomologist

Fig. 3. Heliococcus deserticola, third instar female.

Vol. 50, No. 2, April 1974

187

Fig. 4.

Heliococcus deserticola, second instar male

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cydoniae, H. deserticola and H. kurilensis have only small clusters of

these ducts. H. kurilensis differs from H. deserticola in having the

crateriform clusters scattered over the entire dorsum and in having

multilocular disk pores near the vulva; H. deserticola has the crateri-

form clusters restricted to the marginal areas and has no multilocular

disk pores.

Third instar female (Fig. 3). — Same as for adult female except as follows:

mounted, 1. 4-2.1 mm long, 0.9-1. 3 mm wide. Anal lobes each with 2 large-sized

crateriform tubular ducts.

Dorsum with 18 pairs of cerarii. Anal-lobe cerarii each with 2-5 trilocular

pores. Discoidal pores absent or sparce. Crateriform tubular ducts of 3 sizes:

larger size with marginal ducts sometimes with 2-10 (av. 5.1) associated smaller-

sized crateriform tubular ducts; medium-sized ducts present in medial areas of

some posterior abdominal segments; small-sized ducts associated with larger ducts

and scattered in small numbers in mediolateral and lateral areas.

Venter with crateriform tubular ducts of 2 sizes: larger size present on margin

of abdominal segment VII, surrounded by cluster of small-sized ducts. Oral-

collar tubular ducts absent.

Legs without translucent pores; hind tibia/tarsus ratio 1: 1.7-1: 1.8 (av. 1:1.8) ;

hind tibia + tarsus length 311-415 (av. 366) /a. Antennae 7- or 8-segmented, 421-

519 (av. 458) ^ long.

Notes . — The above description is based on 2 specimens from 2 localities.

Second instar male (Fig. 4). — Same as for adult female except as follows:

mounted 1.1 mm long, 0.7 mm wide. Anal lobes each with 1 large-sized crateri-

form tubular duct.

Dorsum with 17 pairs of cerarii. Anal-lobe cerarii each with 2 or 3 trilocular

pores. Remaining cerarii sometimes indefinite, with widely separated cerarian

setae. Crateriform tubular ducts of 2 sizes: large-sized ducts in small numbers

over surface, with 1-4 associated setae, marginal ducts occasionally with 1-3 as-

sociated small-sized ducts; small-sized ducts scattered over surface. Oral-collar

tubular ducts of 2 sizes: shorter ducts restricted to lateral areas of anterior ab-

dominal segments, thorax, and head, of same size and shape as those on dorsum;

longer ducts present medially near legs.

Legs without translucent pores; hind tibia/tarsus ratio 1:1.4; hind tibia -f-

tarsus length 226 (jl . Antennae 7-segmented, 317 /j. long.

Notes . — The above description is based on 1 specimen.

First instar (Fig. 5; Sexes not distinguished). — Same as for adult female ex-

cept as follows: mounted, 0.5 mm long, 0.2-0.3 mm wide. Anal lobes lightly

sclerotized, without crateriform tubular ducts.

Dorsum with 16—18 indefinite cerarii. Anal-lobe cerarii each with 1 trilocular

pore. Remaining cerarii with 1 associated trilocular pore of same size and shape

as other triloculars; ocular cerarii normally absent, when present, with 2 conical

setae. Trilocular pores scattered over surface, arranged in 4 or 5 pairs of longi-

tudinal lines on abdomen. Discoidal pores and crateriform tubular ducts absent.

Body setae arranged in 4 pairs of longitudinal lines on abdomen and head, median

lines sometimes with additional row on thorax.

Anal ring with 2 rows of pores.

Vol. 50, No. 2, April 1974

189

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Venter with quinquelocular pores normally distributed as in figure 5, sometimes

with only 1 pore near mouthparts, occasionally with 3 pores near anterior spiracle

and 2 near posterior spiracle. Trilocular pores nearly always present near spiracles,

rarely with 1 or 2 present near body margin. Discoidal pores restricted to medio-

lateral areas. Crateriform and oral-collar tubular ducts absent. Body setae bristle

shaped.

Legs without translucent pores; hind tibia/tarsus ratio 1: 1.0-1: 1.1 (av. 1:1.1) ;

hind tibia -f- tarsus length 168-188 (av. 177) /x. Antennae 6-segmented, 225-

243 (av. 234) /x long.

Notes. — The above description is based on 8 specimens from 2

localities.

First instar nymphs of North American Heliococcus species have not

been described previously. The first instars of Heliococcus deserticola ,

Phenacoccus dearnessi King (see Miller and Appleby, 1971), and

Heterococcus spp. (see Miller, in press) are similar. The first instar

of H. deserticola differs from that of P. dearnessi and Heterococcus

spp. in having a circulus, conspicuous ostioles, and one size of trilocular

pore.

Acknowledgments

I thank Robert 0. Schuster, University of California, Davis, Richard

D. Goeden and Donald W. Ricker, University of California, Riverside,

for the loan of specimens.

I thank my wife, Judy, for her assistance in typing the drafts of this

manuscript and for her comments and criticisms. To Louise M. Russell,

Ronald W. Hodges, Robert W. Carlson, and Richard E. White, System-

atic Entomology Laboratory, ARS, USD A, I give thanks for their

criticisms of this manuscript.

Literature Cited

Baerensprung, F. von 1849. Beobachtung fiber einige einheimische Arten aus

der Familie der Coccinen. Ztg. Zool. Zoot. Palaeozool., 1: 165-170,

173-176.

Balachowsky, A. S. 1953. Sur un Heliococcus Sulc nouveau, vivant a 3600

metres d’altitude dans les Alpes du Dauphine et du Tyrol. Mitt. Schweiz.

Entomol. Ges., 26: 236-240.

Bazarov, B. 1963. Two new species of coccids from Tadzhikistan. (In Russian.)

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Borchsenius, N. S. 1941. New pest of mulberry — the central Asiatic cochineal,

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Borchsenius, N. S. 1949. Fauna of USSR. Homoptera, Pseudococcidae. (In

Russian.) Akad. Nauk Zool. Inst. (n.s. 38), 7: 1-383.

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Borchsenius, N. S. 1956. Notes on the Coccoidea of Korea. Entomol. Obozr.,

35: 671-679.

Borchsenius, N. S. 1958. Notes on the Coccoidea of China. II. Descriptions of

some new species of Pseudococcidae, Aclerdidae and Diaspididae. (In

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Borchsenius, N. S. 1962. Notes on the Coccidea of China. XI. New genera

and species of Pseudococcidae. (In Russian.) (Akad. Nauk SSSR

Zook Inst. Trudy, 30: 221-244.

Borchsenius, N. S. and E. M. Tereznikova. 1959. Two new species of mealy

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491-494.

Bueker, E. D. 1930. Phenacoccus wilmattae Ckll. Can. Entomol., 62: 93-94.

Cockerell, T. D. A. 1901. The New Mexico Coccidae of the genus Ripersia.

