Vol. 52 = JANUARY 1976 No. 1

POWELL—Host plant preference, mating and egg development in Synnoma

inosyraninlepideprena: Lontricidae): 20 578 tet. ee 1

MERRITT—A review of the food habits of the insect fauna inhabiting cattle

droppines mmaNorth'.CentralCalifomia, $0. 4 a oS eee 13

SCOTT AND SHEPARD—Simple and computerized discriminant functions

for difficult identifications: a rapid nonparametric method 23

AULT—Observations on the nesting behavior of Belomicrus columbianus Kohl

CHipimeno prerd » \OMUeCIdae ip ements e 29

PURCELL—Seasonal changes in host plant preference of the blue-green sharp-

shooter Hordnia circellata (Homoptera: Cicadellidae) 33

CHEMSAK AND LINSLEY—Further studies among the Mexican and Central

American lepturine Cerambycidae related to Euryptera and Choriolaus

(Coleopperayies 2.5. AUk. Sane) gr Te ieee Ok ae 2 Pik pee eee 38

JOHNSON—Redescription and phylogenetic affinities of Kytorhinus prolixus

(Fall) (Coleoptera: Bruchidae: Kytorhininae) —_ 50

MOORE—Giulianium campbelli, a new genus and species of marine beetle from

Galitomida (Coleoptera: Staphylinidae)! 2. siek. 22 A 56

JOHNSON—A new species of Callophrys (Mitoura) from Mexico (Lepidop-

tericn bycareniGde) eae. LAr BS Sieh 60

RISSING AND WHEELER—Foraging responses of Veromessor pergandei to

changes in seed production (Hymenoptera: Formicidae) — 63

PARKER—A new Proteriades reared from trap stems, its biology and nest

associates (Hymenoptera: Megachilidae) _.. 73

MEIN MOTO SIN COINS rel a baie I 2 Say 3U80) 82.895

BOLUS UM EC IR IC, RES, AMIS Dalccste een” DER OES. 7 OE: Ah RR SS 62

ee ORNL LORE AED, coke ES 3. Na eR i Fu 22, 229409

ZOOLOGICAL NOMENCLAT WRE 2. Sab 2 go aT ' nt ee 28

SAN FRANCISCO, CALIFORNIA ¢ 1976

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

Vor. 52 January 1976 No. 1

Host Plant Preference, Mating and Egg Development in

Synnoma lynosyrana

‘(Lepidoptera: Tortricidae)

JerRY A. POWELL

University of California, Berkeley

In several ways Synnoma lynosyrana Walsingham is unique among

American Tortricinae. Like many Sparganothidini, this species exhibits

marked sexual dimorphism, but in contrast to all other members of the

tribe, the female of Synnoma evidently is flightless. Males possess

strongly pectinate antennae, whitish ochreous forewings flecked with

shining bluish silver, and dark brown hindwings, while females have

filiform antennae, an immensely enlarged abdomen, lack most of the

ochreous scaling on the shining bluish black forewings, and have dull

gray-brown hindwings. Elongate vestiture on the body and appendages

and strongly upraised scaling on the forewings, gives both sexes a

ruffled, shaggy appearance (figs. 1-3).

Synnoma lynosyrana was discovered by Lord Walsingham near Mt.

Shasta in northern California in 1871, when he reared adults from

larvae on Lynosyris (Compositae) (Walsingham, 1879), a shrub now

known as Chrysothamnus. Other than a brief report nearly a century

later (Powell, 1959) and a description of the larva by MacKay (1962),

nothing has been recorded on the biology or distribution of this interest-

ing insect. We now know the species to be widespread in Great Basin

and Mojave Desert regions, from southern Montana, Idaho and Wyoming

through eastern Oregon, northeastern California, Nevada and Arizona

to extreme western Texas, with an apparently isolated population at the

western edge of the Mojave in southern California (map 1). Records

are lacking from eastern Washington, Colorado, and Utah, although

the species no doubt ranges into some of these areas.

A generalized life history is as follows. There is a single annual

generation, adults emerging in the fall, from October in the north to

November in the south. The moths rarely have been observed in the

field, in part owing to their late seasonal occurrence and in part to

Tue Pan-Paciric Entomo.ocist 52: 1-12. January 1976

2 THE Pan-PaciFic ENTOMOLOGIST

Fics. 1-4, Synnoma lynosyrana Walsingham (JAP 73H8-9): 1, 2, female in

characteristic “calling” posture; 3, female perched in foliage, a cryptic position

taken at night or after external disturbance; 4, mating pair.

their diurnal behavior, which precludes light attraction records. The

eges are deposited in elongate clusters on the twigs of certain woody

Compositae and enter diapause or a prolonged temperature related

quiescence, probably until early spring. Young larvae evidently migrate

down the stems and feed on newly developing foliage. Later instar

larvae live in aggregated clusters of tough silken tubes, tying up the

branches into conspicuous bundles. They emerge from these shelters

Vou. 52, No. 1, JANUARY 1976

zy

a

30

| -@& chrysoruannus

@ GUTIERREZIA

| -C>- NO HOST ASSOCIATION

SCALE

200 300 MILES

. i ,

KILOMETERS

fo)

n

T

200

CONIC PROJECTION

Distribution of locality records for Synnoma lynosyrana Walsingham

i a

120

in the western United States. Top half closed circles indicate larval collections

from Chrysothamnus, bottom half closed circles indicate records from Gutierrezia.

Map. 1.

Open circles indicate collections of adults without host plant information.

to forage on leaves and green bark. Pupation occurs in the larval

galleries in the fall.

Host plant selection.—Although Synnoma lynosyrana has been reared

from various composites of the genera Chrysothamnus, Gutierrezia, and

Haplopappus, these plants are not used randomly over the whole distri-

4, THE Pan-PactFic ENTOMOLOGIST

bution of the moth. A general pattern emerging from field work and

museum specimen records indicates that the species is dependent on

Chrysothamnus over the northern half of its range, and in the Great

Basin, and primarily on Gutierrezia in southern parts of its distribution

(map 1). In northern California and northeastern Nevada I found

larvae on both Chrysothamnus nauseosus and C. viscidiflorus (58H2-4,

73H8-9),1 while records from southern Idaho, Montana (MacKay,

1962) and Wyoming (67J6) involve only the latter plant. Southward,

the larval shelters have been collected on Gutierrezia at the northern

edge of the Mojave in south central Nevada (68633), at five localities

in Arizona and northern New Mexico (68F19, 68F40, museum speci-

mens), and in the Big Bend region of Texas (MacKay, 1962). Most

collections from Gutierrezia have been made too early in the season to

enable specific identification of the host plant. A specimen label from

Springer, N.M., gives G. sarothrae, a widespread species, as a host.

Probably various species of the genus are used.

This pattern is supported by negative evidence accumulated during

an extensive survey of these plant genera conducted during 1966-1968.

Chrysothamnus and Haplopappus were examined during late July to

October at numerous stations in Nevada, eastern and southern California,

and Arizona without disclosing evidence of Synnoma. Thus, although

the distributions of the latter two plant genera are broadly sympatric

with that of S. lynosyrana over wide areas, these hosts are not utilized

in the southern, more xeric regions. Exceptions that may represent

incidental or temporary associations occur at the western edge of the

Mojave in southern California. Near Frazier Park a sparse colony of

Synnoma was discovered on Chrysothamnus nauseosus in 1961 (61H2).

However, we were unable to relocate larvae at this site in 1967 and

1973. Approximately 40 km to the east in the Cuyama Valley we found

an extensive population on Gutierrezia in October, 1967 (67K94). Here

C. nauseosus growing abundantly in the riverbed was not used by the

moth, but two plants of nauseosus among the Gutierrezia on slightly

higher ground harbored many Synnoma larvae (67K95). Haplopappus

linearifolius was also utilized in an isolated instance in the Cuyama

Valley. We found no larvae on this plant in 1967, but in October, 1972,

with shelters again common on Gutierrezia, an extensive survey of all

three host plants disclosed one bush of H. linearifolius with several larvae

(72K1). This species is distributed primarily in the Coast Ranges

in somewhat more mesic habitats than Synnoma, and at Cuyama Valley

loses most of its foliage by the end of the dry season. Because of its

1 Data for collections accompanying these rearing numbers is given in the appendix.

VoL. 52, No. 1, JAnuary 1976 3

deciduous habit it seems doubtful that H. linearifolius could support an

extensive colony of larvae, and here Chrysothamnus and Haplopappus

appear to be incidental hosts used by a population of Synnoma which

is essentially dependent on Gutierrezia. Other species of Haplopappus,

particularly H. venetus, H. acradenius and H. tenuisectus (= hartwegii) ,

were examined many times within the range of S. lynosyrana with nega-

tive results.

Diel rhythm and longevity of adults Observed emergences from the

pupa were between 0715 and 0906 PST (67J6, n= 10), and at least

some took place earlier in the morning. Wing expansion and drying

in four timed examples required 16—20.5 minutes (8-9.5 minutes from

breaking of the pupal shell until the wings were fully lengthened, then

7-11 minutes with the wings held erect from the body before being folded

into the normal position over the dorsum).

In contrast to nearly all other Nearctic Tortricinae, moths of the

present species are diurnal. We observed males flying on a clear day in

early November at 1030-1130 PST, near Gorman, California (7412),

despite a stiff, cool breeze. During intermittent calm periods and in

sheltered spots the moths actively fluttered above Chrysothamnus

foliage, apparently independent of observer disturbance. Repeated sur-

veillance of confined individuals made during darkness hours in presence

or absence of artificial lighting revealed virtually no movement of

Synnoma adults (67J6, 67K94.). Occasionally females continued oviposi-

tion in darkness after initiating egg-laying late in the day. Activity in

the laboratory was greatest between 1230 and dusk, ceasing about 30

minutes after sundown (ca. 1830 in September, to 1730 in October)

(67K94, 73H8-9). Occasional movement by both sexes occurred be-

tween 0730-1200, but most individuals remained inactive during the

morning. Some females assumed the “calling” position perched atop

an inflorescence (figs. 1, 2) early in the morning and if not mated

sometimes remained so perched until well after dark. On cloudy days

moths housed in a trailer lab (18°C) remained inactive.

During warmer hours of the day males displayed a positive photo-

tactic reaction, flying from open containers to windows, fluttering at

sides of containers towards sunlight, etc. Although females sometimes

dropped from open bouquets of the plants, they were not attracted to

sunlit windows. Any movement or jostling of the plant usually caused

females at calling perches to quickly back down the stem into foliage,

but none was ever observed to attempt flight. Following mating and

oviposition, several females died at the bottom of containers, even

though there was no closure confining them. Under field conditions, it

6 THE Pan-Paciric ENTOMOLOGIST

is possible that females fly after a portion of the egg complement

has been deposited, but my observations do not suggest this.

Adults were comparatively short-lived, surviving only 5-9 days in

standardized one-gallon breeding jars (Powell, 1964). None was ob-

served to visit the water source that was provided, and dissections re-

vealed the tongue to be extremely reduced and apparently non-functional

in both sexes, with the galea lobes separated and only about 0.3 mm

in length. Thus it appears that Synnoma lynosyrana is adapted for sur-

vival without nourishment as are late fall and winter-flying ethmiid moths

of arid regions in the western Nearctic (Powell, 1971, 1973). Female

Synnoma housed without cage enclosures and without a water source

produced fertile egg masses of apparently normal size, but individual

longevity records were not maintained (73H8-9).

Courtship and mating.—Although viable eggs were obtained from

caged adults, sexual behavior of the moths was successfully observed

only in 1-3 day old individuals placed outside under conditions of

natural air movement (73H8-9).

Pairs were tested at Berkeley in September, between 1500-1730 in a

light westerly wind. Under field conditions mating behavior is prob-

ably related to temperature and may also occur earlier in the day. As

soon as containers with dry Chrysothamus inflorescences and moths

were placed outside in a light breeze, males commenced a characteristic

search behavior in which they quickly crawled over the foliage, rapidly

fluttering their wings. This activity is unlike any other tortricid move-

ments I have seen. Males which took flight sometimes remained aloft

for several minutes, generally hovering up and down 1-3 m above

the ground (females were stationed at a height of 1.5 m). If a male

flew upwind from the female station no reaction was obvious until he

drifted downwind to the east of the female. Downwind, males were ob-

served to detect the pheromone plume, from as far away as 10 m, as

evidenced by a sudden change in flight pattern from hovering in vertical

arcs to a flight directly towards the female perch, wavering only 10—20

cm from side to side. Arriving at the Chrysothamnus, males immediately

initiated the fluttering behavior and rapidly crawled from one in-

florescence to another if the branch was not occupied by the female.

This search behavior sometimes required up to 60 seconds including

three or four pauses of a few seconds duration each.

Once a female was encountered, the courtship sequence was brief.

The male quickly crawled around the female, oriented head to head,

with his abdomen curled towards her and his valvae widely spread.

When genital contact occurred, coupling was immediate, and the

~]

VoL. 52, No. 1, January 1976

Fics. 5, 6, Synnoma Presale (Walsingham) (JAP 73H8-9): 5, mating pair;

6, female perched above egg cluster.

male turned to assume the tail-to-tail position that is characteristic of

all tortricids in copulo (figs. 4-5). No reaction by the female was evi-

dent during courtship. Her wings were held in repose, with the large

abdomen protruding beyond. The whole courtship sequence required

2-10 seconds in several timed examples. Sometimes copulation oc-

curred almost immediately, but in about half the encounters the male

approached head first to the tip of the female’s abdomen at the onset

of contact before orienting in a head-to-head position alongside the

female.

Several attempts were made to elicit courtship response on exposed

Chrysothamnus bouquets in a trailer lab (73H9). Three successive first-

day males were released near a newly emerged female perched in calling

position at 1645-1700 with indirect lighting from late afternoon sun.

Each began the courtship fluttering behavior immediately, moving up

and down in the foliage but mostly well below the female. Brief periods

of pause were interspersed with activity, but the female was not closely

approached. Each male stopped altogether after 1.5-2 minutes and did

not move again for 10 minutes. Lightly blowing across the female

towards the inactive male resulted in some further brief activity but no

closer approach to the female. Apparently, natural air movement, and

8 THE Pan-PaciFric ENTOMOLOGIST

possibly direct sunlight, is necessary for effective tracking of the scent

plume.

Females remained in position, head-upward on the plant terminal dur-

ing courtship and mating, sometimes with the male arranged awkwardly

sideways due to interposing foliage. The duration of copulation was

short compared to other Tortricinae (Powell, 1964:27), lasting 12-19

minutes (avg. 17 min., n = 4) in examples where I witnessed both court-

ship and separation.

Following unsuccessful mating attempts, consisting of coupling for a

few seconds to a few minutes, females were immediately attractive to

the same or another male. On one occasion, a male was attracted in

flight from a distance of about 1 m to a mating pair that had coupled

2-3 minutes previously. At termination of successful mating, the male

flew off while the female either remained stationary or moved a few

cm away from the calling location. In one instance the male was ob-

served to fly around 2-3 minutes, finally coming to rest about 5 m distant

and was not attracted by other females placed at the calling station

during the succeeding 40 minutes. Whether mated females continue

broadcasting the pheromone and are capable of mating again during

the same diel cycle was not ascertained. Dissections of females that

had successfully mated and deposited eggs revealed a single spermato-

phore in each (67K94, n=6). Thus, under laboratory conditions fe-

males evidently mate only once, but individually labelled females were

not tested under outdoor conditions on successive days. The spermato-

phores are amorphous and irregular in size compared to those of other

Tortricidae, so that confirmation of their number is difficult (and is

further complicated by the sticky, black accessory fluid which fills the

corpus bursae). Thus further studies are needed to confirm that fe-

males mate but once.

Oviposition.—The eggs are normally deposited in elongate patches

on the twigs of the host plant, usually about 4—5 cm below the flower

heads (figs. 6-8). Occasional masses are placed near the apex of broken

stems or on lower stems. Each egg mass consists of a double row of

regularly arranged, imbricate eggs. The size of these clusters varies

considerably and in the laboratory may be affected by such factors as

density of moths, observer disturbance, etc. One female in a breeding

jar also housing two males, laid individual patches containing 8—46

eggs (67J6, avg. 29, n= 6), while those deposited by several females

on exposed bouquets of Chrysothamnus were larger, ranging 70-116

(73H8-9, avg. 92, n= 6). Sometimes irregular masses were deposited

VoL. 52, No. 1, January 1976 9

Fic. 7, egg cluster of Synnoma lynosyrana Walsingham deposited in the lab

on dry Chrysothamnus foliage (JAP 67K93.1); fig. 8, closeup of a portion of the

same cluster.

on paper toweling at the bottom of the containers, possibly by older,

weakened females.

Oviposition was observed only twice (67J6, 73H8), at 1730 and 1930

(before sundown and well after dark) in September. Several other

occurrences of egg deposition were recorded between these hours, and

a few later at night (at 18-20°C). In one instance a female under

surveillance placed approximately 26 eggs during 10 minutes in a lighted

room (67J6). Each egg-laying was brief but was followed by a long

period (15-40 seconds) in which the female reconnoitered the position

of the next egg by tapping the surface of the prior two or three eggs

and adjacent stem with the papillae anales.

The precise number and time interval between eggs were difficult to

observe because of a black substance secreted around the eggs that acts

as an adhesive and perhaps as a protective agent. The source of this

material is a pair of huge accessory glands lying alongside the ductus

10 THE Pan-Pactric ENTOMOLOGIST

bursae and filling the tip of the abdomen in virgin females. The liquid

is secreted with each egg, rather than over the whole patch at its com-

pletion, as was evidenced when a female was interrupted after depositing

only a short row which included the normal amount of accessory fluid.

Other Sparganothidini cover the eggs with milky or clear, cement-like

secretions (Powell, 1964:31), and no other American tortricine moths

are known to produce a black substance for the eggs.

The total number of eggs deposited by a female ranges from 230-280.

For example, a single female isolated in a breeding jar deposited 233

eges after the fourth day following emergence (67J6). Owing to the

short lifespan of the adults and the fact that the mouthparts are rudi-

mentary, it might be expected that the eggs are deposited within a few

days with little successive egg maturation involved. Dissection of virgin

females indicated this to be the case. Examination showed 180 full sized

or nearly full sized eggs and about 80 eggs of varying decreasing sizes

in a female on the day of emergence (67K94.), while an unmated female

that lived several days contained about 275 full sized eggs and only a

few smaller ones (73H9). In females that were retained alive for one

or more days following mating, dissections showed no indication of

continued egg production, with 20-220 full sized eggs remaining follow-

ing oviposition (67K94).

Egg development.—When eggs were housed in the laboratory, (20° +

2°C), larvae began to emerge in 36-40 days (58H3-4, 67J6, 73H8-9).

This is a considerably longer period than is normal in Tortricinae, yet

shorter than in the usual obligate diapause (Powell, 1964:34). The

emergence pattern is similar to that of eggs of the winter-active Ethmia

charybdis Powell (Powell, 1971). Experiments with Synnoma indicated

that incubation can be delayed by reduced temperatures. Some of the

73H8-9 eggs were transferred to an outdoor cage at Berkeley after 30

days in the laboratory, and emergence was delayed to 43-47 days. Other

egg masses of this lot were moved to an outdoor shed at Russell Reserve,

an inland station at about 250 m elevation, after 18 days in the labora-

tory. No larval eclosion occurred there, with weekly averages of daily

mean temperatures, in successive weeks: 13, 9.5, 9, 9, 8, 3, 4, 4, 3.5, 4,

6, 0.5, 0.5, 7.5. After 70 days one egg cluster was transferred to the lab

in Berkeley, where larvae emerged 20-24 days later. The final clusters

were removed from the shed on January 20, 1974 after 100 days, and

larval hatching occurred 16-17 days later. Under these conditions, the

percent successful eclosion declined with longer cold storage. Thus it

seems likely that under field conditions, egg maturation is delayed by

the onset of cold temperatures after oviposition in late October to early

VoL. 52, No. 1, January 1976 ll

December, so that larval hatch does not occur through winter but com-

mences with warming conditions in early spring, correlated with vernal

foliation of the foodplants.

Acknowledgments.—This study was in part supported by National

Science Foundation Grant No. GB-6813x, and assistance with field

collections and laboratory observations during 1967 was provided by

P. A. Opler and P. A. Rude while working on that grant project. R. A.

Mackie, now with the Public Health Service, Guam, sent specimens and

host data from southern Idaho in 1956. Cooperation by the following

individuals enabled use of specimens in institutional collections: K. J.

Goeden and R. L. Westcott, (Oregon Department of Agriculture, Salem) ;

R. W. Hodges (National Museum of Natural History, Washington,

D.C.) ; L. M. Martin and J. P. Donahue (Los Angeles County Museum

of Natural History); F. H. Rindge (American Museum of Natural

History, New York); and the late T. N. Freeman and J. E. H. Martin

(Canadian National Collection, Ottawa). Some of the data from museum

specimens were assembled by the late R. L. Lambert of Ottawa in his

manuscript on American Sparganothidini. J. T. Doyen read a draft of

the manuscript and offered useful suggestions. The photographs for

figures 7 and 8 were produced by A. A. Blaker, formerly of the Scientific

Photo Lab, University of California, Berkeley.

LITERATURE CITED

MacKay, M. R. 1962. Larvae of the North American Tortricinae (Lepidoptera:

Tortricidae). Canad. Entomol., Suppl. 28; 182 pp.

Powe tt, J. A. 1959. Note on Synnoma lynosyrana in: Proc. Pacific Coast Ent.

. Soc., 262nd meeting. Pan-Pac. Entomol. 35:62.

PoweLL, J. A. 1964. Biological and taxonomic studies on tortricine moths, with

reference to the species in California. Univ. Calif. Publ. Entomol., 32;

317 pp.

Powe.., J. A. 1971. Biological studies on moths of the genus Ethmia in Cali-

fornia (Gelechioidea). J. Lepid. Soc., 25, Suppl. 3; 67 pp.