Ann. Mag. Nat. Hist. (ser. 7), 8: 51-57.

Danzig, E. M. 1971. New and little-known species of mealy-bugs from the far

east of USSR. (In Russian.) Entomol. Obozr., 50: 366-390.

Danzig, E. M. 1972. Contribution to the fauna of the white flies and scale in-

sects of Mongolia. (In Russian.) Nasekomye Mongolii, Mongol Orny

Shav’zh pp. 325-348.

Ehrhorn, E. M. 1900. New Coccidae from California. Can. Entomol., 32:

311-318.

Ferris, G. F. 1950. Atlas of the scale insects of North America. Series V.

Pseudococcidae (Part I). Stanford Univ. Press, California, 278 pp.

Ferris, G. F. 1953. Atlas of the scale insects of North America. Volume VI.

The Pseudococcidae (Part II). Stanford Univ. Press, California, pp.

279-506.

Goux, L. 1931. Notes sur les coccides de la France. (3 e note). Description

d’une espece nouvelle. Bull. Soc. Entomol. Fr., 1931: 113-118.

Goux, L. 1934. Notes sur les coccides de la France. (9 e note) . Contribution

a l’etude du genre Heliococcus avec description de deux especes nou-

velles. Bull. Soc. Entomol. Fr., 39: 164^171.

Goux, L. 1953. Notes sur les coccides de la France (42 me note). Etude d’un

Heliococcus nouveau. Bull. Mens. Soc. Linn. Lyon, 22: 104^-109.

Green, E. E. 1925. Notes on the Coccidae of Guernsey (Channel Islands),

with descriptions of some new species. Ann. Mag. Nat. Hist., 16: 516-

527.

Hollinger, A. H. 1917. A new species of Phenacoccus. Can. Entomol., 49:

281-284.

Kanda, S. 1934. Two new genera of Japanese Coccidae. Insect World, 38:

308-313.

Kanda, S. 1935. On the genus Heliococcus. Annot. Zool. Jap., 15: 70-77.

Kanda, S. 1959a. Two new species of the Pseudoccidae. Kontyu, 27: 177-180.

Kanda, S. 1959b. Two new genera and two new species of the Pseudococcidae.

Kontyu, 27: 239-243.

King, G. B. 1902. Two new ants’-nest coccids from Texas. Can. Entomol., 34:

285-286.

Kuwana, S. I. 1907. Coccidae of Japan, I. A synoptic list of Coccidae of Japan

with descriptions of thirteen new species. Imp. Centr. Agric. Exp. Stn.

Bull., 1: 177-207.

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Kuwana, S. I. 1917. A check list of the Japanese Coccidae. Yokohama, Japan.

19 pp ‘

Laing, F. 1929. Descriptions of new and some notes on old species of Coccidae.

Ann. Mag. Nat. Hist., 4: 465-501.

Lindinger, L. 1912. Die Schildlause Europas, Nordafrikas und Vorderasiens,

einschlieblich der Azoren, der Kanaren und Madeiras. Ulmer, Stuttgart.

388 pp.

Lobdell, G. H. 1930. Twelve new mealybugs from Mississippi. Ann. Entomol.

Soc. Amer., 23: 209-236.

MacGillivray, A. D. 1921. The Coccidae. Scarab Co., Illinois. 502 pp.

McDaniel, B. 1962. A new species of scale insect from the Rio Grande Valley,

Texas. Tex. J. Sci., 14: 323-327.

McKenzie, H. L. 1960. Taxonomic study of California mealybugs with descrip-

tions of new species. Hilgardia, 29: 681-770.

McKenzie, H. L. 1964. Fourth taxonomic study of California mealybugs, with

additional species from North America, South America, and Japan.

Hilgardia, 35: 211-272.

McKenzie, H. L. 1967. Mealybugs of California with taxonomy, biology, and

control of North American species. Univ. California Press, California.

525 pp.

Miller, D. R. 1974. A systematic revision of the genus Heterococcus , with a

diagnosis of the genus Brevennia. (in press)

Miller, D. R. and J. E. Appleby. 1971. A redescription of Phenacoccus dear-

nessi. Ann. Entomol. Soc. Amer., 64: 1342-1357.

Sanders, J. G. 1902. A new Phenacoccus on Platanus occidentalis. Ohio Nat.,

2: 284-286.

Sulc, K. 1912. Heliococcus bohemicus n. gen. n. sp. Cesk. Spolec. Entomol.

Casopis., 9: 39-48.

Takahashi, R. 1930. Observations on the Coccidae of Formosa. Part II. Dept.

Agric. Gov. Res. Inst. Formosa, Rpt. no. 48, 45 pp.

Takahashi, R. 1951. Some mealy bugs from the Malay Peninsula. Entomol.

Soc. India (1950), 12: 1-22.

Ter-Grigorian, M. A. 1967. Two new species of the genus Heliococcus Sulc

from Armenia. (In Russian.) Dokl. Akad. Nauk Arm. SSR, 44: 134-

138.

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Entomol., 12: 1-79.

Vol. 50, No. 2, April 1974

193

Mealybugs of San Clemente Island, California

(Homoptera: Coccoidea: Pseudococcidae)

Douglass R. Miller

Systematic Entomology Laboratory, USDA, ARS, Beltsville, Maryland 20705

In earlier papers (Miller, 1971, 1973) I described the mealybug

faunas of Santa Cruz and San Miguel Islands and presented brief dis-

cussions relating to the Southern California Island biota.

In April 1973 Dr. Arnold Menke and I collected on San Clemente

Island for eight days. Although this was an inadequate amount of

time to thoroughly sample the island pseudococcids, the material ob-

tained should allow some tentative comparisons of the mealybug fauna

of this island with those of Santa Cruz and San Miguel Islands.

San Clemente Island is located approximately 49 statute miles from

the nearest point on the mainland and is 21 miles from the closest

island, Santa Catalina, which intervenes between the nearest mainland

point and San Clemente. The island is approximately 21 miles long,

about 4 miles across at its widest point, and approximately 56 square

miles in land area (Philbrick, 1967). The long axis of the island is

oriented approximately northwest, nearly parallel to the near mainland

coast. Of the several geological studies of the island, two of the more

detailed ones were published by Olmstead (1958) and Smith (1898).

Geologically, San Clemente is the emergent portion of a tilted, slightly

arched block of the earth’s crust. A major fault is present on the sea

floor at the northeastern edge of the San Clemente block, and, although

movement of this fault is thought to be primarily horizontal, nearly

the entire northeastern side of San Clemente Island is a single, large

fault escarpment. The northeastern side of the island is very steep

in contrast to the southwestern side which generally slopes gently. Both

sides of the island are deeply cut by geologically young, often precip-

itous canyons. The highest point is Thirst (elevation 1,967 feet),

located near the northeastern escarpment. The southwestern side of the

island, up to 1,500 feet, shows some of the best examples of marine

terracing on the Southern California Islands, with as many as 20 separ-

ate terraces visible on some slopes. The island is principally com-

posed of volcanic rocks of probable Miocene age. Sedimentary marine

deposits (Miocene), sand deposits (Pleistocene? and Recent), and

alluvial deposits (Recent) overlying the volcanic rocks are scattered

locally on the island. Normally the soil covering is deposited in rela-

tively thin layers and has a high clay content. San Clemente is apparently

The Pan-Pacific Entomologist 50 : 193 - 202 . April 1974

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no older than the Miocene and may be as recent as the Early to Mid-

Pliocene. Most evidence suggests that this island has never been con-

nected with the mainland or the other islands, although at times of lower

sea levels, particularly during the Pleistocene, San Clemente undoubtedly

was closer to other land masses than at present. As is evident from the

lack of marine deposits and marine terraces above 1,500 feet elevation,

the areas of the island above this level have probably been continually

emergent since Miocene or Pliocene times.

Data on the weather of the island are scant, and have been sum-

marized by Raven (1963) as follows: “ . . . the east (northwestern)

side of the island is apparently moister than the west (northeastern),

and it is likely that the main tracks of the storms from north to south

are often deflected down along the east side of the island. Generally

the wind is from the sea, over the island and onto the mainland; but

during the Santa Ana cycles of winter, violent and sustained winds

blow off the mainland and from Santa Catalina Island toward the

east side of San Clemente. These Santa Ana winds may obviously be

of importance in transporting seeds (or first instar mealybugs) to the is-

land.” Most of the rainfall occurs during the winter months, but mois-

ture producing fogs are common during rain-free times of the year.

No permanent streams are present although intermittent flows are

common during the rainy season. Olmstead (1958) stated that the

deep plunge pools formed by intermittent waterfalls contain water

throughout the year. Additional small impoundments have been con-

structed, but basically the island is without water during the dry

months.

The predominant habitat types on the island are flat grassland areas,

hilly cactus areas, deep precipitous canyons, steep cliffs, volcanic re-

gions often with many small caves, recent sand dunes, fossil sand dunes,

sand beaches, and shingle beaches.

The island reptile fauna includes only two lizard species, one belongs

to an endemic genus known from San Clemente, San Nicolas, and

Santa Barbara Islands ; there are no amphibians ( Savage, 1967 ) . The

indigenous terrestrial mammal fauna of the island is composed of four

bat species, three mice species including one endemic sub-species, and

one fox species which is an insular endemic with an endemic race on

San Clemente (von Bloeker, 1967). The mammals introduced by man

are pigs, goats, cats, and deer. The aquatic mammal fauna consists of

three species of pinnipeds (Bartholomew, 1967) and the sea otter (von

Bloeker, 1967). Diamond (1969) reported 24 species of land birds

on San Clemente in 1968. To my knowledge no detailed study of the

Vol. 50, No. 2, April 1974

195

insect fauna of San Clemente has been published, although several

short papers have appeared in the series “Contributions from the Los

Angeles Museum — Channel Islands Biological Survey.”

Most of the native vegetation of the island has been ravaged by the

feeding of a large population of feral goats. Because of this, the is-

land is sparsely vegetated, and introduced herbs, mostly grasses, pre-

dominate. Plant species once apparently widespread are now either

extinct or are restricted to small populations in areas inacessible to

goats. Some of the best refuges on the island are located in the pre-

cipitous canyons, particularly on the northeastern side. The upper

reaches of these canyons are of particular interest because they have

not been inundated by the sea since Miocene or Pliocene times. A

flora of the island published by Raven (1963) is the source of much

valuable information. Additions to the flora have been published by

Raven (1965, 1967) and Thorne (1969). Of the Channel Islands,

San Clemente has the most interesting flora, despite the larger land

area and seemingly greater habitat diversity of some of the other is-

lands. It supports more insular endemics (43, 13 of which occur ex-

clusively on San Clemente) than the other Channel Islands, evidently

because its isolation has allowed little floral interchange with other

land areas.

San Clemente Island Mealybugs

Knowledge of the unusual San Clemente flora and of the similarities

between the dispersal mechanisms of scale insects and plants (Miller,

1971), encouraged me to look to San Clemente as the most promising

source of unusual mealybugs in the Southern Channel Islands. Un-

fortunately, collecting mealybugs on this island is difficult because

most of the more interesting plants occur in nearly inaccessible areas

and the heavy clay soil generally present is unsuitable for most pseu-

dococcid species. The following list includes 9 genera and 11 species.

No mealybugs have been reported from this island previously. Plant

names are as listed by Raven (1963).

Chorizococcus

cibroniae McKenzie

1. Shifting Sands, 14 May 1973, Abronia. umbellata (roots and decumbent

stems)

Discococcus

simplex Ferris

1. 0.25 mi. N. Boulder, 10 May 1973, Stipa sp. and Avena sp. (on crown)

2. canyon N. Boulder, 11 May 1973, Stipa sp. (in sheaths near crown)

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3. China Canyon, 13 May 1973, Stipa sp. (in sheaths)

4. Eel Point, 8 May 1973, Stipa sp. (in sheaths near crown)

5. Pyramid Head, 12 May 1973, Stipa sp. (on crown)

6. Seal Cove, 9 May 1973, Stipa sp. (in sheaths)

7. near mouth of Wall Rock Canyon, 9 May 1973, Stipa sp. (on crown)

8. near White Rock, 10 May 1973, Stipa sp. and Avena sp. (in sheaths near

crown)

9. Wilson Cove, 8 May 1973, Avena sp. (on crown)

Distichlicoccus

salinus ( Cockerell )

1. China Point, 13 May 1973, Distichlis spicata (on leaf blade)

2. Shifting Sands, 14 May 1973, Distichlis spicata (on leaf blade)

Heliococcus

clemente Miller

1. Pyramid Head, 12 May 1973, under rock

Paludicoccus

distichlium (Kuwana)

1. China Point, 13 May 1973, Distichlis spicata (in leaf blade sheath)

Phenacoccus

eschscholtziae McKenzie

1. China Point, 13 May 1973, Lotus argophyllus subsp. ornithopus (?) (on

main roots)

solani Ferris

1. S. West Cove, 7 May 1973, Franseria chamissonis (decumbent branches)

2. 2 mi. SE West Cove, 11 May 1973, Astragalus miguelensis and Atriplex

semibaccata (roots)

Puto

yuccae (Coquillett)

1. Eel Point, 8 May 1973, Suaeda calif ornica (on roots)

Spilococcus

corticosus McKenzie

1. canyon N. Boulder, 11 May 1973, Artemisia calif ornica (under bark on

stems)

keiferi McKenzie

1. China Point, 13 May 1973, Lotus argophyllus subsp. ornithopus (?) and

Suaeda californica (on roots)

2. Eel Point, 9 May 1973, Astragalus miguelensis, Atriplex semibaccata, and

Suaeda californica (on roots)

3. Pyramid Head, 12 May 1973, Phacelia floribunda (?) and, Suaeda cal-

ifornica (on roots) , and under rock

4. near mouth of Wall Rock Canyon, 9 May 1973, Suaeda californica (on

roots)

5. 2 mi. SE West Cove, 11 May 1973, Suaeda californica (on roots)

Trionymus

smithii (Essig)

1. Wilson Cove, 7 May 1973, Avena sp. (leaf sheath)

Vol. 50, No. 2, April 1974

197

Discussion

Perhaps my most exciting find on San Clemente was the endemic

mealybug, Heliococcus clemente Miller. I believe this is the first of

several endemics which may be discovered when the canyons on the

northeastern side of the island are intensively collected. H. clemente

is most similar to Heliococcus deserticola Miller, a species that occurs

in the deserts of the southwestern U. S. and is relatively close to another

desert species, Heliococcus atriplicis McKenzie. H. clemente appears

to be more similar to xeric southern desert species than to the more

common and widespread mesic species (such as H. adenostomae Mc-

kenzie, H. osborni (Sanders), or H. stachyos (Ehrhorn).