PoweLt, J. A. 1973. A systematic monograph of New World ethmiid moths

(Lepidoptera: Gelechioidea). Smithson. Contr. Zool., 120; 302 pp.

WarsincHamM, Lorp T. pe Grey 1879. Illustrations of typical Lepidoptera

Heterocera in the collection of the British Museum. Part IV, North

American Tortricidae. London, British Museum (Nat. Hist.), 84 pp. +

17 pls.

APPENDIX

Biological Study Collections—JAP 58H2: 1.5 mi NE Weed, Siskiyou Co.,

Calif. VIII-7-58 larvae on Chrysothamnus viscidiflorus typicus. JAP 58H3-H4:

10 mi NE Weed, Siskiyou Co., Calif. VIII-23-58 larvae on C. viscidiflorus and

C. nauseosus albicauis. JAP 61H2: 2 mi W Frazier Park, Kern Co., Calif. VIII-

12 THe Pan-Paciric ENTOMOLOGIST

24-61, larvae on C. nauseosus mojavensis. JAP 67J6: Point of Rocks, Sweetwater

Co., Wyo. IX-4-67, larvae on C. viscidiflorus viscidiflorus. JAP 67K94-K95:

Cuyama Vy., 3 mi S Ozena Jct., Ventura Co., Calif. X-6-67, larvae on Gutierrezia

californica and Chrysothamnus nauseosus mojavensis. JAP 68F19: 3 mi SE

Kingman, Mohave Co., Ariz. VI-3-68, abandoned shelters on Gutierrezia. JAP

68F40: 4 mi S Pine, Gila Co., Ariz. VI-5-68, abandoned shelters and young larvae

on Gutierrezia. JAP 68633: 10 mi SW Clark’s Sta., Nye Co., Nev. VII-20-68,

Larvae on Gutierrezia. JAP 68G67: 3 mi E Litchfield, Lassen Co., Calif. VII-

24-68, larvae on Chrysothamnus nauseosus consimilis. JAP 72K1: Cuyma Vy., 6

mi N Ozena Jct., Ventura Co., Calif. X-5-72, larvae on Haplopappus linearifolius.

JAP 73H8-H9: 2 mi W Wells, Elko Co., Nev. VIII-14-73, larvae on Chrysotham-

nus nauseosus and C. viscidiflorus. JAP 74L2: 8 mi ESE Gorman, Los Angeles

Co., Calif. XI-5-74, adults flying in assoc. Chrysothamnus nauseosus mojavensis.

RECENT LITERATURE

FRAGILE EcosysTeMs: EVALUATION OF RESEARCH AND APPLICATIONS IN THE

Neotropics. E. G. Farnworth and F. B. Golley (Eds.). The Institute of

Ecology. Springer-Verlag, Berlin, etc. 1974. xxvii + 256 pp. $7.80, paper.

TROPICAL EcoLocicAL Systems (EcoLocicaL Stupies, VoLuME II). F. B. Golley

and E. Medina (Eds.). The Institute of Ecology. Springer-Verlag, Berlin, etc.

xvi + 398 pp. 1974. $24.80, paper.

ComMPLEX ADAPTATIONS IN Evotvinc PopuLations. T. H. Frazzetta. Sinauer

Associates, Sunderland, Massachusetts 01375. 288 pp., 51 illustrations. 1975.

$4.95, paper; $11.00, cloth.

AnrmMAL BeHaAvyiorn: AN EvoLutrionary ApproaAcH. J. Alcock. Sinauer Associates,

Sunderland, Massachusetts 01375. 1975. 526 pp., 259 illustrations. $14.00, cloth.

CoEVOLUTION OF ANIMALS AND Piants. L. E. Gilbert and P. H. Raven (Eds.).

University of Texas Press, Austin 78712. 1975. xiv + 246 pp. $12.50.

History oF ENTOMOLOGY IN THE PENNSYLVANIA DEPARTMENT OF AGRICULTURE.

A. G. Wheeler and K. Valley. Pennsylvania Department of Agriculture, Bureau

of Plant Industry, Harrisburg, Pennsylvania 17120. iii + 37 pp., available on

request.

This useful descriptive work summarizes the history, present operations and

biological holdings of the Pennsylvania Department of Agriculture. The present

collection contains approximately 92,000 pinned insects and over 5000 slides and

vials of immatures or soft-bodied adults. Especially well represented are Coleoptera,

with 42,400 identified specimens.—Editor.

A Review of the Food Habits of the Insect Fauna Inhabiting

Cattle Droppings in North Central California’

Ricuarp W. MERRITT?

Division of Entomology and Parasitology

University of California

Berkeley, California 94720

A two-year study (1971-73) in the Sierra Nevada foothills of Cali-

fornia consisted of: 1) a quantitative analysis of the differences in di-

versity and abundance of the insect fauna colonizing and inhabiting

undisturbed cattle droppings in four different pasture and rangeland

ecosystems; and 2) a study of the relationship between the diversity

and abundance of insect inhabitants per cowpat and the rate of pat

degradation (Merritt 1974, Merritt and Anderson, in prep.). During

this time observations were made on the feeding habits of selected dung

insects and a general literature survey was conducted. This report re-

views the known food habits of the insects recorded in the previous

study. Many of the families, genera and species considered here, also

occur in bovine manure throughout North America and other parts of

the world. Table 1 is a general tabulation of the food habits of dung-

associated insects for both adult and immature stages.

Several studies have been conducted on the biology, ecology and

succession of insects inhabiting cattle droppings (Hafez 1939, Mohr

1943, Snowball 1944, Laurence 1954, Poorbaugh 1966, Sanders and

Dobson 1966) ; however, few have dealt specifically with the food habits,

or food webs of insect communities inhabiting bovine manure. Ham-

mer (1941) studied the life histories of various species, based on field

observations, and stressed their biology, food and reproduction. Valiela

(1974) studied qualitative and quantitative changes in species com-

position and in the structure of food webs during the succession of

insects in cattle droppings. He grouped species into trophic “guilds”

or groups of species which fed on similar food and had similar feeding

behavior. He found that neither food nor predators appeared to limit

populations of dung feeders, and that local alterations in the environ-

ment and in the dung may set limits on the numbers of predators and

prey. Merritt (1974,) showed that loco- and microclimatological factors*®

1 Research on this project was conducted while the author was on an NIH Predoctoral Traineeship

in the Dept. of Entomological Sciences, University of California, Berkeley. Research was supported,

in part, by training grant 00218, and the Univ. Calif. Agricultural Experiment Station.

2 Current address: Dept. of Entomology, Michigan State University, East Lansing, Michigan 48824.

3 Lococlimate refers to the conditions prevailing in the environment close to the dung and

microclimate refers to the condition prevailing in the actual dung pile.

Tue Pan-Paciric ENTOMOLOGIST 52: 13-22. January 1976

14, THe Pan-Pactric ENTOMOLOGIST

were most important in determining the diversity and abundance of

insects colonizing dung, except during late spring when adults of

Aphodius fimetarius (L.) (Coleoptera: Scarabaeidae) invaded and

disrupted the droppings, thereby reducing the developmental sites of cer-

tain dipteran larvae.

COLEOPTERA

Hydrophilidae.—According to Sanders and Dobson (1966), and

Hafez (1939), adults and larvae of Cercyon spp. feed on dung; however,

literature on the ecology, life history, and taxonomy of this group is

scarce (McDaniel et al. 1971). I checked numerous specimens of this

group for parasitic nematodes and found that the gut contents often

consisted of a brownish-black material which I assumed was dung.

Therefore, I agree with the above authors that the adults are dung feeders

(Table 1). Feeding habits of the larvae were not observed. Larvae of

Sphaeridium spp. were observed feeding on various Diptera and Coleop-

tera immatures by Mohr (1943), Poorbaugh (1966), and Bourne and

Hays (1968). Hammer (1941) stated that the adults of Spaeridium

scarabaeoides L. were carnivorous and fed on egg heaps of Diptera;

however, more recent studies (Sanders and Dobson 1966, Thomas

1967, Kessler and Balsbaugh 1972, Macqueen 1973, Merritt, pers. obs. )

indicated that adults were generally scavengers (Table 1). Adults of

Sphaeridium spp. played an important role in the ecological succession

of animal life in the dung. They were often observed burrowing in and

out of the dung shortly after pats were dropped, thus providing aeration

to the pat and permitting staphlinids and parasitic Hymenoptera to use

their tunnels to locate prey (Gary and Wingo 1971, Merritt 1974).

Valiela (1969) showed that face fly mortality in the dung was greater

in the presence of Sphaeridium (a burrower) and a predator rather

than a predator alone. Several authors (Mohr 1943, Sanders and Dobson

1966, McDaniel et al. 1971) have discussed the ecological importance

of the tunneling activities of Sphaeridium.

Scarabaeidae.—Adults and larvae of aphodines feed on dung and

digest only dissolved albuminous substances (Madle 1934, Landin

1961) (Table 1). The larvae also digest bacterial albumens which may

account for their subsistence in old dung heaps (Landin 1961).

Staphylinidae.—Little work has been conducted on the food habits

of dung-inhabiting Staphylinidae in the United States (I. Moore, Univ.

Calif., Riverside, pers. comm.) (Table 1). Life history studies have

been conducted on a few species, such as on Aleochara tristis Graven-

horst (Drea 1966) and on Aleochara bimaculaia Gravenhorst (Wingo

TABLE 1, Food habits of dung associated insects in north central California.

Immature

A

o S é Reported Reference

Dung Pred Par Other Dung Pred Par Other Prey Citation

COLEOPTERA

Histeridae

» Peranus bimaculatus L. - x - - - xX - Diptera & Coleoptera larvae Hafez (1939)

Saprinus sp. - xX - - - x - - Diptera & Coleoptera larvae Hafez (1939)

Hydrophilidae a a

Cercyon spp. (2) xX - - - X? - - - Sanders & Dobson (1966)

Sphaeridium spp. (3) x - * - - x - - Diptera & Coleoptera larvae Thomas (1967) - Adults

ee Nar ea Mohr (1943)-Larvae

Rhizophagidae

Monotoma picipes (Herbst.) = ~ x X - - - X? Hafez (1939)

Scarabaeidae

Aphodius spp. (6) xX - - - x - - - Landin (1961)

Staphylinidae

Aleochara spp. (2+) xX xX - - - - xX - Diptera eggs, larvae, pupae Drea (1966)

Aleochara bimaculata Gray - x - = = - xX - Diptera eggs, larvae, pupae Wingo et al. (1967, 1974)

Aleochara bipustulata L. + x ™ = - at xX = Diptera eggs, larvae, pupae Thomas & Morgan (1972a)

Aleocharinae (5+) ~ X? - - - - - - Unknown I. Moore (Pers. Comm.)

Aploderus annectans LeC - X? - - - - Unknown I. Moore (Pers. Comm.)

Hyponygrus sp. x - - - - Diptera larvae & Coleoptera Valiela (1969)

adults

Leptacinus spp. (2) - X? - - - - Diptera larvae Snowball (1944)

Lithocharis ochracea Grav. - X? - - - - Unknown I. Moore (Pers. Comm.)

Oxytelus sp. xX X? ~ - - - Unknown Valiela (1974)

Philonthus spp. (4) x x - = - - Diptera eggs, larvae Valiela (1969)

Platystethus americanus Erich. X xX - -; - - Diptera larvae Mohr (1943)

Quedius sp. - X? - - - - Unknown I. Moore (Pers. Comn.)

DIPTERA

Anthomyiidae .

Calythea micropterx (Thom. ) - - = x - “a Merritt (1974)

Hylemya spp. (2) - xX x - - Mohr (1943)

Scatophaga stercoraria (L.) = x xX xX - - Diptera adults Foster (1970)

S. furcata (Say) - 4 x xX - - Diptera adults

Mohr (1943)

OL6L AUVONVE ‘T ‘ON ‘ZS “IOA

ST

Adult

Pred* par? Other

Dung

Cecidomyiidae

Colpodia sp i = = x

Tetraxyphus sp. ia me “ x

Ceratopogonidae

Forcipomyia spp. (2) - > = xX

Chironomidae

Smittia sp. _ ~ = x

Drosophilidae (1) = - = xX

Emipididae

Drapetis sp. “ x > 2

Muscidae

Pseudophaonia orichalcoides > = = x

Huckett

Haematobia irritans L. - - Xx xX

Hydrotaea spp. (2) x - - x

Morellia micans (Mac.) x - - x

Musca autumnalis (DeG.) x - - X

Myospila meditabunda (P.) x - - x

Orthellia caesarion (M.) x - - 4

Psychodidae

Psychoda spp. (3) x 7 = x

Sarcophagidae

Ravinia spp. (3) x = =

Scatopsidae (1) = i = xX

Sciaridae (2) - - - xX

Sepsidae (8) xX - - x

Sphaeroceridae (9) x ) - x

Stratiomyiidae (3) - = - xX

TABLE 1. (Cont.)

K?

X?

X?

Reported

Prey

Coleoptera adults

Vertebrate host (cattle)

Diptera larvae

Diptera larvae

A

Reference

Citation

Cole (1969)

Cole (1969)

Saunders (1924)

Strenzke (1950)

Baumberger (1919)

Laurence (1952)-Adults

Papp (1971)-Larvae

Merritt (1974)

Hammer (1941)

Hammer (1941) - Adults

Porchinskii (1911)-Larvae

Hammer (1941)°

Hammer (1941)

Hammer (1941) - Adults

Poorbaugh (1966) - Larvae

Hammer (1941)

Laurence (1953)

Mohr (1943)

Baumberger (1919)

Snowball (1944)

Hammer (1941)

Hammer (1941)

Hammer (1941)

91

LSISOTOWNOLNY DIMIDVG-NVg dH,

TAXA

Tipulidae (1)

HYMENOPTERA

Bethylidae (1)

Braconidae

Aphaereta pallipes (Say)

Asobara sp.

Idiasta sp.

Pentapleura sp.

Phenocarpa sp.

Cynipidae

Eucolia rufocincta (K)

Kleidotoma fossa K

Diapriidae

Phaenopria sp.

Figitidae

Figites sp.

Xyalophora sp.

Ichneumonidae

Phygadeuon sp.

a

Predators

Pparasitoids

Adult

mS PS POO

TABLE 1. (Cont.)

Inmature

Dung Pred Par

I

I

me PS OS OS

Other

“Bither phytophagous (e.g., nectar), saprophagous (e.g., fungi, bacteria), or unknown.

a

The number of species representative for that particular taxon given.

and the general nature of this table, each species has not been listed separately.

“2-presumed to feed, but not certain.

Reported

Prey

Unknown

Diptera pupae

Diptera pupae

Diptera pupae

Diptera pupae

Diptera pupae

Diptera Pupae

Diptera pupae

Diptera pupae

Diptera pupae

Diptera pupae

Diptera pupae

A

Reference

Citation

Baumberger (1919)

Blume (1970)

Garry and Wingo (1971)

Mohr (1943)

Mohr (1943)

Merritt (1974)

Turner et al. (1968)

Thomas and Wingo (1968)

Mohr (1943)

Mohr (1943)

Mohr (1943)

Kessler & Balsbaugh (1972)

Combs & Hoelscher (1969)

Due to the lack of information on the food habits of specific species

A complete list of species is given by Merritt (1974).

OL6L ANVANV[ ‘T ‘ON ‘ZS “IOA

LI

18 THE PAn-PAcIFIC ENTOMOLOGIST

et al. 1967). In each of these species the newly hatched larva entered

a fly puparium, developed within a parasite of the pupa, and emerged

as an adult bettle. The adult beetles were predacious on eggs and larvae

of muscoid Diptera (Wingo et al. 1967, Drea 1966). Host records of

parasitic staphylinids belonging to the genus Aleochara in America

were reviewed by Moore and Legner (1971). Members of the genus

Philonthus are predacious as adults and larvae on Diptera eggs and

immatures (Sanders and Dobson 1966, Macqueen 1973). Philonthus

cruentatus (Gmelin) was shown to be one of the most effective predators

on immature stages of the face fly, Musca autumnalis (DeGeer) (Valiela

1969, Wingo et al. 1974) and horn fly, Haematobia irritans (L.)

(Thomas and Morgan 1972b).

DIPTERA

Hammer’s (1941) work is the most complete study dealing with the

food habits of flies inhabiting cattle droppings. Information is lacking

on the adult habits of many species and more evidence is needed re-

garding the larval habits inside the dung (Table 1). Faunal studies

included lists of dipteran species “reared” from dung; however, this

does not necessarily imply that the specific species feeds entirely on

the dung itself (Valiela 1974, in part). The larvae may feed on micro-

organisms (e.g. fungi, bacteria) growing on the dung (Baumberger

1919) or they may be facultative carnivores, requiring live prey for

normal development (Muirhead Thompson 1937, Hammer 1941). For

example, several species of nematoceran Diptera have been reared from

cattle droppings (Hafez 1939, Mohr 1943, Poorbaugh 1966) ; however,

gut content analysis has shown that some members of this group feed

on decomposing vegetable matter (e.g., fungal spores and hyphae)

(Saunders 1924, Strenzke 1950) (Table 1).

Only two genera of Diptera were reported predacious as adults,

Scatophaga spp. (Anthomyiidae) (Hammer 1941, Foster 1970) and

Drapetis spp. (Empididae) (Laurence 1952) (Table 1). In northern

California, Foster (1970) found that Scatophaga stercoraria fed on

dung flies in the field, primarily of the families Anthomyiidae and

Sphaeroceridae. Laurence (1952) recorded an adult Drapetis sp. feed-

ing on a small staphylinid. The larvae of Drapetis are presumed preda-

tors but little is known about their habits (Rogers 1973, Papp 1971).

Other predacious larvae found in cattle droppings in north central

California were members of the genus Hydrotaea, and Myospila

meditabunda (F.) (Muscidae) ; however, they were found in relatively

low numbers (Merritt 1974). In contrast, Poorbaugh (1966) found

VoL. 52, No. 1, JANuary 1976 19

that the larvae of M. meditabunda caused heavy mortality to coproph-

agous fly populations in coastal California. Hammer (1941) re-

ported that Hydrotaea larvae could not live on a diet of dung alone but

required living fly larvae for development. He considered the adults

of Hydrotaea and those of another muscid, Morellia, as facultative

bloodsuckers, since they procure blood and secretions from wounds on

cattle. The majority of adult Diptera feed mainly on the liquids of dung

and often frequent flowers where they suck nectar (Hammer 1941)

(Table 1).

HYMENOPTERA

The Hymenoptera consisted largely of pupal parasitoids of Diptera

(Table 1). The search for biological methods of control against

pestiferous flies has led to much current research on the biology and

population dynamics of parasitic Hymenoptera and their associated

hosts. Theoretical and applied studies on the bio-ecology of natural

enemies are reviewed by Legner and Poorbaugh (1972). Further de-

scriptions of host-parasite associations can be found in Greenberg

(1971). Merritt (1974) found a braconid, Pentapleura sp. (P. tri-

ticaphis (Fitch) or P. foreolata Viereck) parasitizing Scatophaga sterco-

raria and S. furcata during the spring in the Sierra Nevada foothills.

DISCUSSION

In reviewing Table 1, two general observations can be made. First,

except for the empid Drapetis, no Diptera were reported predacious on

Coleoptera, while approximately 50 percent of the Coleoptera and all

the Hymenoptera have been cited as predators or parasitoids of Diptera.

Also, in contrast to northern Europe where Hammer (1941) found

several species of predacious Diptera larvae inhabiting cattle droppings,

only a few species were recorded from droppings in California (Table

1). Anderson and Poorbaugh (1968) implied a relationship between

fewer numbers of horn flies and face flies in Europe with higher levels

of predacious fly larvae, as compared to the situation in the United

States where there are fewer species of predacious larvae and higher

populations of pestiferous flies. Although Blume et al. (1970) showed

*““. .. a significant negative correlation (r = 0.898) between the mean

numbers of horn flies and the mean numbers of insects of other species

produced per dropping... ,” the exact nature of this relationship needs

further quantitative investigation. In Australia the pest fly situation

represented an extreme example where introduced pestiferous flies

breeding in cattle dung experienced little competition for food or losses

20 THE Pan-PactFic ENTOMOLOGIST

due to predation from other arthropods (Bornemissza 1960, 1970).

They have undertaken a major research program there to evaluate the

introduction of exotic dung beetles to reduce the numbers of pest flies

breeding in cattle droppings (Waterhouse 1974). .

The second general observation is that there are limited data regard-

ing the food habits of insect species inhabiting cattle droppings (Table

1). In the United States, life history studies on many groups inhabiting

dung, such as the Aleocharinae and other staphylinid subfamilies, have

not been conducted (I. Moore, Univ. Calif., Riverside, pers. comm.).

This is largely due to the difficulties involved in examining the inter-

specific and intraspecific interactions inside the dung. Clarifying the

several insect trophic levels within the cow dung community will require

more detailed investigations.

LITERATURE CITED

ANDERSON, J. R. AND PoorsaucH, J. H. 1968. The face fly Musca autumnalis ...

a new livestock fly is now moving toward California. Calif. Agric. 22:

4-6.

Baumpercer, J. P. 1919. A nutritional study of insects, with special reference to

microorganisms and their substrata. J. Exp. Zool.; 28: 1-81.

Biume, R. R. 1970. Insects associated with bovine droppings in Kerr and Bexar

Counties, Texas. J. Econ. Entomol., 63: 1023-1024.

Biume, R. R., Kunz, S. E., Hocan, B. F., anp Matter, J. J. 1970. Biological

and ecological investigations of horn flies in central Texas: Influence

of other insects in cattle manure. J. Econ. -Entomol., 63: 1121-1123.

BornemisszA, G. F. 1960. Could dung-eating insects improve our pastures?