An unidentified species, apparently a second endemic, has been en-

countered on both San Clemente and Santa Cruz Islands. On Santa

Cruz it was collected at several localities ranging from the higher eleva-

tions of the north ridge to the coastal areas near Coches Prietos. It

has been collected in spring on a varied range of hosts, on rootlets in

heavy, clay soil. On San Clemente, the apparent same species was found

near Boulder, in heavy, clay soil on small roots of Malacothrix foliosa

(Compositae) , Lupinus bicolor (Leguminosae) , and other small dicoty-

ledonous plants. I found only first instar nymphs at both locations,

and suspect that adults occur much later in the year. Until adult fe-

males can be found, this pseudococcid should remain unnamed, but it

is so different from the known first instar nymphs of any other North

American mealybug, that I am fairly certain it is a Channel Island

insular endemic. It is quite probable that this mealybug represents

an endemic genus.

Using the work of McKenzie (1967) as a source for California

pseudococcid distribution patterns, the 10 nonendemic species occurring

on San Clemente are distributed on the mainland as follows: one is

present throughout most of continental California, three occur exclu-

sively in coastal mountain ranges, one occurs in the coastal ranges and

the southern California deserts, two occur in the coastal ranges and the

ocean beach areas, two occur in saline areas, and one occurs exclusively

on ocean beach areas. These data suggest that most mealybug species

known to occur on San Clemente Island are most closely allied to the

aggregation of species found in coastal areas on the California main-

land; in fact, all of the known San Clemente species except H. clemente

are known to inhabit coastal areas in part.

One species shows a disjunct distribution pattern. Discococcus sim-

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plex is not known on the portion of the mainland closest to San Cle-

mente Island but occurs approximately 190 miles to the north. With

the limited amount of material available, it has not been possible to

make general statements concerning the number of species on San

Clemente compared to a similarly sized area on the mainland. How-

ever, the absolute number of individual specimens on the island is small

compared to the mainland because an average of only three mealy-

bug collections per day was made on San Clemente, whereas I find

an average of about ten mealybug collections per day on the California

mainland.

Discococcus simplex is unusually abundant on San Clemente. Pre-

viously, this species was known by less than 10 specimens from two

mainland localities. On San Clemente, with some diligence, D. simplex

can be collected in most grassland areas on the island; it is normally

quite common on Stipa spp. The same species was collected once on

another island (Santa Cruz), where apparently it is rare.

Spilococcus keiferi is also unusually abundant. It is a fairly com-

mon coastal mainland pseudococcid which seems to prefer the roots and

decumbent branches of Franseria chamissonis in sandy soils. It has

occasionally been found on a few other hosts and has been collected

inland in two instances. Under normal circumstances only one or two

specimens of S. keiferi are collected at mainland localities, and they

usually are mixed with specimens of Chorizococcus abroniae. On San

Clemente this species seems to behave differently. Next to Discococcus

simplex, it was the most commonly encountered mealybug on the is-

land. It was collected on five plant species which are all new host records.

Although populations of Franseria chamissonis were examined exten-

sively, S. keiferi was never found on this host. Infestations of S. keiferi

normally were very large, and they never were mixed with Chorizococcus

abroniae. On San Miguel and Santa Cruz Islands S. keiferi did not

show characteristics of the San Clemente populations.

Similar insular differences in relative abundance and host preference

are known to occur in island populations of Puto yuccae. On the main-

land P. yuccae occurs on a variety of hosts and is common in many

habitats including the Sierra Nevada up to 6,000 feet, in chaparral

areas of the coastal mountain ranges, and in sand areas near the ocean.

On San Clemente the species was collected only once, even though pre-

ferred hosts were examined extensively. On San Miguel P. yuccae

was much more abundant. Although it apparently is restricted to sandy

Vol. 50, No. 2, April 1974

199

beach situations, it occurs on a variety of hosts and is quite common.

On Santa Cruz it is more abundant than on the other two islands. It

occurs in several types of habitats and has been recorded on seven dif-

ferent host plant species.

The mechanisms affecting differences in habitat preference, host

diversity, and relative abundance of mealybug species on islands appear

to be complex. A study of these mechanisms would undoubtedly give

insight into aspects of pseudoccid biology and ecology not presently

understood.

In previous papers on the Southern California Islands, I have not

mentioned morphological variation of insular pseudococcid species,

although some island populations are highly variable. At times this

variation has caused problems in placing the island species, but this

problem was anticipated. If the Southern California Islands possess

endemic forms at generic and specific levels, then they should contain

examples of endemism at intraspecific levels. In fact, this endemism

should be more common than at higher levels. Variation observed in

three mealybug species bears mention. Material identified as Puto

yuccae has shown considerable variation from island to island and from

island to mainland. Some insular specimens possess features of both

Puto echinatus McKenzie, a coastal mainland species, and P. yuccae.

The greatest amount of variation was noted on specimens from San

Miguel Island where typical representatives of both P. ecliinatus and

P. yuccae were found, in addition to many intermediate individuals.

On Santa Cruz variation was less marked, but still considerable. On

San Clemente P. yuccae possessed key characters of both species. With-

out more material, particularly from the mainland coastal areas, the

identity of this species must remain tentative.

There are a number of possible explanations for this type of in-

sular variation. For example, P. echinatus may be a recent coastal de-

rivative of the inland species P. yuccae , or vice versa, with intermediate

forms persisting on the islands; P. echinatus and P. yuccae may be

the same species; annectant forms may be hybrids; and there are other

possibilities. Variation seen in insular populations of Spilococcus keiferi

and Trionymus smithii show the same general pattern as mentioned

above. S. keiferi varies between typical S. keiferi and individuals

approaching S. atriplicis (Cockerell) or S. nototrichus Miller and Mc-

Kenzie. T. smithii varies between specimens typical of California forms

of T. smithii and material approaching T. winnemucae McKenzie or

T. nanus Cockerell. Although annectant specimens cause concern to

200

The Pan-Pacific Entomologist

the taxonomist, they point out possible relationships which otherwise

might go undetected.

Comparison of the Mealybug Faunas of

San Clemente, San Miguel, and Santa Cruz Islands

Notable similarities exist in the three pseudococcid faunas. (1) All

are depauperate aggregations of species similar to the pseudococcid

fauna on the adjacent southern California mainland. (2) All are pri-

marily composed of coastal or coastal mountain species. (3) All contain

representatives of the following species: Chorizococcus abroniae, Dis-

tichlicoccus salinus, Paludicoccus distichlium, Puto yuccae, and Spilo-

coccus keiferi.

The mealybug fauna of San Clemente most closely resembles the

fauna found on Santa Cruz. Of the 12 species (including the unde-

scribed form) known to occur on San Clemente, nine occur on Santa

Cruz while only six are known on San Miguel. The pseudococcids of

both San Clemente and Santa Cruz, with the exception of Discococcus

simplex on San Clemente, occur in localized areas; whereas on San

Miguel, several species are widespread and common. Both San Clemente

and Santa Cruz have representative species which do not occur on the

adjacent mainland but are found farther north in California, and both

islands have specimens of the apparent insular endemic known only

from immatures.

A feature found within the mealybug faunas of San Clemente and

San Miguel, but not within the Santa Cruz fauna, is the presence of

species which show more diversity in habitat and host preference than

they do on the mainland.

The San Clemente pseudococcid fauna is unique in the presence

of an endemic species. Its resemblance to species occurring in the

desert areas of the southwestern U. S. suggests the presence of southern

xeric elements within the San Clemente fauna.

Summary

Nine genera and 11 species of mealybugs are known on San Clemente

Island. Heliococcus clemente is endemic, and another probable insular

endemic known only from immatures, is reported from San Clemente

and Santa Cruz Islands. The pseudococcid fauna of San Clemente is

most like the aggregation of mealybug species found on the adjacent

mainland, but contains elements of both northern and southern faunas.

The flora and mealybug fauna of San Clemente have many similar

attributes. Both show affinities to their respective counterparts on the

Vol. 50, No. 2, April 1974

201

adjacent mainland, both contain endemics, both possess species with

disjunct distribution patterns, and both contain small components of

northern and southern biotas.

Acknowledgments

I am grateful to Lt. Commander W. G. Kay, Officer in Charge, San

Clemente Island Facility, for his assistance and cooperation during

my stay on San Clemente. Thanks are also due to Chief West who

made some of the arrangements for transportation on the island, pointed

out areas which might be of special interest, and allowed us to collect

in normally inaccessible regions.

Special appreciation is expressed to Dr. Arnold S. Menke for his

assistance in collecting scale insects while on the island.

I also thank my wife Judy Miller for her help in typing the drafts

of this article and for her comments on its content. I appreciate the

help of Louise M. Russell, Raymond J. Gagne, Douglas C. Ferguson,

and Ronald W. Hodges, Systematic Entomology Laboratory, ARS,

USDA, for their careful criticisms of this manuscript.

Literature Cited

Bartholomew, G. A., 1967. Seal and sea lion populations of the California

Islands. Pp. 229-244 in R. N. Philbrick ( ed. ) , (For complete citation

see Philbrick) .

Diamond, J. M. 1969. Avifauna equilibria and species turnover rates on the

Channel Islands of California. Proc. Nat. Acad. Sci., 64(1) : 57-63.

McKenzie, H. L. 1967. Mealybugs of California with taxonomy, biology, and

control of North American species. Univ. Calif. Press, Berkeley, 525

pp.

Miller, D. R. 1971. Mealybugs of Santa Cruz Island, California. Pan-Pac.

Entomol., 47 (4) : 293-303.

1973. Mealybugs of San Miguel Island, California. Pan-Pac. Entomol.,

49(3) : 264-269.

Olmstead, F. H. 1958. Geologic reconnaissance of San Clemente Island, Cal-

ifornia. Geol. Surv. Bull., 1071-B: 55-68.

Philbrick, R. N. Introduction. Pp. 3-8 in R. N. Philbrick (ed.), Proceedings

of the Symposium on the Biology of the Californa Islands. Santa

Barbara Botanic Garden, Calif., 341 pp.

Raven, P. H. 1963. A flora of San Clemente Island, California. Aliso, 5(3) :

289-347.

1965. Notes on the flora of San Clemente Island, California. Aliso 6: 11.

1967. The floristics of the California Islands. Pp. 57-67 in R. N. Philbrick

(ed.), (For complete citation see Philbrick).

Savage, J. M. 1967. Evolution of the insular herpetofaunas. Pp. 219-227 in

R. N. Philbrick (ed.), (For complete citation see Philbrick).

202

The Pan-Pacific Entomologist

Smith, W. S. T. 1898. A geological sketch of San Clemente Island. U. S.

Geol. Surv., Ann. Rpt. 18, (1896/97) pt. 2, pp. 459-496.

Thorne, R. F. 1969. A supplement to the floras of Santa Catalina and San

Clemente Islands, Los Angeles County, California. Aliso, 7 : 73-83.

von Bloeker, J. C., JR. 1967. Land mammals of the Southern California Islands.

Pp. 245-263 in R. N. Philbrick (ed.), (For complete citation see

Philbrick ) .

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203

SCIENTIFIC NOTES

The American Toad as Champion Carabid Beetle Collector. — The Ameri-

can Toad ( Bufo americanus Holbrook) has been reported as a predator of Carab-

idae (Coleoptera) by Kirkland (1897. Hatch Exp. Sta. Mass. Agric. Bull.,

46: 1-30; 1904. U. S. Dept. Agric. Farmer’s Bull., 196: 1-16), Garman (1901.

Kentucky Agric. Exp. Sta. Bull., 91: 60-68), Stoner (1937. Florida Entomol.,

19: 49-53), Smith and Bragg (1949. Ecol., 30: 333-349), and also by Bush (1959.

Herpetol., 15: 73-77).

From May to October 1973, I collected and analyzed the stomach contents of

342 American toads from southern Quebec in order to determine the extent of

predation on carabid beetles. Immediately after capture, the animals were killed

in hot water, their stomachs removed and the contents emptied into a small dish

of water. The total number of species of carabid beetles in the toads was 98,

and the maximum of individuals in one toad was 17. In addition, remains of

19 species were found in toads’ excreta. The diet of the American toad is de-

termined mostly by relative availability of prey items (Smith and Bragg 1949),

and selectivity of prey is probably a factor of minor importance. The amphibians

live, on the whole, in the same habitats as carabid beetles, and both are predomi-

nantly active nocturnally, partly explaining the large consumption of Carabidae

by toads.