J. Australian Inst. Agric. Sci., 26: 54-56.

Bornemissza, G. F. 1970. Insectary studies on the control of dung breeding

flies by the activity of the dung beetle, Onthophagus gazella F.

(Coleoptera: Scarabaeinae). J. Australian Entomol. Soc., 9: 31-41.

Bourne, J. R. ann Hays, K. L. 1968. Effects of temperature on predation of

horn fly larvae by the larvae of Sphaeridium scarabaeoides. J. Econ.

Entomol., 61: 321-322.

Coir, F. R. 1969. The flies of western North America. Univ. Calif. Press,

Berkeley. 693 p.

Comps, R. L. anp Hoetscuer, C. E. 1969, Hymenopterous pupal parasitoids

found associated with the horn fly in northeast Mississippi. J. Econ.

Entomol., 62: 1234-1235.

Drea, J. J. 1966. Studies of Aleochara tristis (Celeopten: Staphylinidae), a

natural enemy of the face fly. J. Econ. Entomol., 59: 1368-1373.

Foster, W. A. 1970. Predatory behavior of Scatophaga stercoratia (Diptera:

Anthomyiidae) in California. Ann. Entomol. Soc. Amer., 63: 338-339.

Gary, C. EK. anp Winco, C. W. 1971. Factors affecting parasitism of the face

fly by Aphaereta pallipes in laboratory studies. J. Econ. Entomol., 64:

104-107.

VoL. 52, No. 1, JANUARY 1976 21

GREENBERG, B. 1971. Flies and disease. Vol. I. Ecology, classification and biotic

associations. Princeton Univ. Press, Princeton, New Jersey. 856 p.

Harez, M. 1939. Some ecological observations on the insect fauna of dung. Bull.

Soc. Fouad Entomol., 23: 241-387.

Hammer, O. 1941. Biological and ecological investigations of flies associated

with pasturing cattle and their excrement. Vidensk. Medd. Naturhist.

Foren. Kobenhavyn, 105: 141-393.

KessLer, H. anp Bartspaucu, E. U. 1972. ‘Succession of adult Coleoptera in

bovine manure in east central South Dakota. Ann. Entomol. Soc. Amer.,

65: 1333-1336.

Lanpin, B. O. 1961. Ecological studies on dung beetles. Opuscula Entomol.

Suppl., 19: 1-228.

LaurENcE, B. R. 1952. The prey of some Empididae and Dolichopodidae (Dipt.).

Entomol. Mon. Mag., 88: 156-157.

LAuRENCE, B. R. 1953. Some Diptera bred from cow dung. Entomol. Mon. Mag.,

14: 281.

Laurence, B. R. 1954. The larval inhabitants of cow pats. J. Anim. Ecol., 23:

234-260.

Lecner, E. F. anp Poorsaucu, J. H. 1972. Biological control of vector and

noxious synanthropic flies: A review. Calif. Vector Views, 19: 81-100.

Macqueen, A. 1973. Horn fly breeding, nutrient immobilization associated with

cattle dung in the southern interior of British Columbia. Ph.D. Diss.,

Simon Fraser Univ., Burnaby, B.-C. 211 p.

Manis, H. 1934. Zur Kenntniss der morphologie, Okologie und physiologie von

Aphodius rufipes Lin. und einigen verwandten Arten. Zool. Jahrb., Abt.

Anat. Ont., 58: 303-396.

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habiting bovine manure in a pasture on the Big Sioux River flood

plain in South Dakota. Proc. S. Dak. Acad. Sci., 50: 220-237.

Merritt, R. W. 1974. The species diversity and abundance of insects inhabiting

cattle droppings and their role in the degradation of dung in four

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hills of California. Ph.D. Diss., Univ. California, Berkeley. 273 p.

Mour, C. O. 1943. Cattle droppings as ecological units. Ecol. Monogr., 13:

275-309.

Moors, I. anp Lecner, E. F. 1971. Host records of parasitic staphylinids of the

genus Aleochara in America (Coleoptera: Staphylinidae). Ann. Entomol.

Soc. Amer., 64: 1184-1185.

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the Anthomyiidae. Parasitology, 29: 273-358.

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22. THE Pan-Pacitric ENTOMOLOGIST

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(Diptera: Empididae). M.S. thesis, Univ. Calif., Berkeley.

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Chironomiden. Arch. Hydrobiol. (Plankt.), Suppl. Bd., 18: 207-414.

Tuomas, G. D. 1967. Natural enemies of the face fly Musca autumnalis DeGeer

in Missouri. Ph.D. Diss., Univ. Missouri, Columbia. 131 p.

Tuomas, G. D. anp Morean, C. E. 1972a. Parasites of the horn fly in Missouri.

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ture stages of the horn fly in Missouri. Environ. Entomol., 1: 453-459.

Tuomas, G. D. anp Winco, C. W. 1968. Parasites of the face fly and two other

species of dung-inhabiting flies in Missouri. J. Econ. Entomol., 61:

147-152.

Turner, E. C., Burton, R. P., anp GerHarDT, R. R. 1968. Natural parasitism

of dung-breeding Diptera: a comparison between native hosts and an

introduced host, the face fly. J. Econ. Entomol., 61: 1012-1015.

VALIELA, I. 1969. An experimental study of the mortality factors of larval Musca

autumnalis DeGeer. Ecol. Monogr., 39: 119-225.

VALIELA, I. 1974. Composition, food webs and population limitation on dung

arthropod communities during invasion and succession. Amer. Midl.

Natur., 92: 370-385.

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100-109.

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1514-1517.

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sion and abundance of insects in pasture manure: relationship to face

fly survival. Ann. Entomol. Soc. Amer., 67: 386-390.

RECENT LITERATURE

THe NortH AMERICAN Species OF HeETeRosaruS ROBERTSON (HYMENOPTERA:

AporwEA). P. H. Timberlake. University of California Publications in Entomol-

ogy, 77:1-56. 1975.

In this review of the panurgine bee genus Heterosarus, 55 species are treated,

37 being described as new. Male genital armature and subgenital plate are

figured for all species available.

Simple and Computerized Discriminant Functions for Difficult

Identifications: A Rapid Nonparametric Method

James A. Scott ano Jon H. SHEPARD

Natural Resource Ecology Lab., Colorado State University, Fort Collins, Co. 80521

and R. R. #2, Nelson, British Columbia

Many closely related species cannot be identified using single char-

acteristics alone because some individuals of both species have identical

characteristics. Discriminant functions have been developed that use

combinations of characteristics to identify species that cannot be identi-

fied using single characteristics alone (Dixon, 1965). The procedure

consists of measuring many characters for individuals known to belong

to each of the two species under study. A discriminant function equation

is then found using a computer program. The characteristics of an un-

known individual are then measured and inserted into this equation,

which provides a number that allows identification when compared

to the numbers given by known individuals. The purpose of this paper

is to illustrate a rapid nonparametric method and to compare it to

traditional linear and nonlinear discriminant functions.

Stupy ANIMALS

Papilio glaucus L. and P. rutulus Lucas are known to hybridize in the

laboratory and apparently hybridize in British Columbia and South

Dakota (Clarke & Sheppard, 1955, 1957, 1962; Brower, 1959a, 1959b).

Canadian populations are intermediate in many respects, and P. glaucus

and P. rutulus may in fact be subspecies.

Eight characteristics were quantified for each sex. For males, A, B,

C, D, and E (Fig. 1) were measured lengths of male genitalia

(all lengths are mm). P describes the form of the prong of Figs. 2-4

(1-no lateral processes; 3-a long lateral process; 2-intermediate). F

for males and females is the left forewing length in mm. V for males

and females describes the amount of red in the anterior submarginal

ventral hindwing light spot (l-all yellow; 2-slightly red; 3-half red;

4-mostly red; 5-all red). For females, G, H, and K are measurements

of female genitalic structures (Fig. 7). L describes the form of a lobe

(1-leaflike as in Fig. 6; 3-bladelike as in Fig. 5; 2-intermediate). S de-

scribes the shade of a flange (Fig. 7) (1l-almost transparent; 3-dark

gray; 2-intermediate). B describes the dorsal forewing color (1-pale

yellow; 3-lemon yellow; 2-intermediate).

Tue PAN-PactFic ENTOMOLOGIST 52: 23-28. JANUARY 1976

24, Tue Pan-Pactric ENTOMOLOGIST

A Raprw NonLINEAR NONPARAMETRIC DISCRIMINANT FUNCTION

The simple method involves three steps: 1) choosing characters

which differ between the two species. Characters for males were A/B,

C/F, E/F, B/C, D/F, P, and V; for females, G/F, L, H/F, 5, K/F, V,

and B; 2) determining means for each character for each species using

individuals known to be correctly identified; 3) characters with means

larger in species 1 than in species 2 are multiplied by each other in the

numerator of the discriminant function. Characters with means smaller

in species 1 than in species 2 are multiplied by each other in the

denominator. With P. rutulus as species 1 and P. glaucus as species 2,

the discriminant functions are:

SDF é = (B/C D/F E/F G/F) /(A/B P V),

SDF? = (G/F K/F)/(LS H/F VB)

COMPUTERIZED DISCRIMINANT FUNCTIONS

These discriminant functions were calculated using the computer

program of Dixon (1965). The linear ones are:

LDF é = 1.69A — 1.60B — .63C — 2.92D — .23E + .88P

— .00056F + .61V

LDF? = .30S — 2.16G + 3.35H + .17K —1.14F + 1.43V — .22B

The nonlinear ones are:

NLDF¢é = (8.34LnA — 12.64LnB — 6.53LnC — 5.13LnD

—1.30LnE + 3.14LnP + 2.75LnF + 9.55LnV ) /2.30

NLDF®? = ~.71LnS + 3.45LnG + 3.91LnH + .59LnK

— 21.06LnF + 11.47LnV + .21LnB

In all of these discriminant functions to identify an unknown individual

the characters (letters) are found, substituted into the equations, and

the result is compared to results from individuals of known identity.

The results can also be plotted along a single axis for comparison; for

the simple function this axis should be logarithmic (Fig. 8). These

discriminant functions mathematically maximize the difference between

the results for the two species. The computerized functions also minimize

the squared deviations from the mean result for the known individuals

of each species.

VoL. 52, No. 1, JANuARY 1976 25

<—E—>

{

Fics. 1-7, male and female genitalia. Fig. 1, male genitalia, illustrates mea-

surements A to E, and prong (P). Three shapes of the prong are designated 1

(Fig. 2), 2 (Fig. 3), and 3 (Fig. 4). Fig. 7, female genitalia, illustrates three

measurements and the lobe (L) and flange (B). Two shapes of the lobe are

designated 1 (Fig. 6) and 3 (Fig. 5).

26 THE Pan-Paciric ENTOMOLOGIST

SDF Oo” XX

Xl XXXX 4

1 LUM XDOCX XX XXO XO

SDF?

XX X xX 0)

30 50 100 = 200.~—Ss« 500 1000 2,000 5,000 ‘10,000

LDF Oo” x

x |

x X

dd3 XX X XXXXX + ll

LDF

: ’

Xxx XX XX

-7 -5 -3 -| O | 3 5

NLDFo”

OOX xX XX D4

OOX X X x xX XXXX It It

-4 -2 O 2 4 6 8

NLDF

t x

O XXX XX

-86. -82 -78 -74 -70 -66 -62

Fic. 8. Discriminant function results for Papilio glaucus from eastern United

States to Alberta and Alaska (vertical lines), P. rutulus from California to

Washington (0), and for British Columbia individuals (x). The scale for SDF,

and for LDF, is the same for males and females.

ADVANTAGES OF THE Rapip DISCRIMINANT FUNCTION

There are four advantages: 1) it is simple and rapid, without the

necessity for computer programs, which may be unavailable; 2) it is

nonparametric, which means that the data do not have to conform to

a probability distribution as do the computer methods; 3) since the

method depends only on the ranking of means, it will not vary with the

addition or subtraction of individuals from the group of known indi-

viduals from which the means were derived, except in the unlikely event

that the ranking of the means of a character for the species changes.

If that happens, that character would seem to be of little use for identi-

fication, and it should be expunged. The computerized discriminant

functions will change somewhat with the addition or subtraction of

VoL. 52, No. 1, January 1976 aT

each individual, and will stabilize only with large sample sizes. 4)

Qualitative (arbitrarily rated numerically) characters are easily used

in the simple function, but are very difficult to incorporate into the

computerized functions if they do not vary greatly. For example, one

of the best characters for females is L, yet its lack of variation in the

known individuals prevented it from being used in LDF and NLDF.

Multiplying or dividing any character by a constant (scaling) will

not affect the identifications using SDF in any way. The simple dis-

criminant function therefore does not weight characters, whereas the co-

efficients of the computerized functions are weights. In many taxonomic

applications it may be preferable not to weight characters (Sneath &

Sokal, 1973), and with small sample sizes of known individuals the

computer weights may be unreliable. A possible disadvantage of the

simple function is that the computerized methods may provide better

identification if large numbers of known individuals are used; this re-

mains to be determined.

RESULTS

All three functions provided excellent identification of the known

individuals, and nearly identical identification of the unknowns from

British Columbia (Fig. 8). The British Columbia sample includes both

“species” and individuals in varying degrees of intermediacy. A further

breakdown of the British Columbia sample indicated that central British

Columbia individuals were mostly P. glaucus and southeastern British

Columbia individuals were usually intermediate. We conclude that inter-

gradation does occur between the species in British Columbia. The

results are not sufficient to determine whether the integradation is

introgression between species or simply hybridization between sub-

species with additive or non-additive (Mendelian) inheritance of char-

acters. The results do provide useful methods for further study of this

problem. Full data may be obtained from the authors.

ACKNOWLEDGMENT

The University of California, Davis, provided a small grant for com-

puter time. Specimens are from the collections of J. Shepard, J. Scott,

the California Insect Survey, Berkeley, California, and the University

of California, Davis, California. Both authors were supported by Na-

tional Institute of Health traineeships during the initial stage of the

study.

28 THE PAn-PAcIFIC ENTOMOLOGIST

LITERATURE CITED

Brower, L. P. 1959a. Speciation in butterflies of the Papilio glaucus group.

I. Morphological relationships and hybridization. Evolution 13: 40-63.

Brower, L. P. 1959b. Speciation in butterflies of the Papilio glaucus group.

I. Ecological relationships and interspecific sexual behavior. Evolution

13: 212-228.

CiarKe, C. A. AND SHepparD, P. M. 1955. The breeding in captivity of the

hybrid Papilio rutulus female < Papilio glaucus male. Lepid. News.

9: 46-48.

CiarRKE, C. A. AND SHEPPARD, P. M. 1957. The breeding in captivity of the

hybrid Papilio glaucus female * Papilio eurymedon male. Lepid. News

11: 201-205.

CuarKE, C. A. AND SHepparpD, P. M. 1962. The genetics of the mimetic butterfly

Papilio glaucus. Ecology 43: 159-161.

Dixon, W. J. [Ed.]. 1973. BMD biomedical computer programs. University of

California Press, Berkeley, California. 773 p.

SNEATH, P. H. A. anp Soxat, R. R. 1973. Numerical taxonomy. The principles

and practice of numerical classification. W. H. Freeman Co., San

Francisco, California. 573 p.

ZOOLOGICAL NOMENCLATURE

ANNOUNCEMENT A.N.(S.) 97

Required six months’ notice is given of the possible use of plenary powers by the

International Commission on Zoological Nomenclature in connection with the

following names listed by case number: (see Bull. Zool. Nom. 32, part 3, 22nd

September, 1975).

1003. Chaitophorus Koch, 1854 (Insecta, Hemiptera): designation of type-species.

2060. Xiphidium glaberrimum Burmeister, 1838 and Orchelimum cuticulare

Audinet-Serville, 1838 (Orthoptera) ; suppression; designation of Orcheli-

mum vulgare Harris, 1841 as type-species of Orchelimum Audinet-Serville,

1838.

2107. Polydrusus Germar, 1817 (Insecta, Coleoptera): designation of type-species.

2109. Notozus Forster, 1853 (Insecta, Hymenoptera, Chrysididae): designation

of type-species; Elampus Spinola, 1806: proposed suppression.

Comments should be sent in duplicate, citing case number, to the Secretary,

International Commission on Zoological Nomenclature, c/o British Museum

(Natural History), Cromwell Road, London, S.W.7 5BD, England. Those

received early enough will be published in the Bulletin of Zoological Nomenclature.

—R. V. Metvitte, Secretary to the International Commission on Zoological

Nomenclature.

Observations on the Nesting Behavior of

Belomicrus columbianus Kohl

(Hymenoptera: Sphecidae)

Steven K. Autt!

Department of Parasitology, Liverpool School of Tropical Medicine,

Liverpool L3 5QA, England

Observations on the nesting behavior of Belomicrus columbianus

Kohl were made in late June and early July 1972. The study area was

located on the east slope of Mount Rose, Washoe Country, Nevada at

an altitude of approximately 8000 feet (2462 m.). This arid area

consisted of dry sand and gravel, interspersed with rocks less than 3

cm diameter, and covered with sparse low vegetation.

Belomicrus Costa is a genus of gregarious fossorial wasps, with more

than 50 species, thought to be an evolutionary intermediate between

strictly solitary wasps and semi-social wasps and which today occurs

in the Nearctic, Palaearctic, and Ethiopian regions (Evans and Eberhard,

1973). Very few published accounts of the biology of Belomicrus exist.

Nesting behavior of Belomicrus franciscus Pate and Belomicrus forbesit

forbesii (Robertson) in North America was described by Williams

(1936) and Evans (1969), respectively.

NESTING BEHAVIOR

In this study the height of nest excavation activities for female

B. columbianus occurred about 1300-1330 when ground temperature

was approximately 27°C., after a slow beginning at about 1000 when

ground temperature and wasp activity was minimal. In the nesting area

the east-facing banks were the first to become active with wasps. As

many as 6 hours of the daylight period were required for a female to

commence and complete a nest, provisioned with her eggs and prey.

“Trial digging” was observed in various places within 10 meters of the

final chosen nesting site, with any incompleted burrows left open.

Belomicrus is not known to use pre-existing animal-made holes for

nesting sites (R. M. Bohart, pers. comm., 1972).

The selected site was usually on the leeward side of a sandy bank

with slopes less than 30°. Soil removal commenced with the female

digging rapidly, apparently with mandibles and fore legs, while assuming

a 30° angle to the ground. Pellets of earth were carried between head,

1 Presently Department of Entomology, University of California, Davis.

THe PAN-PactFIc ENTOMOLOGIST 52: 29-32. JANUARY 1976

30 THe PAn-PAciFIC ENTOMOLOGIST

thorax, and the front pair of legs. The ventral posterior surface of the

head bears a fringe of long hairs, termed a psammophore, which

assists in holding the pellet of earth (Evans and Eberhard, 1973). The

female flew obliquely backwards out of the burrow with the pellet,

dumping it about 15 cm from the entrance site, from a height of about

40 cm. This movement required two seconds. No tumulus was formed

around the nest as the soil was dumped at random in a semicircle around

the nest entrance. Immediately after dumping the soil the wasp flew

rapidly back into the burrow to repeat the action. The nest entrance

measured 3-4: mm diameter when complete. During the digging process

occasional short flights (of 1-3 minutes duration) away from the nest

were observed, the wasp returning without prey.

One female B. columbianus was observed entering the incompleted

burrow headfirst without prey. It plugged up the burrow entrance by

grasping, apparently with its mandibles and fore legs, a 3 mm pebble

located immediately outside the entrance and pulling it into the entrance,

partly blocking it. Staying inside the burrow for 15 minutes, it then

removed the pebble, placing it to the immediate side of the entrance.

It then left the nest, head-first, and flew away without a load of earth.

I assume that during the time in the nest the wasp was digging and

filling deep in the ground. The lengthy amount of time was probably

due to the complex of many cells being prepared (as will be described

later).

Disruption of the digging process occurred if one came within about

30 cm of the nest entrance or the wasp. When disturbed the wasp flew

about 2 meters away from the entrance, then returned to land within one

meter of the nest, remaining there 1-2 seconds. This action was repeated

4—5 times, the female coming closer to the entrance each time. No

particular orientation seemed to be chosen, i.e. the wasp did not neces-

sarily face the source of the disturbance or the nest entrance.

Twice a species of the solitary wasp Diploplectron (det. R. M. Bohart)

was observed entering a B. columbianus nest before it was completed,

the duration of the stays being 2 seconds and one minute. Then

Diploplectron left the nest carrying nothing with it. I assume it was

merely resting in the nest since Diploplectron is not known to be para-

sitic or nest in pre-existing burrows (R. M. Bohart, pers. comm., 1972).

Other Crabroninid and Chrysidid wasps and various sarcophagous

“satellite” flies were also observed around the nesting sites but no

interaction was observed between these insects and B. columbianus.

Ants were observed to enter and quickly leave the nest entrance, without

VoL. 52, No. 1, January 1976 51

finding prey; the presence or absence of the wasp in the nest having

no apparent effect on the ant’s behavior.

Immediately after completion of the nest the female would begin

provisioning the nest with prey, this activity usually beginning after

1400. Upon beginning the search for prey the wasp usually plugged,

partly or completely, the entrance to the nest as described previously,

then would fly away, returning in 5-30 minutes with or without prey.

At the wasp’s return, when burdened with prey, it would rest 1-2

seconds on the ground after travelling a few meters, then recommence

these short, hopping flights back to the nest. Prey were observed being

carried between the middle and hind pairs of legs of the wasp, the

abdomen of the prey held to the ventral side of the wasp, head forward

and abdomen sometimes protruding beyond the wasp’s abdomen.

Reaching the nest entrance, if it was plugged with a pebble, the

wasp would place its prey at the immediate side of the entrance then

remove the pebble, which was also placed near the entrance, grasp the

prey with its mandibles and fore legs and enter the nest abdomen first.

If the nest entrance was not plugged the wasp would fly directly into the

burrow. All prey appeared to be immobilized, probably by sting. The

wasp would remain in the nest from 1—25 minutes, leaving the entrance

unsealed. Then it would emerge from the nest head-first, usually sealing

the entrance as before, and fly away in search of further prey.

Nest ARCHITECTURE

At the study site more than 20 individual occupied nests could be

found in an area of 3 meters square, and nest entrances were located

as close as 2 cm to each other. In order to examine the nest architecture

of B. columbianus 12 nests were removed from the study site in whole

cores of earth, each core approximately 20 cm deep and 15 cm diameter.

Removal of the cores from the dry sandy soil necessitated moistening

of the core with water while carefully digging around the edge of the

designated core with a hand trowel. The whole cores were placed in

plastic bags to prevent collapse and loss of moisture while being trans-

ported to the Sagehen Creek Field Station, University of California

(near Truckee, California) for examination.

Seven of the 12 cores examined yielded prey and wasp eggs; the

other 5 contained either incompleted or empty nests. The entrance

tunnel descended vertically to a depth of approximately 4.5 cm, where-

upon it branched into 48 further tunnels (of 2 cm length) in a hori-

zontal or slightly oblique plane, each tunnel concluding in a single

cell of size 10 X 4mm. The cells were 1.6-2.0 mm apart and arranged

32 Tue Pan-Paciric ENTOMOLOGIST

in a semicircle. Tunnel diameter throughout the nest was about 3 mm.

A range of 5-13 prey could be found in each cell, with 7 prey the

mode.

Two undetermined species of immature Hemipterans were the only

prey found in the cells, one being a Mirid bug. No prey or wasp eggs

were found in the passages to the cells. All bugs were semi-

paralyzed, twitching only slightly when probed. The wasp’s egg was

found deposited (one egg per cell) on one of the two prey found at the

far end of the cell, with further bugs placed, presumedly, afterwards.

It is assumed the bug with the egg attached is the first (or second)

bug placed into the cell by the wasp. The egg was always attached in

a transverse position cephalad of the left front coxa of the bug. One egg

measured 1.44 X 0.40 mm, the egg shell being pale white and translucent,

with a pale yellow embryo visible inside.

A total of 11 wasp larvae, of different instars, were found in the

examined nests, often in association with eggs in that nest; but a larva

apparently remains in its cell as none were found with other larvae or

eggs in the same cell. Larvae ranged in size from 2.1 X 0.7 mm to

3.6 X 1.2 mm, appearing a translucent white with green-brown viscera.

ACKNOWLEDGMENTS

I wish to express my gratitude to Dr. R. M. Bohart, University of

California at Davis for identification of the wasp specimens and guidance

throughout the course of the investigation. I am also grateful to Shiela

M. Casey, Douglas Edwards, and Robin McMasters, all formerly of U. C.

Davis, for their invaluable assistance in this project.

LITERATURE CITED

Evans, H. E. 1969. Notes on the nesting behavior of Pisonopsis clypeata and

Belomicrus forbesii (Hymenoptera, Sphecidae). J. Kans. Entomol.

Soc. 42: 117-125.

Evans, H. E. anp Eperuarp, M. J. W. 1973. The Wasps. David and Charles

(Holdings) Ltd., Great Britain. 265 pp.

Wiutams, F. X. 1936. Notes on two oxybelid wasps in San Francisco, California.

Pan-Pac. Entomol. 12: 1-6.

Seasonal Changes in Host Plant Preference of the

Blue-green Sharpshooter Hordnia circellata

(Homoptera: Cicadellidae)

A. H. PurceLyu

Department of Entomological Sciences, University of California, Berkeley 94720

In California’s north coastal counties, the blue-green sharpshooter

Hordnia circellata (Baker) is the most important vector of Pierce’s

disease of grapevines (Hewitt et al., 1942, Severin 1949, Winkler et al.,

1949, Purcell 1975). Freitag (1951) listed 93 species in 23 families of

natural and cultivated plants as hosts of the causal pathogen of Pierce’s

disease. Thus the relative importance of various plant species as food

and oviposition hosts of H. circellata is of considerable interest relative

to the epidemiology of Pierce’s disease as well as to the ecology of its

chief vector in coastal California.

The blue-green sharpshooter is found on many host plant species.

Frazier (in Winkler, 1949) reported that more than 150 species of plants

had been recorded as hosts. Severin (1949) noted that a wide variety

of ornamental species in Berkeley, California were commonly infested.

In the course of trapping and collecting these leafhoppers in Napa

County, California from 1972 to 1975, I noticed that the host plants on

which H. circellata was most commonly found followed a seasonal

sequence.

At least twice per month and usually once a week from April, 1972

to November, 1974. and much less frequently in 1975, I searched for and

collected H. circellata at 4 sites along the Napa river between Yountville

and St. Helena, California, and at 2 sites on the eastern slope of Spring

Mtn. 3 to 5 mi. west of St. Helena. All of these locations were adjacent

to commercial vineyards, described in a previous publication (Purcell

1975). Collecting was by a 15 in. sweep net or direct search of foliage.

RESULTS

Fig. 1 summarizes the most commonly inhabited host plants along

vineyard study sites in the Napa Valley. The length of the apparent

attractiveness of any particular host plant varied as much as three

weeks from year to year, especially from late Aug. until November and

from location to location depending on site factors such as shade and

soil moisture, as will be discussed in a later section.

Nymphal Host Plants.—The most important hosts for oviposition and

Tue Pan-PactFic ENTOMOLOGIST 52: 33-37. JANUARY 1976

34 THE Pan-Pactric ENTOMOLOGIST

subsequently for nymphal development were relatively few compared to

the number of food hosts (Fig. 1). Blackberries, especially Rubus

procerus P. J. Mueller and R. vitifolius Chamisso and Schlechtendal;

elderberry, Sambucus caerulea Rafinesque and S. mexicana Presl; wild

grape, Vitis californica Bentham; Mugwort, Artemesia Douglasiana

(vulgaris of Jepson) Besser (in Hooker); and nettle, Urtica spp., were

the most common breeding hosts as shown in Fig. 1. I occasionally

collected nymphs on cocklebur, Xanthium strumanium (L.); California

figwort, Scrophularia californica Chamisso and Schlechtendal; and Ash,

Fraxinus sp. In the mountains of the northern Coast Range, a few

nymphs were found regularly on Rubus parviflorus Nuttall, usually in

moist, shaded sites.

Condition of Plant Growth.—In addition to plant species, the condition

of plant growth and site factors such as shade and soil moisture seemed

to influence host plant selection. Frazier (in Winkler, 1949) noted the

feeding preference of this and other Cicadellinae (Tettigellinae) for the

succulent tips of growing stems. This habit is quite pronounced in

cultivated grapes (Purcell 1975) and extends to most other hosts. The

only nymphs I have found on oak, Quercus spp. and walnut, Juglans

californica Watson, were on vigorously growing shoots or suckers

induced by severe pruning of the trunk or large branches. H. circellata

can imbibe quantities of xylem sap daily that are hundreds of times its

own body weight (Mittler 1967). Soil water tension and host plant

physiological condition may affect the rate of flow of xylem sap and

thus mediate host plant attractiveness for xylem feeding insects.

During the hot, dry California summer, H. circellata may be commonly

found on blackberries, artemesia, and cocklebur only in shaded sites

where soil moisture is sufficient to sustain continued rapid plant growth

and is rarely found in open sites such as along railroad beds, roads or

fencerows where these plants may be abundant but produce very little

new growth as soil moisture is depleted. Cocklebur, for example, growing

in full sun in unirrigated vineyards in Napa Valley seldom exceeds

1 m in height, whereas cocklebur growing in the shaded understory along

the edge of a river often exceeds 2 m in height. Only the latter plants are

commonly inhabited by adult and occasionally nymphal H. circellata.

Cultivated grapes, Vitis vinifera L. are normally grown in full sun

without irrigation in the Napa Valley and can support large numbers

of the blue-green sharpshooter. Grapes, however, are heavily pruned

and respond with luxuriant new growth as long as sufficient moisture

is available. On shallow soils—typically in stony hillside vineyards—or

VoL. 52, No. 1, JANuARY 1976 39

Xanthium strumarium

Vitis californica

Artemesia douglasiana

Rubus spp.

Urtica spp.

Rosa californica

Sambucus spp.

Salix spp.

Scrophularia californica

Symphoricarpus rivularis

Baccharis pilularis

Malva parviflora

Vinca major

Umbellularia californica

Mentha arvensis

Chenopodium spp.

Fic. 1. Seasonal occurrence of Hordnia circellata on plant hosts, Napa Valley.

Heavy solid line indicates period when H. circellata is commonly found; solid line

indicates period of frequent but much less common occurrence; dashed line

indicates only occasional occurrence. Vertical spikes indicate nymphs common.

Plants identified according to Munz and Keck (1970).

in abandoned vineyards where grapevines produce much less succulent

growth, Hordnia is only rarely found.

Fall Transition Period.—From late August to mid-September, as

cultivated grapevines are senescing, H. circellata move to natural

vegetation from vineyards as shown by sticky trap catches of flying

adults (Purcell 1975). Normally at this time of year adults can be

found on a variety of plant species which are either senescent or not

adding new growth. Coyote bush, Baccharis pilularis De Candolle, and

walnut, Juglans spp., are frequently fed upon during the early autumn or

late summer despite their lack of recent growth. Freitag and Frazier

(1954:) observed adults leaving vineyards to feed on nearby Persian

walnut Juglans regia L. I found these sharpshooters feeding on the

fruits of leafless horse chestnuts, Aesculus californica Nutall, in Novem-

ber 1975 along the Napa River. Relatively large catches on yellow

sticky traps at this time of year (Purcell 1975) contribute to the

supposition formed by direct observation that adults may change hosts

36 THE Pan-PaciFiIc ENTOMOLOGIST

in rapid succession during the fall transition to cooler, wetter weather.

Cool temperatures seem to limit flight activity below 16°C (my

unpublished data).

Winter Hosts—From early to mid-November until late March,

sharpshooters are difficult to locate. A few can be found on Urtica spp.

growing in damp localities, Malva parviflora L., laurel, Umbellularia

californica Nuttall, chicory, Cichorium intybus L., Chenopodia spp.

including C. murale L. and C. album L., and rarely on other herbaceous

hosts. Ornamental species escaped from cultivation such as Vinca major

L. and ivy, Parthenocissus tricuspidata Planchon, are also fed upon

during winter. Feeding adults are found most frequently during this

period beneath trees (leafless or not) or along stands of brush along

drainage ditches or streams. They are conspicuously absent from the

usually dense growth of herbs and grasses in open vineyards, pastures,

or meadows.

The gradual disappearance of adults in the fall and their rather

sudden reappearance in the early spring in the same location suggests

that adults do not normally disperse more than a few hundred meters

from fall until spring. I have not recovered adults from leaf litter,

beneath logs, bark, or other such suspect sheltered sites despite a great

number of attempts. Adults may be found at all times during the winter

in Berkeley, which has a mild winter climate, although they are difficult

to locate on very cold days.

Effects of Host Plants on Sharpshooter Distribution.—The geographic

distribution of H. circellata extends from Mexico and Arizona (DeLong

and Severin 1949, Nielson 1968) to northern Oregon and probably at

least as far north along the coast as Puget Sound or beyond. In essence

Hordnia is an oasis dweller, whether its habitats are ornamental and

shade trees in the desert town of Indio, California where it is an

abundant nuisance pest, on stream-bank vegetation in the Chiracahua

Mountains (Nielson 1968), a small bog at the bottom of a mountain

ravine on the western slope of the Napa Valley, or the well-tended home

gardens and parks of the San Francisco Bay area. DeLong (1965) has

described the influence of site factors such as soil moisture and shade

upon leafhopper food and habitat selection. These same factors seem

to influence the host plant preference of H. circellata profoundly.

ACKNOWLEDGMENTS

This research was supported by the Napa Valley Viticultural Research

Fund. I thank Dr. N. W. Frazier for his critical review of the

manuscript.

VoL. 52, No. 1, JANuARY 1976 37

LITERATURE CITED

DeLonc, D. M. 1965. Ecological aspects of North American leafhoppers and

their role in agriculture. Bull. Entomol. Soc. Amer. 11(1): 9-26.

DeLonc, D. M. ann H. H. P. Severtn. 1949. Characters, distribution, and food

plants of leafhopper vectors of virus causing Pierce’s disease of grape-

vines. Hilgardia 19(6): 171-186.

Freitac, J. H. 1951. Host range of the Pierce’s disease virus of grapes as

determined by insect transmission. Phytopathology 41: 920-934.

Freitac, J. H. anp N. W. Frazier. 1954. Natural infectivity of leafhopper vectors

of Pierce’s disease of grapes in California. Phytopathology 44: 7-11.

Hewitt, W. B., N. W. Frazier, H. E. Jacop anp J. H. Frerrac. 1942. Pierce’s

disease of grapevines. Calif. Agric. Exp. Sta. Circ. 353. 32 pp.

Mirtiter, T. E. 1967. Water tensions in plants—an entomological approach.

Ann. Entomol. Soc. Amer. 60(5): 1074-1076.

Muwnz, P. A. anp D. D. Kecx. 1970. A California flora. Univ. of Calif. Press.

Berkeley. 1681 pp.

Neitson, M. W. 1968. The leafhopper vectors of phytopathogenic viruses

(Homoptera, Cicadellidae) taxonomy, biology, and virus transmission.

U.S. Dep. Agr. Tech. Bull. 1382. 386 pp.

PurceLtt, A. H. 1975. Role of the blue-green sharpshooter, Hordnia circellata,

in the epidemiology of Pierce’s disease of grapevines. Environ, Entomol.

4(5): 745-752.

Severin, H. H. P. 1949. Life history of the blue-green sharpshooter, Neokolla

circellata. Hilgardia 19(6): 187-189.

Winker, A. J. (Ed.). 1949. Pierce’s disease investigations. Hilgardia 19(7):

207-264.

SCIENTIFIC NOTE

A New Record of Purpuricene Cerambycid from America North of

Mexico.—Two specimens of a very distinct cerambycid were collected by the

author and P. H. Sullivan, at Kitt Peak in the Baboquivari Mountains, Pima

County, Arizona, in September 1969. Through reference with the Biologia Centrali

Americana, these specimens have been tentatively identified as Mannophorus

forreri, Bates. Records of this species have been limited to Durango, Mexico, and

the discovery of these specimens can be considered an important addition to the

knowledge of the cerambycid fauna of the United States. Since the initial record

in 1969, numerous specimens have been taken on various species of Compositae

at the Kitt Peak site. Specimens are in the collections of D. G. Marqua, A. E.

Lewis, F. T. Hovore, E. M. Geisbert, California Academy of Science Sciences,

San Francisco, and the University of California, Berkeley—Davin G. Margua,

Los Angeles County Nature Centers, 1000 N. Durfee Avenue, S. El Monte,

California 91733,

Further Studies Among the Mexican and Central

American Lepturine Cerambycidae related to

Euryptera and Choriolaus

(Coleoptera)

Joun A. CHEMSAK AND E. G. LINSLEY

Division of Entomology and Parasitology, University of California, Berkeley

During the preparation of the volume on Lepturinae for the Ceram-

bycidae of North America, it has been necessary to re-examine the

generic status of many of the representatives of this subfamily which

have been described from Mexico and Central America. Some of these

studies, as they relate to genera and species in the Euryptera-Choriolaus

complex have been published previously (Linsley, 1961, 1970; Linsley

and Chemsak, 1971; Chemsak and Linsley, 1974). Generic assignments

in these and publications of earlier writers are now revised and updated

and some of the more significant new genera and species now available

are described.

This study was supported by the National Science Foundation (Grant

GB-BM574.) for a monograph of the North American Cerambycidae.

Authorities at the following institutions and collections are acknowl-

edged for the loan of specimens from their own or collections in their

care: California Academy of Sciences, San Francisco; Essig Museum

of Entomology, University of California, Berkeley; Los Angeles Museum

of Natural History; National Museum of Natural History, Washington,

D. C.; G. Nelson; and J. M. Campbell. Celeste Green prepared the

illustrations.

GENUS CHORIOLAUS BATES

Choriolaus Bates, 1885, Biologia Centrali-Americana, Coleoptera, 5:283; Boppe,

1921, Genera Insectorum, 178:107; Chemsak, 1964, Pan-Pacific Ent., 40:232

(designation of type species) ; Linsley and Chemsak, 1971, Arquivos de Zoologia,

21:6.

Form moderately robust, elytra not attenuated, sides subparallel or very feebly

expanded before rounding to apices. Head moderately elongate; eyes emarginate;

antennal insertions adjacent to upper inner edge of lower lobe of eyes, widest

anterior portion of lower lobes extending slightly beneath insertions but lobes

not embracing them; antennae filiform, outer segments not distinctly swollen or sub-

serrate, first four and one-half segments shining, moderately finely, densely

punctate, usually with coarse, semi-erect black hairs, segments five to eleven dull,

minutely, densely puncate, densely clothed with very short, appressed pubescence,

third segment elongate, subequal in length to scape, nearly as long or as long

Tue Pan-Paciric Entomo.tocist 52: 38-49. JANuARY 1976

VoL. 52, No. 1, January 1976 39

as fifth segment, fourth segment slightly shorter than both third and fifth,

poriferous areas of outer segments small or obsolete. Pronotum narrowly campanu-

liform, sides slightly rounded at middle, length subequal to basal width, punctation

of disc as coarse as or coarser than that of base of elytra, posterior angles acute,

extending over humeri; prosternum very narrow and laminiform between anterior

coxae, coxae prominent, exserted, cavities closed behind; mesosternum with

intercoxal process at midpoint between coxae distinctly narrower than vertical

diameter of base of intermediate femora. Elytra without costae; apices rounded to

outer angle which is feebly to distinctly dentiform. Legs with posterior tarsi

usually slightly longer than tibiae in the male and subequal in length to tibiae

in female. Abdomen with last sternite rotundate-truncate in male, rounded in

female.

Type species—Choriolaus latescens Bates (Chemsak designation, 1964).

As restricted above, Choriolaus may be distinguished among related

genera here discussed by the moderate, rather than small or large size,

filiform antennae which are not distinctly enlarged or subserrate

apically, and the moderately coarsely punctate pronotum with the

punctures subequal in size to, or larger than, those of the base of the

elytra. In addition to C. latescens Bates and C. aegrotus Bates from

Oaxaca, both originally included in the genus by Bates, the following

species fall within the present definition of the genus: celestae Chemsak

and Linsley, new combination (Megachoriolaus) (Chiapas), filicornis

Linsley and Chemsak, new combination (Megachoriolaus) (Chiapas),

gracilis Chemsak and Linsley, new combination (Megachoriolaus)

(Costa Rica), sabinoensis Knull, new combination (Euryptera) (Ari-

zona), and the new species described below.

KEY TO THE SPECIES OF CHORIOLAUS

iF Posterior tarsi distinctly shorter than posterior tibiae WW. 2

Posterior tarsi subequal to or longer than posterior tibiae 3

2(1). Body wholly rufo-testaceous except antennae, apices of femora, tibiae and

tarsi which are black; eyes small; antennal sockets completely outside

of ocular emargination — sabinoensis

Body dark brownish-black; face above clypeus, vertex, neck, pronotum

at sides, intercoxal process of prosternum, and elytra golden yellow; eyes

moderate, antennal sockets adjacent to and slightly within lower lobe of

eye, but not embraced by eye We frees) ee al, ss Se filicornis

3(1). Head yellowish-orange or yellowish-testaceous —_ 4

oo Eee te aN ET) "sg Ee ae cea. LORY SRR een A Lty eR eW NEE Le ie. Eme Ha rate 6

4(3). Elytra red or scarlet; elytra with punctures mostly separated by one or

TU ERT 8 eae ec wrsrenaonacti genie nesdannaentccdic genta seo ttanatias sts ean ead

Body yellowish-testaceous except eyes, antennae, extreme apices of

femora, tibiae and tarsi which are black, and the sterna which are dark

brownish-black margined with yellow; elytra with punctures mostly

separated by less than one diameter —_--- aegrotus

40 THe Pan-Pactric ENTOMOLOGIST

5(4). Head and pronotum yellowish-orange; elytra scarlet; legs, including

coxae, mesosternum, metasternum and abdomen black; pronotum finely,

densely punctate, punctures mostly contiguous _ celestae

Head, entire ventral surface, and femora except apically, yellowish-

orange; pronotum and elytra reddish; pronotum with punctures mostly

Saf Ug! Eee OR SPER Ns IRE Oe ce PE ne Oe: gracilis

6(3). Dark brown, pronotum and prosternum, basal three-fifths of elytra, and

femora golden yellow; elytra with external angle of apices acutely

[SURAT 1301 100) Pe, al Ae EL EP eS yl CO ee Re latescens

Black, pronotum and elytra rufo-testaceous; elytra with external angle

bere 1G am” Lea ¥: 1} OE a En OTD ORIENT ame SOE ETS derhami

Choriolaus derhami, new species

Male: Moderate sized; integument black, pronotum and elytra rufo-testaceous,

appendages brownish-black. Head moderately elongate, distance from anterior

margins of antennal sockets to apex of clypeus shorter than width of face across

genae; widest anterior portion of lower lobe of eye extending slightly beneath

antennal insertions; vertex minutely, densely puncate, with coarse, confluent

punctures superimposed, pubescence more or less transversely appressed, not

obscuring surface; frons concave, with a median longitudinal groove, surface

subopaque, densely, variolately, confluently punctate, sparsely clothed with

appressed pubescence; clypeus minutely, densely punctate with coarser punctures

superimposed; antennae extending nearly to apical one-fifth of elytra, first four

and one-half segments shining, moderately finely punctate, with some appressed

coarse black bristles, remaining segments dull, minutely, densely punctate, clothed

with very short, appressed pubescence, scape slightly shorter than third segment,

second segment short, broader than long, third segment longest, fourth segment

shorter than scape but only slightly shorter than fifth, sixth segment slightly

shorter than fifth, subequal to fourth, remaining segments successively shorter to

eleventh, which is distinctly longer than tenth with apex subconical, pale orange.