The number of each species of prey is listed below. Numbers in parentheses

indicate additional individuals found in excreta. Agonurn sp., 6 (28) , A. creni-

striatum Leconte, 47, A. cupripenne Say, 12 (1), A. decentis Say, 1, A. harrisi Le-

conte, 1, A. melanarium Dejean, 8, A. placidum Say, 41 (6), A. propinqiium Gem-

minger and Harold, 1, A. puncticeps Casey, 1, A. retractum Leconte, 6, Amara sp.,

40 (8), A. aenea DeGeer, 41 (8), A. angustata Say, 2, A. apricciria Paykull, 2, A.

avida Say, 3, A. convexa Leconte, 2, A. discors Kirby, 2, A. familiaris Duftschmid,

1, A. impuncticollis Say, 3, A. latior Kirby, 4 (1), A. littoralis Mannerheim, 1, A.

musculis Say, 5, A. obesa Say, 14 (1). A. quenseli Schonherr, 33 (8), A. rubrica

Haldeman, 2, Anisodactylus discouleus Dejean, 1 , A. harrisi Leconte, 5, A. kirbyi

Lindroth, 1, A. rnerula Germar, 21, A. nigerrimus Dejean, 2, A. nigrita Dejean, 1,

A. rusticus Say, 3, A. sanctaecrucis Fabricius, 6, Anisotarsus nitidipennis Leconte, 2,

A. terminatus Say, 2, Badister notatus Haldeman, 1 , Bembidion frontale Leconte, 2,

B. nitidum Kirby, 3, B. patruele Dejean, 2, B. punctato striatum Say, 1, B. quadri-

maculatum oppositum Say, 8, B. stephensi Crotch, 1, B. tetracolum Say, 2, Bradycel-

lus sp., 1, B. nigriceps Leconte, 1, B. rupestris Say, 1, B. semipubescens Lindroth,

1, Calathus sp., 1, C. gregarius Say, 2, C. ingratus Say, 1 (2), Calleida punctata Le-

conte, 3, Calosoma calidum Fabricius, 1 (1), Carabus nemoralis Muller, 4, C. ser-

ratus Say, (1), Chlaenius lithophilus Say, 2, C. pensylvanicus Say, 1, C. tomentosus

Say, 4, C. tricolor Dejean, 1, Clivina sp., 1, C. fossor Linne, 13, Cymindis americana

Dejean, 1, C. borealis Leconte, 1, C. cribricollis Dejean, 3 (4), C. pilosa Say, 1,

Dicaelus politus Dejean, 1, Diplocheila obtusa Leconte, 2, Dromius piceus Dejean, 1,

Dyschirius globulosus Say, 4, Geopinus incrassatus Dejean, 22, Harpalus sp., 6 (3),

H. affinis Schrank, 72 (11), H. bicolor Fabricius, 9, H. caliginosus Fabricius, 1, H.

egregius Casey, (1), H. erraticus Say, 12, H. fallax Leconte, 1, H. fuliginosus Duft-

schmid, (1), H. funerarius Csiki, 1, H. herbivagus Say, 13, H. indigens Casey, 7,

H. lewisi Leconte, 3, H. opacipennis Haldeman, 5, H. pensylvanicus DeGeer, 13, H.

pleuriticus Kirby, 2 (2), H. viduus Leconte, 1, Loricera pilicornis Fabricius, 8,

204

The Pan-Pacific Entomologist

Metabletus americanus Dejean, 3, Notiophilus aquaticus Linne, 1, Olisthopus par-

matus Say, 1, Omophron americanum Dejean, 3, Patrobus longicornis Say, 1,

Pseudamara arenaria Leconte, 1 , Pterostichus sp., (6), P. adstrictus Eschscholtz,

1, P. corcicinus Newman, 1 (1), P. corvinus Dejean, 9 (1), P. lachrymosus New-

man, 1, P. leconteianus Lutshnik, (2), P. luctuosus Dejean, 4, P. lucublandus

Say, 27 (30), P. melanarius Illiger, 11, P. mutus Say, 10, P. pensylvanicus Leconte,

7 (2), P. tristis Dejean, 1, Selenophorus gagatinus Dejean, 3, Sphcieroderus lecontei

Dejean, 1 , Stenolophus comma Fabricius, 3, S. conjunctus Say, 4, Synuchus im-

punctatus Say, 11, Tachys incurvus Say, 1, undetermined species, 31 (17).

These data suggest that the American toad may not be entirely beneficial as

a predator of noxious insects, since many Carabidae, especially larger species, are

important predators of lepidopterous larvae and other destructive insects (Blatch-

ley, 1910. The Coleoptera or Beetles known to occur in Indiana; Balduf, 1935.

The Bionomics of Entomophagous Insects).

I am grateful to the following persons for their technical assistance: Pierre

Cloutier, Stephane Couture, Jean Dube, Paul Duval, Claude Gelinas, Jean-Pierre

Grenier, Raymond Hutchinson, Frangois Marcotte, Herve Perrault, Jean-Pierre

Pilotte, Paul Phaneuf and Jean-Pierre Tchang. — Andre Larochelle, College

Bourget, C. P. 1000, Rigaud, Quebec.

Notes on the general biology of the flatheaded fir borer Melanophila

drummondi Kirby reared from ponderosa pine (Coleoptera: Buprestidae).

— Melanophila drummondi Kirby is a common metallic wood borer of dying, slow

growing, freshly cut, or wind-thrown Douglas-fir, Pseudotsuga menziesii (Mirb.)

Franco, and western hemlock, Tsuga heterophylla (Raf.) Sarg., in Washington

state. Chamberlin (1926, The Buprestidae of North America exclusive of Mexico)

has cited many other host species. Barr (1971, in Hatch, The beetles of the Pacific

Northwest, Part V. University of Washington Press, Seattle, Washington) adds

white spruce, Picea glauca (Moench) Voss, to Chamberlin’s list of hosts for M.

drummondi. Anderson (1966, Forest and shade tree entomology) mentions pine

as being a host for M. drummondi, but does not indicate which species, nor could

I find any other reference identifying pine as a host for this buprestid. I recently

found M. drummondi to occur on ponderosa pine, Pinus ponderosa Laws.

Infestation of host material by M. drummondi occurs shortly after spring cut-

ting or windfall and extends through early fall at low elevations. At higher eleva-

tions, infestation may be delayed by a month or more due to slower maturation of

larvae and later emergence of adults. Maturation of larvae generally occurs within

a year, but may take several years when live trees are attacked (Anderson, 1966).

On March 15, 1973, I placed several 1 m long bolts of ponderosa pine cut from

a tree felled in the spring of 1972, 2 miles northeast of Liberty, Kittitas County,

Washington, into a plywood rearing chamber maintained at 20.6 °C and 78%

relative humidity. The first M. drummondi emerged on April 22, and emergence

continued until July 4, 1973, yielding a total of 12 males and 10 females.

Melanophila gentilis LeConte, a common buprestid inhabitant of ponderosa pine,

was also reared from the ponderosa pine bolts (30 $ $ , 22 $ $ ) .