Pronotum wider than long (1.28:1.00), surface, except midline, densely, moderately

coarsely punctate, thinly clothed with moderately long, appressed, golden pubes-

cence; posterior disc slightly impressed, shining, sparsely punctate; thoracic sterna

shining, finely moderately densely punctate, clothed with moderately long appressed

and suberect golden pubescence which does not obscure the surface. Elytra

about 2.4 times as long as basal width; surface rather densely punctate, punctures

distinctly smaller than those of pronotum; apices rounded to outer angle which

is distinct but at most feebly dentate. Legs with posterior tarsi not quite as

long as tibiae, first segment longer than remaining segments together. Abdomen

with sternites shining, finely, moderately densely punctate, more sparsely so at

middle of first three, pubescence moderately long, appressed and suberect, sparser

medially; last sternite feebly longitudinally impressed at middle over basal one-half,

apex rotundate-iruncate. Length, 11.5 mm.

Holotype male——30 MILES NORTHEAST OF TEHUANTEPEC, Oaxaca, Mexico, 8 July

1955, Derham Giuliani (CAS).

This species differs immediately from C. latescens Bates in coloration

(in C. latescens the basal half of the elytra, the anterior coxae and the

42 THE Pan-Pactric ENTOMOLOGIST

Oaxaca, 8 July 1955 (Derham Giuliani) which extends the range signifi-

cantly. Forty-five additional specimens have been taken at the type

locality, five miles north of Mazatlan and nine miles north of Mazatlan

(J. A. and M. A. Chemsak, E. G. and J. M. Linsley and A. E. and

M. M. Michelbacher) at flowers of Buddleia wrightii and Jatropha

curcas. In all of these, the Lycus-like elytral pattern is remarkably

constant.

Nemognathomimus, new species

Body small, narrow, elongate; elytra not attenuated, sides at most very gradually

narrowed from behind humeri to apical one-sixth, then gradually rounded to

apices. Head moderately elongate, distance from anterior margin of antennal

sockets to apex of clypeus a little shorter than width of face across genae (1.0:1.1) ;

eyes emarginate; antennal tubercles adjacent to upper inner edge of lower lobe

of eyes, insertions not embraced by eyes; antennae with segments one to five

shining, black, coarsely punctate, clothed with coarse black bristles, segments six

to eleven dull, dark brown, minutely punctate, clothed with short, fine pubescence,

segments, three to five cylindrical, third segment longer than fifth, fifth longer

than fourth, segments six to eleven thickened but not subserrate. Pronotum

narrowly campanuliform, not quite as long as basal width, surface polished, very

sparsely and finely punctate; prosternum with intercoxal process very narrow,

laminiform, anterior coxae prominent, exserted, cavities closed behind; meso-

sternum depressed between coxae and arcuately but somewhat flatly declivous

in front, intercoxal process at middle of coxae distinctly narrower than vertical

diameter of base of middle femora. Legs slender; posterior tarsi filiform, longer

than tibiae in the male, a little shorter than tibiae in the female, first segment

longer than remainder of tarsus, third segment deeply incised, lobes elongate.

Elytra with apices obliquely rotundate-truncate. Abdomen with last sternite

emarginate at apex and feebly triangularly impressed with the lateral angles feebly

produced in the male, the apex entire and rotundate-truncate in the female.

Type species.—Choriolaus pallidulus Linsley

This genus differs from the related genera discussed here in the

elongate, slender form, the structure of the antennae, and the highly

polished, scarcely punctate, sparsely pubescent pronotum.

Nemognathomimus pallidulus (Linstey), New combination

Choriolaus pallidulus Linsley, 1935, Trans. American Ent. Soc., 61:82; Linsley

and Chemsak, 1971, Arquivos de Zoologia, 21:18 (synonymy).

Choriolaus pubicollis Linsley, 1935, Trans. American Ent, Soc., 61:83.

This species occurs in at least four distinctive color forms (1) a

typical Nemognatha-like coloration, being pale testaceous with the

eyes, antennae, tibiae and tarsi black, (2) similar but with elytral apices

narrowly black, (3) black, with the prothorax rufo-testaceous, a color-

VoL. 52, No. 1, January 1976 Al

femora are golden yellow), and the feebly dentate lateral apical angles

of the elytra. From C. celestae (Chemsak and Linsley) and C. gracilis

(Chemsak and Linsley), C. derhami may be distinguished by having

only the pronotum and elytra reddish.

CHORIOLAUS GRACILIS (CHEMSAK AND LINSLEY)

Megachoriolaus gracilis Chemsak and Linsley, 1974, Pan-Pacific Ent., 50:39.

Since this species was described based upon three male specimens,

five additional males and three females have been received from Dr.

Paul A. Opler, collected at the type locality in late May and early June,

1973 and 1974, mostly at flowers of Caesaria nitida. The females are

a little larger and more robust than the males, with the elytra less than

twice as long as width behind humeri (nearly 2.2 times as long in males)

and the last sternite broadly subtruncate (narrowly rounded in the

male). The coloration in the entire series is remarkably constant.

GENUS MEGACHORIOLAUS LINSLEY

Megachoriolaus Linsley, 1970, Pan-Pacific Ent., 46:123; Linsley and Chemsak,

1971, Arquivos de Zoologia, 21:3; Chemsak and Linsley, 1974, Pan-Pacitic

Entomol., 50:35.

This genus is closely related to Choriolaus Bates, but differs in having

the anterior coxal cavities open behind, the intercoxal process of the

mesosternum prominent, not recessed, abruptly arcuate or subvertically

declivous in front and at midpoint between coxae as wide as or wider

than vertical diameter of base of middle leg, the posterior tarsi distinctly

shorter than the tibiae, and the antennae with the outer segments

distinctly thickened or subserrate with the basal segments usually abbre-

viated and clothed with coarse black setae. As redefined, the following

species are included: M. breviceps (Linsley) (Arizona), M. chemsaki

Linsley (Sinaloa to Oaxaca), M. cruentus (Martin) (Arizona), M. flam-

matus Linsley (Morelos), M. ignitus (Schaeffer) (Arizona), M. imita-

trix Linsley (San Luis Potosi), M. lineaticollis Chemsak and Linsley

(Panama), M. patricia (Bates) (Vera Cruz to Panama), M. spiniferus

(Linsley) (Panama), M. texanus (Knull) (Texas), M. unicolor (Bates)

(Guerrero), and M. nigricollis Chemsak and Linsley (Mexico).

MEGACHORIOLAUS CHEMSAKI LINSLEY

Megachoriolaus chemsaki Linsley, 1970, Pan-Pacific Ent., 46:129.

Since this species was described from near Mazatlan, Sinaloa; we

have received specimens from 30 miles northeast of Tehuantepec,

Vou. 52, No. 1, January 1976 43

ational type suggestive of Nemognatha zonitoides Dugés, and (4) black

with the elytra metallic greenish, as in some forms of NV. caeruletpennis

Perty (see Champion, 1891-93).

Material studied by us represents these color phases as follows:

MEXICO: Sinaloa: Mazatlan, 5 and 9 miles north, at flowers of

Buddleia wrightii and Jatropha curcas, elytra yellow, 30, elytra black,

39, Elota, 8 mi. S., elytra black, 1; Nayarit: Arroyo Santiago, near

Jesus Maria, elytra yellow, 3, elytra black, 1; 3 mi. northwest of Santa

Maria del Oro, elytra yellow with apices black, 1; Mexico: Temescal-

tepec, elytra yellow, 1, Bejucos, elytra black; Oaxaca: 30 miles northeast

of Tehuantepec, elytra yellow, 3, elytra black, 4; Chiapas: 17 kilometers

north of Tuxtla Gutiérrez, elytra yellow, 1; Yucatan: Chichen Itza,

elytra yellow, 1. EL SALVADOR: Quezaltepeque, elytra metallic

greenish, 1.

Mordellistenomimus, new genus

Body small, oblong; elytra not attenuated, sides subparallel to apical one-third,

then gradually rounded to apices. Head with muzzle short, broad; eyes emarginate;

antennal tubercles adjacent to upper inner edge of lower lobe of eyes, insertions

not embraced by eyes; antennae filiform, outer segments not swollen, second

segment cylindrical, third segment longer than scape, fourth segment distinctly

shorter than third, fifth segment longer than third and sixth. Pronotum broadly

campanuliform, narrowed gradually from base to middle, then broadly rounded to

apex; surface densely punctate, the punctures larger than those of vertex of head,

smaller than those of elytra. Prosternum with intercoxal process narrow, lamini-

form, coxae prominent, exserted, cavities closed behind, Elytra rugoso-punctate,

not costate; apices separately rounded.

Type species.—Choriolaus nanus Bates

This genus may be distinguished from other genera in the Choriolaus-

Euryptera group by the filiform antennae with the outer segments not

enlarged or subserrate, the fifth segment longer than the third and the

sixth, the pronotum broadly, rather than narrowly campanulate, and

the elytral surface rugoso-punctate. The following is the only known

species.

MOoRDELLISTENOMIMUS NANUS (BATES), NEW COMBINATION

Choriolaus nanus Bates, 1885, Biologia Centrali-Americana, Coleoptera, 5:284,

pl. 20, fig. 8 (Q); Linsley and Chemsak, 1971, Arquivos de Zoologia, 21:17 (2).

Type locality—Mexico.

We have seen only the type specimen in the British Museum (Natural

History), London. The head, except the eyes and antennae, the pro-

thorax, and the femora, except the apices, are yellow; the elytra are

4A, THE Pan-PactFic ENTOMOLOGIST

black; and the mesosternum, metasternum and abdomen brownish. The

pubescence is golden on the yellow areas of the integument, otherwise

black.

As remarked by Bates, this species resembles a mordelled beetle of the

genus Mordellistenus. Among the Mexican species figured by Champion

(1891-1893), it is suggestive of M. atripennis Champion or M. rubri-

collis Champion.

GENUS CHONTALIA BATES

Chontalia Bates, 1872, Trans. Ent. Soc. London, 1872: 233; Bates, 1885, Biologia

Centrali-Americana, Coleoptera, 5:283; Linsley and Chemsak, 1971, Arquivos

de Zoologia, 21:22.

Chontalia differs from the other genera of small Neotropical lepturines

with the posterior basal angles of the pronotum extending over the

elytral humeri by having the antennal insertions within the ocular

emargination and partially embraced by the eyes, and the antennae with

segments three, four and five becoming successively longer.

CHONTALIA CYANICOLOR BATES

Chontalia cyanicolor Bates, 1872, Trans. Ent. Soc. London, 1872: 233; Bates, 1885,

Biologia Centrali-Americana, Coleoptera, 5:40, pl. 5, fig. 3; Linsley and Chemsak,

1971, Arquivos de Zoologia, 21:23.

We have seen only the type specimen of this species, a female in the

collection of the British Museum (Natural History), London. The

integument is black, with the head, prothorax and episterna cyaneus

and the basal half of the elytra yellow. The type has been illustrated by

Bates (loc. cit.).

Lycidocerus, new genus

Body small, elongate; elytra with sides straight, parallel-sided in male, gradually

widened to apical one-sixth in female, then rounded to apices. Head moderately

elongate, distance from a line connecting anterior margin of antennal insertions

to apex of clypeus only slightly less than width of faces across genae; temples not

prominent, narrowed behind eyes to neck, which is distinct; antennal insertions

below ocular emargination, adjacent to upper inner edge of lower lobe of eye;

antennae stout, segments broad, flattened, three to five densely clothed with coarse,

suberect black setae in male, three to six in female, third segment shorter than

scape, fifth shorter than third, longer than fourth, eleventh segment appendiculate.

Pronotum distinctly shorter than basal width, narrowly campanuliform, sides feebly

rounded at middle, basal angles extending over humeri, punctation on disk finer

than that on base of elytra; prosternum with intercoxal process very narrow,

laminiform, coxal cavities open behind; mesosternum with intercoxal process

VoL. 52, No. 1, January 1976 45

deeply recessed, gradually arcuate in front, narrow between the coxae where at

midpoint it is distinctly narrower than tarsal segments. Legs with posterior tarsi

a little shorter than the tibiae in the male, distinctly so in the female. Abdomen

with last sternite narrowly emarginate at apex in male, broadly truncate in

female.

Type of genus.—Lycidocerus sanguineus new species.

This genus is apparently related to Lycochoriolaus but differs

from that genus and others in the Choriolaus group in the much more

abbreviated basal segments of the antennal flagellum, with the third

segment distinctly shorter than the scape.

Lycidocerus sanguineus, new species

(Figure 1)

Male: body narrow, predominantly black; head, except apex of clypeus, labrum,

ocular emargination, antennal tubercles, vertex, and mid-line on neck rufo-testa-

ceous; pronotum, except for median longitudinal black line which continues on to

neck, rufo-testaceous; elytra, except for sutural margin which is narrowly black

over basal two-thirds in male, basal one-half in female, rufo-testaceous; anterior

and intermediate coxae rufo-piceous, base of anterior and intermediate femora base

of first segment of tarsus, apex of last segment and claws testaceous or rufo-testa-

ceous; pubescence black in dark areas, golden in pale areas. Head with vertex and

frons above antennal sockets, including ocular emargination finely punctate and

more or less transversely rugose; lower frons dull, minutely, densely punctate;

clypeus, labrum and genae shining, minutely, sparsely punctate; ventral surface

shining, area adjacent to eyes rugulose; antennae reaching to apical one-fourth

of elytra. Pronotum minutely densely punctate; prosternum shining, finely, very

sparsely punctate and pubescent; metosternum finely punctured and pubescent,

coarsely punctate at side and on episterna. Legs with femora moderately coarsely

punctate, tibiae finely punctate, tarsi minutely punctate. Elytra with surface

granulate, more coarsely punctate than pronotum, clothed with depressed golden

pubescence which does not obscure the surface; apices rounded to outer angle

which is feebly dentiform. Abdomen shining; sternites finely sparsely punctate,

thinly clothed with depressed pale hairs; last sternite narrowly emarginate at

apex. Length, 9.5 mm.

Female: form more robust; antennae not quite attaining apical one-third of

elytra; elytra with narrow black sutural line extending from scutellum to nearly

middle; abdomen broadly truncate at apex. Length, 9 mm.

Holotype male and allotype (Los Angeles Museum of Natural History ).—five

miles south of San Vito de Java, Puntarenas Province, Costa Rica (R. W.

McDiarmid).

This species somewhat resembles a Lycochoriolaus in general form,

but in addition to the generic characters it differs from the described

bicolored species in that genus in the dark red, rather than testaceous

or golden, elytra.

46

Fic.

\P

THE Pan-Paciric ENTOMOLOGIST

Lycidocerus

sanguineus Chemsak and Linsley, allotype @.

VoL. 52, No. 1, January 1976 47

Leptochoriolaus, new genus

Body moderate-sized, elongate, opaque; elytra narrowed from behind humeri

over basel one-third, expanded before narrowing then parallel-sided to apical

one-sixth, then narrowed to apices; apices rounded internally, external angle

feebly dentiform. Head elongate, distance from anterior margin of antennal

sockets to apex of clypeus distinctly longer than width of face across genae;

antennal tubercles adjacent to upper inner edge of lower lobe of eye but outside

of ocular emargination, widest part of lower lobe not extending beneath them;

clypeus somewhat prominent; antennae not filiform, outer segments thickened but

not quite twice as wide as apex of third segment, basal segments with some coarse

short, depressed bristles, third segment twice as long as fourth, one-third longer

than fifth, outer segments with small apical poriferous areas. Pronotum campanu-

liform, shorter than basal width, apical constriction narrow, basal angles acute

extending over elytral humeri. Prosternum with intercoxal process very narrow,

laminiform; anterior coxae prominent, exserted; mesosternum with intercoxal

process moderately narrow, recessed arcuately but somewhat flatly declivous in

front. Legs slender, posterior tarsi filiform, distinctly shorter than tibiae, first

segment longer than remainder of tarsus, third segment deeply incised, lobes

elongate.

Type species.—Leptochoriolaus opacus new species

This genus differs from others in the Euryptera-Choriolaus complex

in the very elongate elytra which are narrowed behind the humeri and

then subparallel nearly to the apices and the opaque integument, and in

the form and proportion of the antennal segments.

Leptochoriolaus opacus, new species

(Figure 2)

Female(?): Integument opaque, head and appendages black, neck, pronotum,

prosternum and elytra at base rufo-testaceous, remainder of elytra, metasternum

and abdomen brown. Head with vertex and upper frons between eyes concave,

very finely, very densely punctate, with a median, longitudinal polished line;

face and clypeus finely, very densely punctate, punctures of clypeus a little larger

than those of lower face; antennal tubercles elevated; antennae extending over

basal half of elytra, scape elongate, subcylindrical, distinctly longer than vertical

diameter of eye, second segment about as long as broad, third segment narrow over

basal one-half, subequal in length to scape, fourth segment one-half as long as

third, subequal to sixth, fifth segment one-third shorter than third, one-third

longer than fourth and sixth, segments six to ten gradually decreasing in length,

eleventh a little longer than tenth, apex acute. Pronotum finely, very densely

punctate, clothed with very short, golden pubescence which does not obscure

surface, midline longitudinally impressed behind middle and in front of base;

prosternum and metasternum finely, very densely punctate. Legs with femora

finely punctate but somewhat shining. Elytra moderately coarsely punctate over

a very densely, minutely punctate surface, clothed with short depressed golden

pubescence which does not obscure surface. Abdomen yery finely, densely

AS THe Pan-PaciFic ENTOMOLOGIST

Fic. 2. Leptochoriolaus

opacus

Chemsak and Linsley, holotype

Tees

VoL. 52, No. 1, JAnuary 1976 49

punctate, clothed with short depressed pale pubescence which does not obscure

surface. Length, 10 mm.

Holotype female(?) (United States National Museum of Natural History) —

Turrialba, Costa Rica (E. A. Bottcher) (Tippmann Coll. ’57, 213112).

The aspect of this species is unlike that of any other described species

of Central American lepturine.

LITERATURE CITED

Bates, H. W. 1872. On the longicorn Coleoptera of Chontales, Nicaragua.

Trans. Entomol. Soc. London, 1872: 163-238.

1879-85. Longicornia. Biologia Centrali-Americana, Coleoptera, 5: 1-436.

pls. 1-25.

Boprr, P. L. 1921. Genera insectorum. 178. 119 pp., 8 pls.

Cuampion, G. C. 1890-91. Mordellidae. Biologia Centrali-Americana, Coleop-

tera, 4(2): 250-350.

1891-93. Meloidae. Biologia Centrali-Americana, Coleoptera, 4(2): 364-450.

Cuemsak, J. A. 1964. Lectotype designations of Cerambycidae in the British

Museum (Natural History). J. Kans. Entomol. Soc., 40: 73-81.

Cuemsak, J. A. AND E. G. Linstey. 1974. Notes and descriptions of some Lycid-

like Neotropical Lepturine Cerambycidae. Pan-Pac. Entomol., 50:35-43,

2 figs.

Linstey, E. G. 1935. Studies in the Longicornia of Mexico. Trans. Amer.

Entomol. Soc., 61: 61-102.

1961. The North and Central American species of Euryptera and a related

new genus. J. New York Entomol. Soc. 69: 131-141, figs.

1970. New genera and species in the lepturine complex related to Euryptera

and Choriolaus. Pan-Pac. Entomol., 46: 128-141, 4 figs.

Linstey, E. G. anp J. A. CHemsAk. 1971. An attempt to clarify the generic

status of some Neotropical species currently assigned to Furyptera,

Chontalia and Ophistomis (Coleoptera, Cerambycidae). Arquivos de

Entomol. 21: 1-40.

Redescription and Phylogenetic Affinities of

Kytorhinus prolixus (Fall)

(Coleoptera: Bruchidae: Kytorhininae)

CLARENCE DAN JOHNSON

Department of Biological Sciences, Northern Arizona University, Flagstaff, Az 86001

_Kytorhinus prolixus (Fall) was described in 1926 and to this date

it is the only species in the subfamily Kytorhininae known to occur

in the New World (Decelle, 1971). There are about 15 species of

Kytorhinines in the Old World, the subfamily having a Holarctic distri-

bution. This is an aberrant group of Bruchidae, not only in morphology

but in its northern distribution. Most species of bruchids occur in the

tropical and semitropical areas of the world.

Only Luk’yanovich and Ter-Minasyan (1957) and Decelle (1971)

have dealt in any detail with Kytorhinus since Bridwell (1932) placed

prolixus in Kytorhinus and named the subfamily Kytorhininae. Luk’

yanovich and Ter-Minasyan provided a key to 8 species and Decelle re-

viewed the literature on Kytorhinus and described 3 new species.

Because K. prolixus has never been completely redescribed, had its

distribution delimited, or had its host range discussed, these subjects

are presented here. Hopefully this will stimulate further studies on the

taxonomy, ecology, and behavior of Kytorhinus.

KYTORHINUS PROLIXUS (FALL)

(Figs. 1-8)

Mylabris (Bruchus) prolixus Fall, 1926:204 (near McKinley Park Station, Alaska).

Kytorhinus prolixus: Bridwell, 1932:101; Bridwell, 1946:53; Blackwelder and

Blackwelder, 1948:45; Brown, 1952:342; Bottimer, 1961:293; Johnson, 1968:1269;

Bottimer, 1968:1036, 1039; Decelle, 1971:106.

Kytorrhinus (sic) prolixus: Teran, 1967:307, 311, 319, 335.

Length (pronotum-—elytra) 1.9-2.7 mm. Width 1.0-1.4 mm. Maximum thoracic

depth 0.9-1.4 mm. ;

Mate.—Head and body dark brown to black, antennae and legs brown to black;

body slender, elongate with recumbent, uniform sparse white hairs; sometimes with

scattered golden brown hairs; scutellum with short, dense white hairs.

Head.— (fig. 4). Short, narrow, densely punciulate; frons usually with a thin line

extending from frontoclypeal suture to vertex; head usually with bulge between

dorsal margins of eyes followed by a vague transverse sulcus between upper limits

of eyes; eye large, rounded, 2.2 to 3.5 times as wide as frons; ocular sinus long,

dividing dorsal surface of eye for about 0.8 its length; posterior margin of eye

protruding from adjacent surfaces; postocular lobe broader near dorsum of eye,

Tue Pan-PaciFic ENTOMOLOGIST 52: 50-55. JANUARY 1976

VoL. 52, No. 1, January 1976 ol

Fics. 1-5. Kytorhinus prolixus. 1. Antenna of male. 2. Antenna of female.

3. Dorsal aspect. 4. Head. 5. Hind leg.

tapering toward venter, rounded, not angulate; muzzle short, distance from venter

of eye to apex of labrum ¥% to ¥4 as long as distance from upper limits of eyes

to apex of labrum; clypeus with large, shallow depression, lateral margins with

carinae extending from apex of clypeus to base of antennae; antennal segment 1

about 0.9 as long as length of eye, segment 2 short, about 0.2 as long as segment

1, segments 3-10 pectinate, segment 11 slender, about as long as ramus of segment

2 THE PAN-PAciFIC ENTOMOLOGIST

yy a

HH naval

ju MIAN

Fics. 6-8. Kytorhinus prolixus male genitalia. 6. Median lobe, ventral view.

7. Lateral lobes, ventral view. 8. Median lobe and lateral lobes, lateral view.

10; ramus of segment 3 about 0.5 as long as segment, ramus of segment 4 about

1.5 times as long as segment, rami of segments 5-10 about twice as long as seg-

ments, rami longest on segments 6-10 (fig. 1); antenna long, reaching to slightly

beyond apex of abdomen.

Prothorax.—Disk subcampanulate (fig. 3), with many punctations in no ap-

parent pattern; cervical sulcus deep, extending from near coxal cavity almost to

midline of pronotum; lateral prothoracic carina evident only at base, often with

blunt ridge extending to coxal cavity; short median impressed line on median

basal lobe; prosternum separating procoxae for about 0.5 their length. .

Mesothorax and Metathorax.—Scutellum elongate, almost 2 times longer than

broad, bifid at apex, clothed with dense recumbent white hairs; elytron almost

3 times longer than broad; striae shallow, punctulate, strial intervals rugulose;

striae 3 and 4 closer to one another at base than to adjacent striae, others subequal

at base; humerus rugulose, swollen, extending obliquely toward midline, ter-

minating between striae 6 and 7, sometimes with small, round pit near center,

VoL. 52, No. 1, January 1976 53

glabrous; undersurfaces punctate; about 0.5 of lateral portion of hind coxa punctate,

medial 0.5 smooth, without punctations; hind femur long, narrow, constricted

basally and apically, expanded only slightly medially to about 0.8 width of coxa

(fig. 5); vague longitudinal carina on inner ventral margin, femur otherwise un-

marked; femur extending to beyond apex of abdomen; tibia unmarked, single short,

broad flat spine at lateroventral margin of apex, without mucro; Ist tarsomere with

only ventral glabrous carina.

Abdomen.—I|st sternum about 0.3 as long as abdomen, posterior margin straight,

sterna 2-4 unmodified, 5th emarginate to receive apex of pygidium, produced

ventrally at apex; pygidium small, punctate, convex in lateral view; exposed

tergite unmarked.

Genitalia—(Figs. 6-8). Median lobe moderate in length, tubular, slightly

flattened, without dorsal hood at apex; ventral valve elongate, pointed, sides con-

cave, base about 0.5 as wide as apex of median lobe, arcuate and perpendicular to

median lobe in lateral view; without hinge sclerites; armature of internal sac

consisting of 2 pairs of clumps of spinules basally, 1 dorsal, 1 ventral, and a

clump of large, heavily sclerotized structures medially, medial cluster composed

of 2 pairs of curved, pointed structures laterally, a pair of elongated spines slightly

medial to curved structures, and 2 elongated structures along midline, dorsal struc-

ture slightly curved, pointed, ventral structure flattened dorsoventrally at both

base and apex. Lateral lobes expanded at apex, gradually constricted to about 0.3

from apex, only slightly cleft at apex (fig. 7). (See also Teran, 1967, p. 311).

FEMALE.—Similar to male except eyes smaller, about 1.4 times as wide as width

of frons; antennal segment 1 (fig. 2) about as long as length of eye, segment

2 about 0.3 as long as 1, segments 3-10 serrate, 11 acuminate at apex; antenna

extending to about 0.66 the length of elytron; abdominal sternum 1 about 0.3

as long as abdomen, segments 2-5 unmodified, 5 gently rounded, not emarginate;

abdomen longer, extending well beyond apex of hind femur; not bent downward

at apex.

Host Plants.-—Unknown.

Types.—Museum of Comparative Zoology, Harvard University. The male speci-

men bearing the following labels is here designated Lectotype for this species:

“McKin. Pk. Alaska, 7-10-24, TYPE prolixus, M. C. Z. Type 25059, LECTOTYPE,

Mylabris (Bruchus) prolixus Fall By C. D. Johnson 1976.” Three female speci-

mens bearing the labels “McKin. Pk. Alaska, 7-9-24” which make up the remainder

of the type-series of Fall are paralectotypes.

Specimens examined.—l04, from the following localities, all June, July:

Manitoba. 30 miles N. Roblin. Saskatchewan. Lake Madge. Alberta. McMurray,

on Hedysarum americanum; Drumheiler; Grande Prairie; Elkwater Park; Kanana-

skis; Calgary; Pt. Saskatchewan; Cypress Hills; Edmonton. British Columbia.

Atlin; Pouce Coupe; Summit Lake, mi 392 Alaska Hwy., 4700’. Northwest Terri-

tories. Reindeer Depot; Mackenzie Delta, on Hedysarum alpinum americanum;

Aklavik; Norman Wells; Wrigley. Yukon Territory. Ross River, 132°30’, 61°56’,

3000’. Alaska. Ft. Wainwright; near McKinley Park Station.

Distribution.—South Dakota. Manitoba. Saskatchewan. Alberta. British Colum-

bia. Northwest Territories. Yukon Territory. Alaska.

Discussion.—F all (1926) described this unique species from a series

of 4 specimens that he collected near McKinley Park Station, Alaska.

54, THE Pan-Pactric ENTOMOLOGIST

Since that time it has been found to occur also in western and central

Canada and South Dakota. Intensive collecting will probably show

that it has a wider distribution in the United States because L. J. Bot-

timer found an egg that is probably prolixus on a pod of Hedysarum

from Idaho in a herbarium.

About 16 species of Kytorhininae and its only genus Kytorhinus have

been described. All species whose hosts are known feed in the Legu-

minosae. The Old World species K. pectinicornis Melichar feeds in the

seeds of Hedysarum and, according to published descriptions, closely

resembles K. prolixus.

Apparently Kytorhinus had an Old World Origin because it is most

abundant there. The present distribution of prolixus does not suggest

a recent introduction into the New World.

In this paper the external morphology and male genitalia have been

described for K. prolixus. Not only will this allow easier identification

of the species but will allow other workers to more easily compare

prolixus with Old World species. The subfamily characters listed by

Bridwell (1946) separate K. prolixus from all other New World

Bruchidae.

The male genitalia of K. prolixus were figured by Teran (1967) and

are also figured here. The shape of the median lobe and ventral valve

resemble some New World Bruchinae but the lateral lobes and armature

of the internal sac are unique. The mass of elongated, curved structures

in the internal sac more closely resembles members of the Amblycerinae.

Although K. prolixus has yet to be reared from the seeds of a host,

it has been collected by sweeping on numerous occasions from species

of Hedysarum (Leguminosae), its probable host. Dr. A. G. Raske col-

lected seeds of Hedysarum sp. from Calgary, Alberta, Canada, in Septem-

ber 1969 but no bruchids were reared from these seeds. I collected seeds

of Hedysarum boreale Nuttall, our rare Arizona boreal species, from

4. miles NE Lukachukai, Chuska Mountains, Apache County on 28 July

1975. There is no evidence of bruchid eggs or exit holes and to this

date no bruchids have emerged from these seeds. The plants were also

thoroughly swept and no bruchids were collected from them. In addi-

tion, I have examined several herbarium specimens of H. boreale from

other localities in Arizona and I have found no bruchid eggs or damage.

Apparently, then, K. prolixus does not occur as far south as Arizona.

ACKNOWLEDGMENTS

I wish to thank the following persons for the generous loan of speci-

mens for this study: G. E. Ball, University of Alberta; E. C. Becker and

VoL. 52, No. 1, January 1976 90

D. E. Bright, Canadian National Collection of Insects, Ottawa; J. F.

Lawrence and Jan Scott, Museum of Comparative Zoology, Harvard

University; H. B. Leech, California Academy of Sciences, San Francisco;

and J. W. Smith, University of Missouri, Columbia.

I am also grateful to A. G. Raske for collecting Hedysarum seeds

for me in Canada.

Partial financial support for this study was provided by the Systematic

Entomology Laboratory, IIBIII, Agriculture Research Division, U.S.

Department of Agriculture under Grant 12-14-100-9970 (33).

LITERATURE CITED

BLACKWELDER, R. E. anp BLAcKWELDER, R. M. 1948. Fifth supplement to the

Leng Catalogue of Coleoptera of America, north of Mexico. John D.

Sherman, Jr., Mt. Vernon, N.Y. 87pp.

Bottimer, L. J. 1961. New United States records in the Bruchidae, with notes

on host plants and rearing procedures (Coleoptera). Ann. Entomol. Soc.

Amer., 54: 291-298.

Bottimer, L. J. 1968. Notes on Bruchidae of America North of Mexico with a

list of world genera. Can. Entomol., 100: 1009-1049.

BRIDWELL, J. C. 1932. The subfamilies of the Bruchidae (Coleoptera). Proc.

Entomol. Soc. Wash., 34: 100-106.

BRIDWELL, J. C. 1946. The genera of beetles of the family Bruchidae in America

north of Mexico. Jour. Wash. Acad. Sci. 36: 52-57. .

Brown, W. J. 1952. Some species of Phytophaga. Can. Entomol. 84: 335-342.

DecELte, J. 1971. Trois nouvelles especes de Kytorhinus (Col. Bruchidae) du

Tibet et du Bhutan. Bull. Ann. Soc. R. Entomol. Belg., 107: 105-115.

Fatt, H. C. 1926. A list of Coleoptera taken in Alaska and adjacent parts of

the Yukon Territory in the summer of 1924, Pan-Pac. Entomol., 2 (4) :

191-208.

Jounson, C. D. 1968. Bruchidae type-specimens deposited in United States

museums, with lectotype designations. Ann. Entomol. Soc. Amer., 61:

1266-1272.

Lux’yanovicH, F. K. anp Ter-Minasyan, M. E. 1957. Bruchidae, Zool. Inst.

Akad. Nauk SSSR, n.s. No. 67. Fauna SSSR, t. XXIV, n. 1. 208pp.

Terran, A. L. 1967. Observaciones sobre las estructuras genitales de los machos

de diversos generos de Bruchidae (Coleoptera). Acta Zool. Lilloana,

22: 307-336.

Giulianium campbelli, a New Genus and Species

of Marine Beetle From California

(Coleoptera: Staphylinidae)

Ian Moore

Staff Research Associate, Division of Biological Control,

University of California, Riverside, California 92502

The coleopterous fauna of the seashore of Pacific North America

contains numerous staphylinids, including intertidal species, as well as

those inhabiting wrack and other supratidal situations. The unusual

species considered here is known from a single beach locality in northern

California. It is described in order to make the name available for future

treatment of the maritime Staphylinidae.

Giulianium new genus

Form.—Small, parallel, not robust. Integuments finely densely reticulate, shining

and pubescent.

Head.—Orbicular, narrowed behind to a distinct neck, with a very faint

nuchal constriction across the dorsal surface. Eyes small, not at all interrupting

the side margin of the head. Antennae long, not incrassate; all the segments

longer than wide; their fossae located at the front margin of the head above the

bases of the mandibles. Mandibles acute at apex. Labrum transverse, apex

deeply emarginate in an even arc so that the labrum appears to be bilobed.

Maxillary palpus four-segmented; first segment short; second segment a little

longer than wide, narrow at base, widest at apex, slightly arcuate; third segment

as wide as apex of second segment, somewhat shorter than wide; fourth segment

about as long as the second and third together, slightly wider than third segment,

bulbous in basal half, narrowed to apex which is hooked outward and slightly

downward, apex narrowly rounded. Inner lobe of maxilla shorter than outer

lobe, narrow, without teeth on inner edge densely pubescent at apex. Outer lobe

of maxilla slender, inner edge simple, apex densely pubescent. Labial palpus

three-segmented, each segment longer than wide, first segment widest, last segment

about twice as long as wide. Mentum transverse, narrowed to apex. Submentum

horizontal, distinctly defined, rapidly narrowed to a very narrow base which is

separated from the gula by a suture. Gular sutures parallel for about one-fourth

of their length, thence widely divergent to base of head. Infraorbital carina

lacking.

Thorax.—Pronotum subquadrate. Prosternum long, without longitudinal carina,

its process short, acute, extending only a short distance between the coxae.

Metasternum moderate, simple, its process acute, extending a short distance

between the coxae. All the coxae large exserted, contiguous. Tibiae without

spines. Tarsi 5-segmented, first 4 segments short, subequal, fifth segment about

as long as the 3 preceding together.

Tue Pan-Pactric ENTOMOLOGIST 52: 56-59. JANUARY 1976

VoL. 52, No. 1, JANUARY 1976 o7

Table 1. Tabular key to the Nearctic genera of the Phloeocharinae

(for construction and use of tabular keys, see Newell, 1970, 1972).

Characters are as follows: 1. Shape of head; PROD = produced as a

beak which conceals labrum; ARC = arcuate, not produced as a beak.

2. Anterior margin of labrum; EMARG = deeply emarginate; ARC =

arcuate; CONC = concealed by beak. 3. Orientation of mentum; VERT

= vertical; HORIZ = horizontal, flat; IMPR = horizontal, impressed.

4. Lateral margin of pronotum; PRES = prominent; OBSOL = obsolete.

9. Trochantin size; LGE = large, conspicuous; SMALL = minute, in-

conspicuous. 6. Separation of middle coxae; CONT = contiguous;

SEP = separated by a long process of the mesosternum.

a ie 2. 3, 4, a 6.

Genus

Rimulincola Sanderson ARC ARC IMPR PRES - LGE SEP

Vicelva Moore & Legner PROD CONC VERT PRES SMALL CONT

Ecbletus Sharp ARC ARC HORIZ PRES SMALL CONT

ARC EMARG HORIZ OBSOL LGE CONT

Giulianium new genus

Abdomen.—First 5 visible segments with paratergites. Fifth tergite a little

longer than preceding. Sixth visible tergite produced in the central third of

its anterior margin in a short broadly truncate lobe which is normally hidden

beneath the apex of the preceding tergite. Tergites not impressed at base.

Second sternite absent. First visible sternite without a keel between the coxae.

Sternites not constricted at base.

Type species.—Giulianium campbelli new species.

This genus is named in honor of its collector, Derham Guiliani. The generic

name is masculine.

This genus is placed tentatively in Phloeocharinae, a small subfamily

which contains a few genera with poorly displayed affinities. Giulianium

is unusual in the form of the fourth segment of the maxillary palpus, the

distinct submentum and the poorly defined hypomera. Other characters

are compared in Table 1.

Giulianium goes to couplet No. 24 in the key to the subfamilies

(Moore and Legner, 1975) where it fits neither half of the couplet. To

include it the key may be altered as follows:

24a. Superior lateral line of pronotum rudimentary, discernible only at

i se Mee Sorte, PA ls eee ag | Giulianium (Phloeocharinae)

Giulianium campbelli, new species

Color.—Piceous throughout.

Head.—Slightly wider than long; surface flattened, slightly depressed on disc,

58 THE Pan-PActFIcC ENTOMOLOGIST

ul

3

Fic. 1-3. Giulianium campbelli. 1. Dorsal aspect of beetle. 2. Maxillary

palpus. 3. Aedeagus.

densely reticulate, moderately pubescent; sculpture of under surface similar to

that above. Eyes small, of about six or eight facets in diameter. Tempora about

twice as long as eye, arcuate. Antenna almost reaching to apex of elytra, slender,

all segments longer, than wide; first segment longest, almost three times as long

as wide; following segments progressively shorter through tenth segment which

is less than twice as long as wide; eleventh segment about twice as long as wide,

pointed at apex.

Thorax.—Pronotum about as wide as head, about one-seventh longer than wide,

widest near apical fifth; base and apex almost straight; sides gently arcuate,

somewhat narrowed to base; angles narrowly rounded; disc flattened and slightly

depressed; sculpture and pubescence much as on head. Elytra conjointly slightly

wider and about as long as pronotum; basal angles narrowly rounded; sides

gently arcuate; outer apical angles rounded into the arcuate apex; inner apical

VoL. 52, No. 1, January 1976 59

angles broadly rounded. Sculpture and pubescence much as on head and

pronotum.

Abdomen.—Parallel. Sculpture and pubescence similar to that of thorax.

Length—3.0 mm.

Holotype male—Bear Harbor, Humbolt County, California, June 1964, Derham

Giuliani, collector. Taken on the beach. Deposited in the collection of the

Entomology Research Institute, Ottawa, Canada. Two paratypes, sex undetermined,

same data as holotype, one in the Entomology Research Institute, Ottawa, Canada

and the other in the collection of the University of California, Riverside.

The two paratypes are dark ferrugineous with the abdomen parily

piceous. They may be teneral. This species is named for J. Milton

Campbell to whom I am indebted for many favors.

LITERATURE CITED

Moorg, [an anp Lecner, E. F. 1975. Keys to the genera of the Staphylinidae of

America north of Mexico exclusive of the Aleocharinae (Coleoptera:

Staphylinidae). Hilgardia 42: 548-563.

Newe.t, I. M. 1970. Construction and use of tabular keys. Pac. Insects

12: 25-37, 3 figs.

1972. Tabular keys, further notes on their construction and use. Trans.

Conn. Acad. Arts Sci. 44: 259-267, 4 figs.

RECENT LITERATURE

AMERICAN MUSEUM OF NATURAL HISTORY (Central Park West at 79th

Street, New York, New York 10024)

BULLETIN:

Vol. 154, art. 1, The eyeless beetles of the genus Arianops Brendel (Coleoptera,

Pselaphidae), by Thomas C. Barr, Jr., 51 pp., 1974. Price $2.25.

Vol. 155, art. 1, A revision of the spider genus Gnaphosa (Araneae, Gnaphosidae)

in America, by Norman I. Platnick and Mohammad U. Shadab, 66 pp,

1975. Price $2.35.

Vol. 155, art. 2, A taxonomic study of African allodapine bees (Hymenoptera,

Anthophoridae, Ceratinini), by Charles D. Michener, 140 pp., 1975. Price

$8.55.

Vol. 155, art. 3, Revision and phylogeny of the monogeneric subfamily

Pseudopsinae for the world (Staphylinidae, Coleoptera), by Lee H. Herman,

Jr., 65 pp., 1975. Price $3.40.

A New Species of Callophrys (Mitoura) from Mexico

(Lepidoptera: Lycaenidae)

Kurt JOHNSON

Museum of Natural History, University of Wisconsin, Stevens Point 54481

During a survey of genitalia of Nearctic Callophrys which feed on

Cupressaceae, a single male specimen which apparently represented a

new species, was discovered from the Juniperus flaccida Schlecht. habi-

tats of Chihuahua, Mexico. Further examination showed that this speci-

men lacked forewing scent scales (characteristic, with unique excep-

tions, in males of Callophrys and other Theclinae), differed distinctly

in pattern and coloration, and showed genitalic traits intermediate be-

tween those of the Nearctic Cupressaceae-feeding group [C. (M.)

nelsoni (Boisduval), siva (Edwards) , loki (Skinner), gryneus (Hiibner),

and hesseli (Rawson and Ziegler)| and the Nearctic-Neotropical

Loranthaceae-feeding group [C. (M.) spinetorum (Hewitson) and

johnsont (Skinner) ]. It is described as a new species below.

Callophrys (Mitoura) turkingtoni, New Species

(Fig. 1)

Diacnosis. Compared to all Callophrys (Mitoura): male forewings, androconia

absent; under surface, hindwings, distinct marking, distal end of cell; aedeagus,

caudal end, three distinct pointed tips (not two). Compared to C. (M.) siva

(Edwards), gryneus (Hiibner), and spinetorum (Hewitson): upper surface wing

coloration burnt sienna, not chamois [males, topotypical g. castalis (Edwards) ],

honey yellow [males, Pima Co., Arizona gryneus ssp.], zinc orange [males, topo-

typical s. siva], or dusky dull bluish green [males, all Arizona spinetorum] (all

colors Ridgway, 1912). Under surface, hindwings, burnt sienna to argus brown,

not green (gryneus and siva) or darker brown (spinetorum). Under surface,

hindwings, no postbasal markings (gryneus, some siva); under surface, fore-

wings, no apical marking of cell (spinetorum).