The first insects noted to emerge were two individuals of the parasitoid

Vol. 50, No. 2, April 1974

205

Hymenoptera, Coeloicles brunneri Viereck on April 12. The emergence of bark-

or wood-inhabiting insects continued through July 16, 1973. Although C. brunneri

is primarily an ectoparasite on bark beetle larvae, it is sometimes associated with

other larvae as well. Four adults of the hymenopteran Atanycolus longifemoralis

Shenefelt emerged between May 7 and May 27. A. longifemoralis is a primary

ectoparasite on both M. drummondi and M. gentilis (Shenefelt, 1943. Res. Stud.

State Coll. Wash., 11: 51-163.).

Freshly emerged Melanophila adults were placed in cylindrical plastic con-

tainers for observation and were fed fresh ponderosa pine twigs and needles. Pine

bark and wood chips were provided for the beetles to walk on. A small specimen

vial cap was filled with tap water and placed in each container to supply moisture

and drinking water. Nearly all individuals of both M. drummondi and M. gentilis

fed on fresh ponderosa pine needles or twig phloem when needles became scarce.

In a feeding-choice experiment I found that adults of M. drummondi, reared

from Douglas-fir, fed on needles of Douglas-fir, western hemlock, or ponderosa

pine. However, closer examination of the needles of the three species revealed

feeding to be greater on Douglas-fir than on the others. Although this evidence is by

no means conclusive, it is conceivable that feeding preference is directly re-

lated to the host tree species in which the brood develops. — Donald W. Scott,

College of Forest Resources, University of Washington, Seattle, 98195.

Tolerance of Neotoma and Rattus to the Feeding of Triatoma . — Evidence

is accumulating that wild rodents can tolerate feedings of large numbers of blood

sucking triatomid bugs without lethal effects. These hematophagous insects normally

inhabit the houses of wood rats, Neotoma, with associated species of white-

footed mice, Peromyscus. Wood (1943, Am. J. Trop. Med. 23:315-320) reported

survival of an adult 2 Neotoma a. albigula after 14 feedings by a total of 1016

Triatoma during 168 days of laboratory confinement. The maximum number of

bugs fed in one day was 148. Experiments reported here using a larger subspecies

of Triatoma protracta indicate larger volumes of blood consumed in toto and for

all stadia.

A wild 2 Neotoma fuscipes captured 1 August 1970 in a horse barn in Juniper

Hills, northern Los Angeles County, California, was transferred to the Los Angeles

City College laboratory. It was fed water and rat chow with carrots, lettuce and

Bermuda grass supplied at irregular intervals. Except for removal to hardware

cloth cylinders for bug feedings, the wood rat was confined in a cage 28 X 21 X 22

cm resting on SAN-I-CEL bedding in an open cardboard box until released

alive 13 August 1971 near the original site of capture. During this time 1789

Triatoma protracta navajoensis (513 $, 644 2 and 612 nymphs) fed on this

wood rat at 39 different times at approximately weekly intervals. The largest

number of bugs fed at one time was 201 on 9 September 1970, the smallest 13

on 12 January 1971, the average was 49.5 per feeding. Adult bugs averaged 30.1

per feeding, nymphs 15.6 at any one time. The 232.5 g weight of the wood rat

before first feeding on 3 August 1970 did not indicate an abnormal size but

on 17 August it weighed 248.5 g and on 24 August 269 g. One young born on

30 August was found dead. Weights before and after the 36 bug feedings after

birth of the young averaged 218.5 g (range 209.5 to 230) before and 214 (207-

206

The Pan-Pacific Entomologist

221.5) after feeding revealing an average weight loss of 4.5 g per feeding. Greatest

weight loss from bug feedings was 12.5 g 11 January 1971, the smallest was 0.5

g 16 July 1971. Although most of the weight loss must have been from the obvious

loss of blood, some would be due to natural metabolic processes.

I have used an engorgement formula of 5 quantities as follows: fully engorged,

%, %, % and %. The formulae for the 1789 bugs fed were as follows:

1177 adults (790: full, 133:%, 128:%., 61:% and 65:%) ; 349-5th instars (54-60-72-

102-61) ; 151-4th instars (28-19-37-41-26) ; 74-3rd instars (6-7-11-33-17) ; 27-2nd in-

stars (7-3-5-8-4) and 11-lst instars (8-0-3-0-0). Triatoma classified as full or %

were successful in molting so as noted above 114-5th, 47-4th, 13-3rd, 10-2nd and

8-lst instars molted in the laboratory. The 923 engorged adults produced numerous

eggs after feeding.

Xenodiagnosis for Chagas’ trypanosome was negative in 65 Triatoma (3 $, 5 2,

54-5th and 3-4th instars) examined in August 1970.

Laboratory temperatures at the time of 27 feedings averaged 23 °C, the range

being 19° to 29 °C. Exposure times for Neotoma for 36 feedings varied from 60

to 390 min, the average being 168. Degree of irritability of the rat in the hard-

ware cloth feeding cylinders may be indicated by the number of fecal pellets

released during feeding. Extremely nervous rats may defecate more frequently.

This 2 Neotoma was remarkable in releasing only 6 pellets during the 39 day-

time feeding exposures.

One 85.5 g $ Neotoma fuscipes was killed 13 June 1968 by exposure to the

feeding of 14 $ and 40 2 T. p. navajoensis. Weight loss was 6.5 g, exposure

time was 240 min and there were no fecal pellets in the feeding chamber. Linsdale

and Tevis (1951, The Dusky-footed Wood Rat, U. C. Press, Berkeley) noted

that wood rats being handled released fecal pellets as a sign of nervousness. They

reported that pellets were released at night when the wood rats were active and

one female studied for 24 days averaged 124 pellets per 24 hour period (range

7 to 208) .

One 94 g $ Neotoma fuscipes was killed 13 June 1968 by the feeding of 33 $

and 19 2 T. p. navajoensis and 10 <5,2 2 and l-5th instar T. rubida uhleri. Weight

loss was 8.5 g, exposure time 150 min and there were 27 fecal pellets released.

Wood (1947, Bull. So. Calif. Acad. Sci. 46:144-155) reported survival of albino

Rattus norvegicus to feedings of 130 Triatoma over 7 hours, 168 over 3.4 days

and 317 over 59 days. Recently, one 305 g $ CFN (Carworth, Inc.) albino

Rattus norvegicus survived the feedings of 4 $ , 97-5th, 84-4th, 12-3rd and 1-lst

instar nymphs of T. p. navajoensis on 13 August 1969 and 38 $ , 62 2 and 19-5th

instar nymphs of T. p. navajoensis and 2 $ and 5 2 T. rubida uhleri on 18

August 1969. Weight losses were 38 and 27 g, exposure times 510 and 360 min

and pellet counts were 12 and 34, respectively. Another 358 g S CFN rat sur-

vived the feeding of 64 $ and 41 2 T. p. navajoensis on 11 September 1969.

Weight loss was 15.5 g, exposure time 180 min, and pellet count 15.

The capability of wood rats in providing blood meals for numerous adult and

5th instar Triatoma is important in replenishing stocks of bugs in nature. Recent

experiments with Peromyscus also verify a supporting role of these mice as hosts

for early instars of Triatoma (unpublished data).