Mate.—Upper surface wing coloration burnt sienna confined to basal submar-

ginal area, both wings; limbal area, hindwings, brighter; under surface of fore-

wings burnt sienna (base to postmedian line), apical area parrot green between

veins; under surface of hindwings burnt sienna to argus brown from base past

mesial line, green marking distal end of cell; mesial line with white and brighter

brown markings nearly continuous; limbal area with blue, orange and black

markings; six marginal black markings between veins caudad from 2a to M; (first,

third: large; second, overshadowed blue; latter three, small); five submarginal

black markings between veins caudad 2°A to Maz; limbal orange marking intense;

area distad along outer angle parrot green between veins. Male Genitalia (Fig. 2)

similar to those of other Callophrys (Mitoura), differing markedly as follows:

distal end aedeagus with three pointed tips; valvae longer, more slender; labides

Tue Pan-Paciric ENToMoLocistT 52: 60-62. JANUARY 1976

VoL. 52, No. 1, January 1976 61

Ficure 1.

more steeply pointed, not gradually angled; two sculptured distal indentions at

suture between vinculum and base of uncullum. Forewing length 13 mm.

FooppLant.—Possibly Juniperus flaccida.

Holotype male, 10 miles east of Namiguipa, CHIHUAHUA, Mexico [west slope,

San de las Tunas Mountains], 3 July 1947 (W. Gertsch, M. Cazier), in collection

of the American Museum of Natural History. Genitalia in vial KJ # 169 at

same institution.

One specimen is known, its distinctness assured by genitalic and

phenotypic comparison of all currently known Nearctic and Neotropical

congeners. The species’ geographic and foodplant associations are

discussed in Johnson, 1975. Other Mexican material, referrable to

C. (M.) siva and gryneus was studied, all from Juniperus deppeana

Steud. habitats, one from only 60 kilometers southwest of the turkingtont

type locality.

I am priviledged to name this species after Fr. William R. D. Turking-

ton, O.H.C., first teacher of biology at the old St. Andrews School,

founded for the children of the Tennessee mountains by the Order of

the Holy Cross.

I am greatly indebted to Dr. Frederick H. Rindge, Curator of

Lepidoptera, American Museum of Natural History, New York, for

advice and facilities for studying Nearctic Callophrys (Mitoura) and

62 THE Pan-PactFic ENTOMOLOGIST

to the Museum of Natural History, University of Wisconsin, Stevens

Point (Dr. Charles A. Long, Director), which supported this work.

LITERATURE CITED

Jounson, K. 1975. Geographic distributions and foodplant diversity in four

Callophrys (Mitoura), Lycaenidae. J. Lepid. Soc. 29 (in press).

Ripcway, R. 1912. Color standards and nomenclature. Washington, D.C. (by

author), iii and 43 pp., 52 pls.

BOOK NOTICE

INSECTS AND OTHER ARTHROPODS OF MeEpiIcaL ImporTANcE—Fdited by Kenneth G.

VY. Smith. British Museum (Natural History), Cromwell Road, London, 1973.

XIV + 561 pp. $15.75.

This present work replaces A Handbook for the Identification of Insects of

Medical Importance by John Smart, first issued in 1943, and useful to medical

entomologists during World War II. The new book has coverage on a world wide

basis of major groups of medically important insects and other arthropods. Each

of the sixteen contributors is a specialist on the taxonomy of the group concerned.

It is intended primarily for identification with keys provided to the following

groups: Diptera (adults and larvae), Culicidae, Simuliidae, Phlebotomidae,

Ceratopogonidae, Tabanidae, Glossinidae, Muscidae, Calliphoridae, Sarcophagidae,

Siphonaptera, Heteroptera, Dictyoptera, and some Arachnida. Arthropods of minor

medical importance are also treated systematically. Special sections on insect

structure, methods of collecting and preserving, classification and nomenclature,

forensic entomology, insects and hygiene, and arthropod vectors (including vector-

pathogen tables and disease distribution maps) are included. Detailed discussions

on biology, ecology and medical aspects, as well as extensive bibliographies are

provided for many groups.—R. W. Merrit, Depariment of Entomology, Michigan

State University, East Lansing, 48824.

Foraging Responses of Veromessor pergandei to Changes

in Seed Production

(Hymenoptera: Formicidae)

STEVEN W. Risstnc! AND JEANETTE WHEELER?

Laboratory of Desert Biology, Desert Research Institute,

University of Nevada System, Reno, Nevada 89507

The desert harvester ant, Veromessor pergandei (Mayr), is a major

component of the ecosystem in the Mohave Desert. In Death Valley

National Monument, nests can be very extensive and long-lived, even

though this is the northern limit of its range (Wheeler and Rissing in

press). In Death Valley V. pergandei has a biomass per unit area

approximately equal to that of the total rodent population in the same

area (Went et al. 1972). Colonies of this species survived in the

Coachella Valley of California even after twelve successive years of

severe drought (Tevis 1958).

The abundance of this species and its ability to withstand long pe-

riods of drought and the consequent scarcity of food must be the result

of an efficient food gathering strategy developed for the desert en-

vironment. We have observed this strategy following a profuse bloom

of winter annuals in 1973 and a significantly less abundant bloom

in 1974.

During the early summer V. pergandei has two foraging periods.

One starts in the early morning often before there is any indication of

sunlight in the sky to the human observer; this period ends when the

temperature becomes too high (apparently). The second period starts

in the afternoon after the temperature has fallen from the noon high

and may continue after dark on warm evenings. By consecutive forag-

ing periods we are referring to any combination of possible time-ad-

jacent foraging periods. However, we omitted numbering the periods

when no foraging activity was observed. At Nest 1 this occurred 7

out of 13 and at Nest 2, 4 out of 13 possible evening foraging periods.

Foraging columns were observed at Nests 1, 2 and 4 during all possible

morning foraging periods.

Materials and Methods: Observations were made on selected nests

at a site in the north end of Death Valley National Monument near

Grapevine Ranger Station at 800 m elevation (see Wheeler and Rissing

1 Present address: Department of Zoology, University of Washington, Seattle, Washington 98195.

2 Supported by National Science Foundation Grants GB-17731X and GB-37241, Dr. Frits W. Went,

Principal Investigator.

Tue Pan-Paciric ENTOMOLOGIST 52: 63-72. JANUARY 1976

64 THe Pan-Pactric ENTOMOLOGIST

in press for full description). The nest numbers and locations are the

same as those described previously; since 1973 Nests 5 and 8 have

been disturbed by road construction, Nest 10 has become inactive, and

Nest 4 (15 m east of Nest 2) has been added to the list of nests studied.

Several properly timed rains which produced abundant germination of

winter annuals preceded the 1973 study period. Later adequate rains

favored plant growth and a profuse setting of seed during the spring

and summer of 1973. In comparison, the rains preceding the spring

and summer of 1974, were less and resulted in poorer germination,

plant growth, and seed production for most plants. For this study,

all evaluations of germination and seed production were very kindly

provided by Dr. F. W. Went from his long-term observations at this

and other sites in Death Valley (personal communication).

Regular observations were made from 19 May to 21 June 1973, and

from 10 June to 23 June 1974. In 1973, the directions of the foraging

columns at four nests were recorded. Periodic samples of the items

foraged at eight nests in the study area were taken by aspirating re-

turning foragers from a smooth surface near the nest entrance at the

height of foraging activity. The items were later separated from the

ants and identified under a stereomicroscope. In 1974, the column

direction and a sample of items foraged were taken each foraging period

at Nests 1 and 2. Observations were begun on the column direction

at Nest 4 on 15 June 1974.

Results: Column descriptions and results of samples of items for-

aged for the 1973 portion of this study are reported elsewhere (Wheeler

and Rissing in press). Figures 3, 4, and 5 show the direction of for-

aging columns at Nests 4, 1, and 2 in 1974. The numbers and percent-

ages of each foraged item are shown in Table 1, which also includes

the cumulative data from the 8 nests observed in 1973.

Discussion: 1. Direction Change And Foraging Method: Studies

have been conducted concerning the foraging pattern of V. pergandei

(Bernstein fide Carroll and Janzen 1973; Went et al: 1972; Wheeler

and Rissing in press). Went e¢ al. reported that foraging column di-

rection changed 15° between consecutive foraging periods. Bernstein

reported a change of approximately 20°, but only when seeds were

scarce; when seeds were abundant she reported random foraging was

preferred to column foraging.

Fig. 1 and 2 show the frequency of degrees of change in column

direction between successive foraging periods in 1973 and 1974 respec-

tively. The data from both years fail to support the observation that

there is some constant by which column direction changes between

VoL. 52, No. 1, January 1976 65

1973 COMBINED DATA (NESTS |, 5, 8, 10)

45 N=222

OBSERVED-1I973

TIMES a°

0 10 =20 30 40 50 60 70 80 90 100 10 120 130 #140 #150 160 i170 180

4° BETWEEN CONSECUTIVE FORAGING COLUMNS

Fic. 1. Histogram showing the frequency of degrees changed between consecu-

tive foraging columns, 1973 data.

successive foraging periods. It appears, however, that in 1973 there

was usually little if any change in column direction between successive

foraging periods and that when this direction did change the amount

was usually small. In 1974 column direction changed more often, and

the amount of change between successive periods was quite variable.

The difference between the two years may be in response to the differ-

ence in seed production in 1973 and 1974. In 1973 seeds were very

abundant and probably uniformly scattered over the foraging area.

In 1974 it may have become necessary to explore all available areas

for local and temporary concentrations of seeds.

In 1973, when seed production was enormous, random foraging was

rarely observed; almost always columns were long and populous. In

1974, when seed production was much less, the column was still the

eas

2 1974 COMBINED DATA (NESTS |, 2, 4)

a 15 N=146

>

a

w

@ 104

oO

°

a

5

w

Ww

=

FO

eT lola ith he iy nie eth) lpi ech, LIS oer Le eRe Ale RIG

0 10 20 30 40 50 60 70 80 90 109 = 110 120 130 140 [50 {i60 I70 180

A° BETWEEN CONSECUTIVE FORAGING COLUMNS

Fic. 2. Histogram for 1974 columns. (See Fig. 1.)

66 THE Pan-Pactric ENTOMOLOGIST

O 5

N

meters

ll, la

34

10 9

14

D.C.

15 “Entrance

14 D.C.

|

: D.C)

6 8

2 4

5

Fic. 3. Observed directions (clockwise from north) and lengths of foraging

columns during consecutive foraging periods at Nest 4. D. C.: double column.

most common foraging pattern; however, infrequent periods of random

foraging were noted, especially at Nest 1. This does not agree with

Bernstein’s report of column foraging only when seeds were more

abundant. It is also interesting that Tevis (1958) noted no shift to

random foraging in 1957, when: seed production was high for the first

time in twelve years. :

While the amount of change between consecutive foraging periods

was variable in 1974, succeeding columns generally rotated about each

individual nest in a characteristic clockwise or counterclockwise fash-

ion. The simplest case during this study was Nest 4 (Fig. 3) where

successive columns rotated in a counterclockwise fashion. This, how-

VoL. 52, No. 1, January 1976 67

Entrance

Ig

16 & i

Fic. 4. Observed columns at Nest 1. For explanations see Fig. 3. During

foraging periods 3, 8, and 10 no columns were formed and random foragers were

22

seen.

ever, was achieved in uneven steps and included some clockwise “back-

stepping” (e.g., foraging periods 8 and 15).

The direction change of the columns at Nest 1 (Fig. 4) was more

complicated, but seemed to occur in a general clockwise rotating pat-

tern. During periods 3, 8, and 10, the typical long, straight column of

foragers was not seen; instead individual foragers left the nest in ran-

dom directions (= random foraging). Columns from this nest gener-

ally did not forage in the area to the west and northwest of the nest

entrance because there were no plants under the Desert Research In-

stitute’s mobile laboratory nor in the gravel road. The columns “swept”

the available area from 0° to 220°; one such sweep was completed

during foraging periods 12 through 19. As was true at Nest 4, some

changes did occur in the direction opposite to that typical for this

nest (i.e., periods 16 and 19).

The columns at Nest 2 (Fig. 5) swept an even smaller area. This

avoided foraging (1) in the adjacent gravel road (which runs past

68 THe Pan-Paciric ENTOMOLOGIST

2 eee

meters

N 20

FA 1 Fe

' at 8 1I8 D.C

4 II a ets

a Ks !°

5

13 12

Entrance

Fic. 5. Observed columns at Nest 2. For explanation see Fig. 3.

this nest from southwest to northeast), and (2) in the foraging area

of Nest 4 (15 m to the east), and (3) in the foraging area of Nest 1

(12 m to the south). Two full counterclockwise sweeps of the area

about Nest 2 are shown in Fig. 5, along with the start of a third. At

this nest clockwise “backstepping” occurred during periods 10, 15, 17

and 20 and for one of the double columns which occurred during pe-

riods 3 and 8. It is very interesting that during periods 8 and 18 double

columns were seen: one column at 218° and the other at 10° (period

8) and 218° and 45° (period 18). These particular columns seemed to

mark the end of one sweep and the start of the next.

2. Response To Diminished Annual Seed Production: Veromessor

pergandei responded to the lower seed production by winter annuals

in three ways: (a) increased collection of non-seed plant material, (b)

VoL. 52, No. 1, January 1976 69

decreased collection of seeds which had become scarce, and (c) in-

creased collection of “less desirable” seeds. (See Table 1.)

(a) Non-seed Plant Material: In 1973, 12.6% of all items collected

was non-seed plant parts; this increased to 27.2% following the lower

seed production of 1974. In both seasons most of this material was

flower parts of Lygodesmia spinosa Nutt., smaller amounts of unidenti-

fiable flowers, leaves, and stems were also found. The exact use of

this and other non-seed material (arthropod exoskeletons and fecal

material) is unknown, but they were rarely found in the refuse pile.

The suggestion by Went et al. (1972) that these materials might be

used in some sort of fungus garden is a possibility.

(b) Scarce Seeds: The general decrease in seed production between

1973 and 1974 is reflected by the fact that 85.4% of all items collected

by the foragers in 1973 were seeds; in 1974 only 71.5% were seeds.

In April 1972, Clark and Comanor (1973) observed that 88% of 252

foraged items were seed and fruits. Tevis (1958) observed, during

a period of drought and poor seed production, that 68.2% of all items

collected by this same species were seeds. This increased to 91.9%

following timely winter rains and a productive spring.

Among the different species of seeds there were notable differences

between 1973 and 1974. Fewer plants of Mentzelia albicaulis Doug].

ex Hook. grew in the study area in 1974, and the seeds of this species

were collected one tenth as often. Mohavea breviflora Cov. and Oe-

nothera clavaeformis Torr. & Frém. had fewer and smaller plants in 1974,

and the percentages of seeds of both of these species decreased. Some

plants, like Phacelia calthifolia Brand and Ph. crenulata Torr., simply

did not germinate in 1974. Differences like that for Chaenactis stev-

ioides H. & A. between 1973 and 1974 may be a result of fewer nests

observed in the latter year. This composite was found only near Nest

3 which was not observed in 1974.

A notable exception to decreased collection of annual seeds occurred

with Cryptantha angustifolia (Torr.) Greene which represented 9.7%

of all items collected in 1973 and 20.7% in 1974. While the plants of

this species were much smaller in 1974, they were still numerous and

produced large numbers of seeds.

In 1973 the seeds of 29 species of plants representing 24 genera

were collected by the foragers. This compares with seeds from 20

species in 16 genera in 1974. Clark and Comanor (1973) reported that

seeds from four species of plants (representing 4 genera) were collected

by this species at our study site in April 1972. Meaningful comparisons

with this study are unfortunately precluded by its brief duration (4

70 THE Pan-Pactric ENTOMOLOGIST

Table 1. Samples of items (number and percent) from foraging

columns at Nests 1 and 2 in 1974 and from all nests in 1973.

Nest 1, 1974 Nest 2, 1974 Total 1974 Total 1973

Seeds of Species Number % Number % Number % Number %

Malvastrum rotundifolium 1 0.1 J. Qe 2 O01

Larrea divaricata > O38 5 0.2 8 03

Menizelia albicaulis 3 0.3 10 (0.5 13 0.4 114 3.9

Lepidium lasiocarpum 7 0.2

Chorizanthe brevicornu 10 «1.1 87 4.0 97 3.1 26 «60.9

Chorizanthe rigida 4 0.2 4 0.1

Eriogonum inflatum 97 10.3 240 11.0 337 10.8 83 28

Eriogonum pusillum 6 0.6 87 4.0 93 3.0 47 1.6

Chenopodium pumilio 149 15.8 1376.3 286 9.1 252 8.5

Atriplex hymenelytra 13 14 5 60.2 18 0.6 55 619

Plantago insularis

fastigiata 3 60.1 3 «60.1

Gilia filiformis 2 02 2 O01 4 0.1 6 0.2

Gilia latifolia 1 0.1 1A eit 26 860.9

Gilia sp. 3 0.1 3 0.1

Phacelia calthifolia 7 0.2

Phacelia crenulata 25 «60.9

Cryptantha angustifolia 203 215 447 20.4 650 20.7 285 9.7

Cryptantha holoptera 76 82.6

Cryptantha nevadensis 2 0.2 2 0,1 4 0.1 18 0.6

Nicotiana trigonophylla 2 O71

Mohavea breviflora 68 2.3

Dalea sp. 8 0.4 8 0.3

Oenothera clavaeformis 177. (18.7 129 =5.9 306 «9.8 951 32.2

Nemacladus glanduliferus 1 0.1

Bebbia juncea 8 0.3

Hymenoclea_ salsola 2 O2 2 0.1 4 0.1 4 0.1

Franseria dumosa 17 18 69 3.2 86 2.7 5301.8

Chaenactis stevioides 2 0.2 2 1 4 0.1 Idl fl

Monoptilon bellioides 4 0.1

Psathyrotes ramosissima 1 0.1 1 0.1

VoL. 52, No. 1, JAnuARY 1976 71

Table 1 (cont.)

Nest 1, 1974 Nest 2, 1974 Total 1974 Total 1973

Seeds of Species _ Number % Number % Number % Number %

Peucephyllum schottii y tee 1 0.1 19 0.6

Lygodesmia spinosa 40 42 251 11.5 291 89.3 192 6.5

Calycoseris parryi 4 0.1

Atrichoseris platyphylla 9 0.3

Unknown Seeds 3 03 7 0.3 10 0.3 30. =—:1.0

TOTAL SEEDS 730 77.2 1503 69.0 2233 71.5 2525 85.4

Non-seed Plant Parts 195 20.6 609 27.8 804 25.7 372 12.6

Arthropod Parts 8 0.9 40 18 48 1.5 381.38

Lizard and Rodent

Droppings 9 1.0 34 §=1.6 43 1.4 16 0.6

Gravel 3 01

TOTAL NON-SEED 212 22.5 683 31.2 895 28.6 429 14.6

TOTAL NUMBER

OF ITEMS 945 2186 3128 2954

NUMBER OF SAMPLES 16 25 41 69

days), number of nests observed (one), low number of items identi-

fied as they were carried past the observer (N = 252), and what seems

to be aberrant behavior of the single nest watched. The largest number

of foragers reported in a column never exceeded 250—an unusually low

number compared to our experience at this site. Tevis (1958) reported

seeds from 14 genera of plants for his entire study. By comparison,

Rogers (1974) found that the harvester Pogonomyrmex occidentalis

(Cresson) collected seeds from 39 species of plants representing 32

genera at the IBP Grassland Biome, Pawnee Site, in northeastern Colo-

rado. Wheeler and Wheeler (1963) reported seeds of 30 species in 26

genera collected by P. occidentalis in North Dakota.

(c) “Less Desirable” Seeds: Chorizanthe brevicornu Torr., an an-

nual with a hard, thick seedcoat, germinated and set seed equally well

in 1973 and 1974, yet it was collected more often in 1974. A closely

related species Chorizanthe rigida (Torr.) T. & G. was abundant both

years but the refuse piles of nests (other than those studied intensively)

in the study area contained large amounts of this seed in 1974, whereas

te THE Pan-PaciFiIc ENTOMOLOGIST

none were found in any refuse piles in 1973. Most of these seeds

had been opened by the ants. Seeds from perennials, such as Franseria

dumosa Gray and Lygodesmia spinosa, were equally abundant in both

years yet both were collected more often in 1974. While these seeds

increased in relative abundance, they may have been ignored origi-

nally due to excess energy required to extract the seed from a hard and

difficult-to-open seed coat. They were collected only when other seeds

were less plentiful.

Conclusions: We have stated previously (in press) that V. pergandei

is very plastic in its behavior: e.g., in nest construction and interchange-

ability of worker roles. This plasticity is seen again in the ability of

this species to make changes in its foraging in response to the varying

productivity in the desert plant community. Indeed, this behaviorial

plasticity must be a major reason why this species is one of the most

abundant animals in the Mohave Desert even during extreme drought.

Certainly such behavior is a beautiful example of adaption for the ex-

tremely varied conditions of the Mohave Desert and contributes to the

predominance of VY. pergandei in that environment.

LITERATURE CITED

Carrott, C. R., and D. H. Janzen. 1973. Ecology of foraging by ants. Ann.

Rey. Ecol. & Syst., 4: 231-257.

CLark, W. H., and P. L. Comanor. 1973. A quantitative examination of

spring foraging of Veromessor pergandei (Mayr) in northern Death

Valley, California. Amer. Midland Natur., 90: 467-474,

Rocers, L. E. 1974, Foraging activity of the western harvester in the shortgrass

plains ecosystem. Environ. Entomol., 3: 420-424.

Tevis, L. 1958. Interrelations between the harvester ant Veromessor pergandei

(Mayr) and some desert ephemerals. Ecology, 39: 695-704.

Went, F. W., JEANETTE WHEELER, and G. C. WHEELER. 1972. Feeding and

digestion in some ants (Veromessor and Manica). BioScience, 22:

82-88.

WHEELER, G. C., and JEANETTE WHEELER. 1963. The ants of North Dakota.

Univ. North Dakota Press, Grand Forks. 326 p.

WHEELER, JEANETTE, and S. W. Rissinc. The natural history of Veromessor per-

gandei (Mayr). Pan-Pac. Entomol., 51: 205-216; 303-314.