I thank Jack C. von Bloeker, Jr., for specific identification of the adult wood

rat. — Sherwin F. Wood, 614 W. Shenandoah St., Thousand Oaks, California

91360.

Vol. 50, No. 2, April 1974

207

Ovipositional Behavior of Chloealtis aspasma (Orthoptera: Acrididae).

— At the time of its description the ovipositional behavior of Chloealtis aspasma

Rhen and Hebard was unknown. The structure of the ovipositor suggests that

this species might oviposit in wood, a phenomenon common to some other species

in the genus (Jago, 1969, Proc. Acad. Nat. Sci. Philadelphia, 121: 229-335).

The ovipositional behavior of C. aspasma was studied 16 miles northeast of Ash-

land, Jackson County, Oregon. The study site is an old logged-over clear-cut area

surrounded by mixed conifer forest. In recent years selective cutting has continued

immediately adjacent to the clearing. Consequently the area is littered with slash,

logs and stumps, all in varying stages of decay. Vegetation in the clearing consists

of grasses, forbs and one perennial shrub, elderberry, Samhucus caerulea Raf.

On July 19, 1971 a number of elderberry stakes were driven into the ground to

serve as nesting sites for solitary bees. When the stakes were examined in April,

1972 the presence of grasshopper eggs in the center pith was noted.

On July 6, 1972 a large number of elderberry stakes were placed in the study

area and monitored periodically in an effort to identify and study the grasshopper.

The first female C. aspasma visited the stakes and oviposited on July 18. Ovi-

position continued throughout the study period but declined in frequency before

the study was terminated on August 11. Each female spent 22-46 minutes in-

serting her ovipositor and depositing a single pod of eight eggs. No female was

observed to deposit successive egg-pods.

Periodic surveys were conducted to determine the distribution of C. aspasma

within the area and to identify oviposition sites. Females and nymphs were found

throughout the clear-cut area. They were less frequently encountered in the brushy

interface between the clearing and forest. Males were apparently restricted to the

brushy interface area. They were not found in either the cleared or forested area.

Potential ovipositional sites in the area were examined. These included all

available types of dead wood, the stems of dead herbaceous plants and elderberry

stems. Females were attracted only to the exposed pith of the numerous dead and

broken stems around the base of each elderberry plant. Examination of old stems

revealed ovipositional scars from previous years. No such marks were evident on

the other materials examined. Undisturbed habitats in the immediate vicinity were

also surveyed. Elderberry was the only ovipositional substrate utilized in un-

disturbed areas.

Earlier literature (Gurney, Strohecker and Heifer, 1964, Trans. Amer. Entomol.

Soc., 84: 119-137) suggests that large series of C. aspasma are not common. How-

ever, the population in the study area was large. The numerous females and nymphs

were conspicuous and easily collected. It is plausible that clearcutting and the

subsequent large-scale invasion of elderberry has enhanced the population’s de-

velopment. In adjacent, undisturbed habitats elderberry plants are widely scat-

tered and restricted to open, south-facing slopes. Grasshopper populations in un-

disturbed areas are much smaller than in the study area.

Stakes used in the study were allowed to overwinter in the field. They were re-

turned to the laboratory in late April, 1973. Examination of and rearings from this

material revealed that two insects commonly prey upon the eggs of C. aspasma.

The larva of a chloropid fly, Goniopsita oophaga Sabrosky, destroyed an average

of 67.1 percent of the eggs examined while the larva of a clerid beetle, Trichodes

oregonensis Barr, destroyed 5.3 percent.

I wish to thank Drs. Ashland B. Gurney and Curtis W. Sabrosky, both of the

208

The Pan-Pacific Entomologist

National Museum of Natural History, Washington, D. C., for their respective iden-

tifications of C. aspasma and G. oophaga, and Dr. W. F. Barr, University of Idaho,

who helped with certain aspects of the field work. — David E. Foster, Entomology

Section, College of Agriculture, Texas Tech University, Lubbock, 79409.

ENTOMOLOGICAL EXCHANGES AND SALES

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Advertising Manager, J. A. Chemsak, Division of Entomology and Parasitology,

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FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca,

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EXCHANGE. Eastern U.S. and exotic Scarabaeidae, Cerambycidae, or Cicindeli-

dae for western U.S. Cicindelidae. John D. Glaser, 6660 Loch Hill Rd., Baltimore,

Md. 21239.

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Catalogues on request. State your interests (Lepidoptera, Coleoptera, new/ second

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Beginning January, 1975 (Vol. 51, No. 1), The Pan-Pacific Entomologist will no

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Vol. 50, No. 2, April 1974

209

Ralph Ward Dawson. Dr. Dawson was born in Lincoln, Nebraska, May 1,

1887, and died from a heart attack at Folsom, California, March 1, 1974. He at-

tended public schools at Lincoln and obtained a B. S. degree at the University of

Nebraska in 1912. He was instructor and assistant professor respectively in ento-

mology at Nebraska until 1922, when he attended graduate school at the University

of Minnesota, obtaining his doctorate in 1929. Within a year he was asked to

join the Department of Zoology at Minnesota, where he remained until his re-

tirement. He then lived in Colorado and Arizona and in 1959 moved to Pullman,

Washington, where he accepted an appointment as Collaborator in Entomology

at Washington State University. In June of 1973 he joined a retirement com-

munity at Folsom. He never married and is survived by a niece in Cleveland.

Dawson’s tenure of almost 14 years as Collaborator at WSU was of tremendous

benefit to our Department. In his honorary appointment without pay, he worked

regularly running Malaise traps, sorting and mounting material and making de-

terminations of groups within his own specialty. His acquisitions to our insect

collection were substantial. In addition, he prepared considerable class material,

occasionally assisted in the laboratory and gave a few lectures. Unfortunately,

during his last few years he experienced declining vision which reduced much of

this activity, but his mental acuity remained excellent until the end.

Professor Dawson was one of the vanishing breed of old time naturalists. He

had a most profound knowledge of natural history but especially insects and

birds. He was also an accomplished nature photographer, a gifted pianist and

a weaver of Navaho rugs. At the University of Minnesota where he had charge

of huge beginning classes in Zoology, he had the reputation of being the best

lecturer on the entire Minnesota campus.

Ralph was reserved and on the quiet side but a most interesting and stimu-

lating conversationalist when discussing his principal interests — birds and insects.

He maintained a wide correspondence with his former students and materially

aided those in financial need. He made lasting friendships and was well liked.

His taxonomic interests were quite broad. He published a number of papers

210

The Pan-Pacific Entomologist

on the genus Serica (Scarabaeidae) . His extensive personal collection of these

beetles was donated to the California Academy of Sciences a number of years ago.

He also maintained an active interest in the Syrphidae, bumblebees, butterflies,

meloids and other groups. H. S. Telford, Washington State University, Pullman,

WA 99163.

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