A New Proteriades Reared From Trap Stems,

Its Biology and Nest Associates

(Hymenoptera: Megachilidae)

FRANK D. PARKER

Bee Biology and Systematics Laboratory, ARS, USDA

Utah State University, Logan, Utah 84322

An intensive trap-nest study conducted earlier (Parker and Bohart

1966, 1968) yielded abundant information about the nesting habits and

associates of many western twig-nesting bees and wasps, including 9

species of Proteriades. These trap-nest studies are continuing. As a

result, nests of an unnamed Proteriades were collected during the 1973

and 1974 season. A description of the adults, the nesting biology, and

the nest associates of the new species is reported here. Information

concerning other species will be presented in a forthcoming paper on

the biology of the genus.

In the following description of the adults, the term abdomen is the

“apparent abdomen”’ called the Metasoma by some authors.

PROTERIADES SHOSHONE, new species

(Figs, 1-8)

Holotype Male.—Black, mandible apically, apical tarsomeres, tegula, base of

abdominal sterna J-V reddish brown; abdominal terga I-IV mostly red except

dark basal spot, spot progressively larger on succeeding terga; terga V, VI with

narrow apical reddish brown band; tergum VII mostly red brown, darker

medioapically; wings hyaline. Pubescence white, moderate, not longer than

scape except for thinner, longer golden hairs on vertex, scutum; mandible with row

of yellowish setae beneath; golden setae bordering apical margin of sterna;

median basal thick pads of short, golden setae on sterna IJ-IV; terga with patch

of hair at sides, extending apically, medially; apical hair band interrupted medially

on terga I-III, entire on IV, V, absent on VI, VII. Punctation moderate, uniform,

pits small, generally separated by a pit diameter, interpunctural integument shiny;

punctures on clypeus, frons, interocellar area course, nearly contiguous; those on

tibiae sparser, larger; dorsal area of propodeum shagreened basally, shiny, impunc-

tate anteriorly; terga more densely punctate basally, apically; sterna finely, densely

punctate in depressed area, less so apically. Scape twice as long as broad, flagel-

lomeres shorter basally, apically longer; flagellomeres II-VI produced beneath over-

lapping base of succeeding article, more so on basal articles; clypeus slightly

protuberant, shallowly convex, slightly overlapping base of labrum; clypeal margin

convex, minutely sinuate, sublateral margin bent inward; head slightly broader than

long, interantennal distance greater than antennocular distance; interocellar distance

greater than antennocular distance; distance from base of antenna to median ocellus

equal to distance from anterior ocellus to margin of vertex; distance between ocelli

equal to distance from lateral ocellus to margin of vertex; genal width equal

Tue PAN-PaAciFic ENTOMOLOGIST 52: 73-80. JANUARY 1976

74, THE Pan-Pactric ENTOMOLOGIST

to ocellar width; inner eye margins nearly parallel; mandible bifid apically; width

of mandible apically %4 basal width; hypostomal carina raised entire length;

labrum longer than broad, subtruncate apically; propodeum with large oval

depression ending at propodeal orifice; mid coxa with small projecting basal

flange; hind coxa with raised carina along inner ventral margin; femora bulbous,

thickened submedially; tibiae bulbous subapically; terga I-V 3 times as broad as

long; medioapical sulcus on tergum I deep, narrow, extending to midpoint;

anterior margin of tergum VI produced, impunctate, curved to encompass tergum

VII; tergum VII produced apically, its lateral margin with two short, blunt peg-like

projections, margin between pegs sinuate, concave medially (Fig. 4), with shallow

impunctate depression subapically; sternum I with apical margin produced into

low concave keel; sternum II depressed over most of its length, basin-like; apical

margin thickened, especially laterally, more sparsely punctured, broadly rounded;

sterna IIJ—-V similar except apical margins concave, sternum IV more so; margin

of sterna IJ-IV dense fringed with golden hairs; sternum VI with median

protuberance arising mediobasally, covered with stout spine-like projections (Fig.

1); sterna V, VII, VIII as in figures 2, 3, 5; genitalia as in Fig. 6; body length

7mm; wings 4.4mm.

Female.—Like male except: mouthparts with longer setae, bent medially

and apically, twisted; labrum with patch of long narrowed medioapical setae;

hair bands on terga as follows: I, II interrupted medially, III-V entire, V and VI

covered with white recumbent plumose setae; scopal hair not longer than hind

hind basitarsus, rising from raised median horizontal area on each sternum;

sternum VI densely bordered apically by short curved yellowish setae; interocellar

area less punctate; clypeus with prominent swelling basally, with indistinct median

longitudinal line (Figs. 7, 8), apical margin smooth, slightly indented medially;

mandibles 3-toothed, apical teeth closer together than to inner one; mid coxa

without flange; legs less bulbous, keel on sternum I less produced, conical;

apical margins of sterna entire; sternum, tergum VI with broad, round apical

margins; length 5-6mm.

Variation—Two color forms occur in both sexes, one with the abdomen all

black or with small areas of red on tergum I and the other with red markings on

first 4 terga. The pattern of the red markings is diagnostic: terga I-III all red

except for mediobasal black spot, black spot on tergum IV much larger. Most

specimens (80% of males and 66% of females) are the black form. Both color

forms were reared from the same cell series. I have reared similar color forms from

nest series of another Proteriades, P. bullifacies Michener.

Types.—Holotype male and 79 paratypes (14 male, 65 female): 5 mi. E. Wells,

Elko County, Nevada. All specimens reared from trap stems placed in a roadcut

adjacent Interstate 80. Holotype male deposited in the collection at the U.S.

National Museum, No. 73496. Paratypes are in the collections at the USNM,

University of Kansas, and the Logan Bee Laboratory.

Range—FEastern Nevada.

Systematics —P. shoshone is related to bidenticauda Timberlake and Michener,

but the former can be separated in the male by the indented apical margin of

tergite VII and the absence of oblique swellings on abdominal sternite IJ. Females

of shoshone can be separated from those of bidenticauda by the former having

golden pubescence on the vertex and scutum, the distance between ocelli less than

the ocellocular distance, and more distinct hair bands on the first 4 tergites.

VoL. 52, No. 1, January 1976 75

BIOLOGY

Nesting Site—All nests (65, containing 157 cells) were recovered

from prebored elderberry trap stems each with a bore diameter of 3 or

6mm. The stems were placed at the type locality ca. 5 mi. E. Wells,

Elko Co., Nevada, adjacent to Interstate Highway 80 on the east side. The

site is a broad sloping roadcut dominated by sweet clover near the road

and scattered Cryptantha plants above. The trap stems were placed

in a long line at the juncture of the roadcut and the natural vegetation

[dominated by rabbitbrush (Chrysothamnus), sagebrush (Artemisia) ,

and juniper shrubs (Juniperus) |. More nests were recovered the first

year (51) than the next (14).

Nest Construction.—This bee preferred the small side holes (3mm)

for nest building; in 1973, 69% and in 1974, 80% of the nests were

located in these holes. Some nests, 14% in 1973 and 13% in 1974, were

made in medium-sized holes (6mm). Nests made in end holes were near

the top of the boring; an empty space 20-80mm long was under the

first cell. Cells were initiated by partitioning end burrows with thin

discs (Imm) of a mixture of gravel, soil, and coarse sand stuck together

with macerated plant parts (Fig. 11). The partitions were 4—-7mm

apart depending on width of the boring. In wider borings, the cells

were 6 X 4mm (Fig. 11), but in narrower holes the cells were 4 X 7mm

(Fig. 9). Nests in side holes were initiated at the end of the burrow,

and the end of the hole was not supplemented with nest-building

material. Many of the side holes were enlarged and tunneled out (Fig.

10) to hold as many as 7 cells in a series. One burrow was wide

enough to accommodate cells made side by side instead of the usual

pattern of one cell above the other. It is uncertain whether P. shoshone

actually enlarged the borings or if it merely utilized borings of other

aculeates. Bees nesting in end holes never utilized all the available

space in the holes and many loose pieces of pith were not removed,

so it may be that these bees do not modify the nesting burrow. Some

cell partitions were made diagonally across the burrows, especially in

wide borings. Nest entrances were capped by the same mixture as

used in the cell partitions, and some plugs were 5mm thick. The nest

entrance was capped flush with the outer rim in side holes, but end

holes generally were capped 6—l1]lmm inside the borings.

Provisions.—The pollen ball was cylindrical, 3mm wide and 5mm

high (Fig. 9) in nests with horizontal cell partitions or those at ends

of borings. The egg was placed on top of the pollen loaf near the edge

and extended inward.

In sloping cells the bottom of the pollen ball conformed to the slope

76 THE Pan-PaciFic ENTOMOLOGIST

Fics. 1-8. Fig. 1. Sternum VI of male. Fig. 2. Sternum V of male. Fig. 3.

Sternum VII of male. Fig. 4. Tergum VII of male. Fig. 5. Sternum VIII of

male. Fig. 6. Genitalia of male. Fig. 7. Dorsal view of female head. Fig. 8.

Female face.

Vout. 52, No. 1, January 1976 7

©

Fics. 9-14. Fig. 9. Cell with 2nd instar larva dead on pollen ball. Fig. 10.

3-celled nest made in side of trap stem. Fig. 11. 2-celled nest made in end

burrow. Fig. 12. Cocoon, note fussy appearance and fecal pellets near the top.

Fig. 13. Nest in side hole, bottom cell with Stelis cocoon, note nipple at top of

cocoon. Fig. 14. Cocoon of Stelis (Stelidina) sp.

of the cell, but the sides of the pollen loaf were cylindrical and the top

was round. The pollen loaf was compact and composed entirely of

Cryptantha pollen. Not all the provisions were consumed by some larvae,

as small amounts clung to the bottom of the cocoon.

Feces.—Most fecal material was pellet-shaped, orange-brown, uniform,

0.5mm long (about twice as long as broad), with ends usually rounded,

sometimes pointed, an impressed mediolongitudinal line on the surface.

Some of the feces were flattened into long strands and incorporated

into the cocoon (Fig. 12), giving it a striped appearance when held to

the light.

Cocoons.—The oval cocoons appeared fuzzy on the outside due to

718 THe Pan-Pactric ENTOMOLOGIST

the abundant short silk strands that attached it to the cell walls (Fig.

10-12). Cocoons filled most of the cell in short cells, but not in

longer ones. The bottom of the cocoon was round, but the top flatter

with longer twisted silk strands covering the surface. The cocoon

on the inside was smooth and shiny and composed of 2 layers with

fecal material deposited between the layers. Cocoons were usually amber

but rarely white.

Immature Forms.—Prepupae were white and not very active when

disturbed.

Sex Ratio—An unusual ratio of females to males emerged, 4.3:1.

Most nests had the typical pattern of females at the bottom and males

above; but in some nests, the sexes were mixed and in others all were

of one sex. Sex ratios of females to 1 male in the cell series, beginning

with the first cell, were: 3.5, 6, 5.5, 2.3, 2.1, no males, no males.

The range in time for emergence to the adult stage from overwintering

larvae incubated at 72°F was: males 23-25 (24.2) days, females —

25-29 (24.5) days.

Nest Associates and Mortality—In 1973 only 3% of the cells were

parasitized, all by the megachilid bee, Stelis (Microstelis) coarctatus

Crawford. One cell was destroyed by an unknown predator. Cell

mortality due to unknown causes totaled 11%, with 5% occurring in the

ege stage and 6% in larval stages. Parasitism increased markedly in

1974, to 52%, but this year all the parasites were another species of

Stelis, an unnamed species of the subgenus Stelidina (Figs. 13-14).

Cell mortality due to unknown causes during the 1974 season was

only 4%, and this occurred in the larval stage.

Supercedure.—Two nests in 1973 and 1 in 1974 were constructed

above existing nests of another megachilid bee, Hoplitis productus

(Cresson).

DIscussION

I believe this to be the first description of an actual nest of a

Proteriades. A brief note on the rearing of P. xerophila (Cockerell)

from old cells of Anthophora linsleyi Timberlake (Linsley and MacSwain,

1942) prompted Timberlake and Michener (1950) to presume that all

Proteriades species nest in the ground. A solely ground-nesting habitat

for the genus was invalidated when Parker and Bohart (1966, 1968)

established. that P. xerophila and 8 other species of Proteriades nested

in a variety of situations above ground including old galls on Tetradymia,

burrows in stems of Tetradymia and Oenothera, and trap stems.

Linsley and MacSwain (1942) described P. xerophila nesting habits as

Vout. 52, No. 1, January 1976 79

follows: “cells are constructed with macerated plant material—the

young larvae spin a tough yellowish cocoon.” I have reared P. xerophila

from 33 nests, and none of the cells were made from macerated plant

material alone nor were the cocoons yellowish white and tough.

Cells of P. xerophila are constructed with coarse sand and macerated

plant material, and the cocoons are delicate and very white. Linsley

and MacSwain’s description is accurate for cells of Hoplitis biscutellae

(Cockerell), a common bee which these authors also reared. Thorp

(1968) reported on a nest of P. bunocephala Michener, but I question

the identity of the nest builder since: (1) no larvae in the nest were

reared to the adult stage, thus making it difficult to positively establish

what the inhabitants were because of our lack of information on

characters that differentiate larvae among these bees; (2) the female

observed entering the nest could have been searching for a nesting site

or just beginning a nest above the one described as a Proteriades; (3)

nest partitions were “masticated leaf and plant materials.” All 15

species of Proteriades I have reared (800 nests, 3014 cells); as well as

4, species of Hoplitina including bunocephala used coarse sand in nest

construction; and (4) the description of the nest and larvae apply more

to Ashmeadiella than to Proteriades. Both previous reports on nesting

of Proteriades were based on nests obtained from old nesting material,

and it is extremely difficult to state precisely nesting characteristics of

one species if the other species have occupied and altered the nesting

burrow! Abundant information on the biology and nest associates

of other reared Proteriades will be presented in a forthcoming paper on

the biology of the genus.

ACKNOWLEDGMENTS

I would like to thank Dr. C. D. Michener (Univ. Kansas) for

examining the new species and reviewing the manuscript. Thanks are

also due Dr. P. D. Hurd (Smithsonian Institution) and G. E. Bohart

(Utah State Univ.), who also reviewed this paper. Mr. W. P. Nye and

H. Potter took some of the photographs. Thanks are also due to H. Potter

and D. Edgar for drilling holes in the trap stems.

LITERATURE CITED

Linstey, E. G. ano J. W. MacSwarn. 1942. The parasites, predators, and

inquiline associates of Anthophora linselyi. Am. Midl. Nat. 27:402-417.

Parker, F. D. ano R. M. Bowart. 1966. Host-parasite associations in some

twig-nesting Hymenoptera from western North America. Pan-Pac.

Entomol. 42:91-98.

80 THE Pan-PAciFIcC ENTOMOLOGIST

Parker, F. D. anp R. M. Bowart. 1968. Host-parasite associations in some

twig-nesting Hymenoptera from western North America. Pan-Pac.

Entomol. 44:1-6.

THorp, R. W. 1968. Ecology of a Proteriades and its Chrysura parasite, with

larval descriptions. J. Kans. Entomol. Soc. 41 (3) :324-331.

TIMBENLAKE, P. H. anp C. D. MicHENER. 1950. The bees of the genus Proteriades.

Univ. Kans. Sci. Bull. 33:387-440.

SCIENTIFIC NOTE

Aprostocetus diplosidis, a Parasite of the Sorghum Midge Found in Cali-

fornia (Hymenoptera: Eulophidae).—Aprostocetus diplosidis Crawford, a

parasite of the sorghum midge Conéarinia sorghicola (Coquillett), was first de-

scribed from material collected near Baton Rouge, Louisiana (Dean, 1911: USDA

Bull. 85 (IV): 39-58). Its distribution generally follows that of its host although

several years may elapse between the discovery of the midge and the appearance of

A. diplosidis (Dean, 1911: Ibid). A. diplosidis was introduced into Texas (San

Antonio) in 1908 (Dean, 1910: J. Econ. Ent. 3: 205-7) and although it is very

aggressive, it appears to have been displaced by another parasite, Eupelmus popa

Girault (Walter, 1941: USDA Tech. Bull. 778). The latter species has been re-

ported by Woodruff (1929: J. Econ. Ent. 22: 160-7) to feed on A. diplosidis

larvae.

C. sorghicola was first recorded in California (Tulare Co.) in 1960 by Lange,

Marble, Pendery, Burton (1961: Calif. Agri. 15(1): 7-9) but no parasites were

found at that time. During field investigations in 1972 I found several small wasps

in emergence cages used to determine the midge infestation level in sorghum heads.

These were tentatively identified by Dr. R. L. Doutt (Dept. of Entomological Sci-

ences, University of California, Berkeley) as belonging to the genus Aprostocetus,

“species probably diplosidis.” Subsequent examination of additional material and

reference to Burks’ key (1967: Ann. Ent. Soc. Amer. 60: 756-60) confirmed that the

specimens were A. diplosidis. The first individuals emerged 28 Sept., 1972 from sor-

ghum heads collected near Ivanhoe (Tulare Co.) on 11 September. A. diplosidis also

emerged from sorghum heads collected from two other fields, one in the Ivanhoe

area, sampled on 11 Sept., the other 5 miles south-east of Dinuba (Tulare Co.)

sampled on 18 September. No A. diplosidis emerged from 480 sorghum heads

collected from sorghum at the Kearney Horticultural Field Station, Parlier (Fresno

Co.), California. Based on emergence data of C. sorghicola and A. diplosidis in

the three fields from which it was collected, the rate of parasitism was < 1%.

During 1973 I surveyed sorghum fields throughout Madera, Fresno, Tulare, Kings,

and Kern counties. A. diplosidis was found in only two locations, one near Ivanhoe

on 25 Sept., adjacent to a field where it was collected in 1972 and a second on 28

Sept. from a sorghum trial at the Kearney Horticultural Field Station. As in 1972,

parasitism was <. 1%. No specimens of A. diplosidis were found during a similar

survey in 1974. The present known distribution of A. diplosidis in California is

shown in Fig. 1.

VoL. 52, No. 1, January 1976 81

FRESNO

| Contarinia sorghicola

| Aprostocetus diplosidis

Fic. 1. Distribution of Aprostocetus diplosidis in relationship to its host Con-

tarinia sorghicola in the southern San Joaquin Valley.

It is not known if the absence of A. diplosidis in the 1974 survey indicates that

the parasite has failed to become established. Even if A. diplosidis becomes estab-

lished, it is not likely to contribute significantly to the regulation of C. sorghicola

populations. [t does not appear until late in the season (after 11 Sept.) by which

time the majority of sorghum midge damage has been done. In other areas of

the U.S. where it is established it is not considered to be of major importance in

controlling midge populations (Young, 1970: p. 235-87. In Wall and Ross [eds.]

Sorghum production and utilization. AVI Publ.).

This report constitutes an extension of the range of A. diplosidis, and a new

record for California. The previous extent of its westward distribution as listed by

Burks (1967: Loc. cit.) was Kansas and Texas. A. americanus Ashmead is the

only other member of the genus reported to occur in California (Burks, 1967:

Ibid.).

The assistance of Dr. R. L. Doutt is greatly appreciated. Thanks also to Dr. R.

E. Rice for his comments on the manuscript.—CHARLES G. SuMMERS, Department

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

82 Tue Pan-PacitFic ENTOMOLOGIST

SCIENTIFIC NOTE

Observations on the Habitat of Aegialites (Coleoptera: Salpingidae).—

Aegialites is known from 4 species occurring in maritime habitats in North America

and Japan: (Spilman, 1967. Pacific Insects, 9: 1-21). These beetles are rarely

collected, and are almost unknown ecologically.

In October 1974 I found large numbers of Aegialites californicus (Motschoulsky )

on the vertical granite cliffs at Cape Flattery at the northwest tip of Washington.

The beetles occurred in and around cracks in the rock from the high-water mark

and splash zone to the level reached only occasionally by spray. None were found

into the mussel or seaweed zone nor in areas exposed to water-propelled sand.

Some of the lowest individuals would have been submerged at every high tide but

the upper ones were permanently above the water—an “elevation” range of 3-5 m.

Five beetles kept alive were active only at night, clustering on the underside of a

piece of bark during the day.

In 1964-65 I attempted to discover the range of Aegialites in California. Though

specimens of A. fuchsi Horn could not be located in Mendocino County (the type

locality) two colonies turned up on Point Reyes Peninsula, Marin County. The

largest of these, on the northern tip of the peninsula, was first encountered atop

the cliff under a thin crust on soft soil in a patch of barren ground. Apparently

spray reached this spot during a storm at high tide and beetles moved off the

rock into it. During a later visit, this soil was drier with no trace of the beetles.

The main body of the colony was located on the vertical face of the granite cliff,

wet from windblown spray, under conditions as described for A. californicus. These

two species are apparently restricted to rocky points exposed to the violent surf

of the open Pacific.

Aegialites subopacus (Van Dyke) occurred in varying numbers at nearly every

intertidal rocky area from Moss Beach, San Mateo County to Point Arena,

Mendocino County. It ranged vertically about 5 feet and appeared to tolerate more

sand than A. fuchsi. The two species occurred sympatrically at both localities

mentioned above, but swbopacus was closer to the water. In one case, both species

were found beneath a single stone, separated by about 10 cm. The largest A.

subopacus population, with thousands of individuals, was on Point Diablo, Marin

County. The two smallest insect colonies I’ve ever seen consisted of this species.

One was atop a solitary rock inside San Francisco Bay near Richmond (with at

most 100 individuals on an area of less than 5 square feet); the other was on a

rounded granite boulder half buried in sand on a San Francisco beach (less than

50 adults).

Where fresh water seeped through or over the rocks intertidal beetles were

seldom found, and they often became much less numerous after the first heavy

fall rains. Possibly as the result of the relatively constant temperature of sea water,

I found adults of intertidal insects throughout the year along with various instars

of staphylinid, carabid and other larvae. Collection months for A. fuchsi were

July, October; for A. subopacus January, March, May, June, July, August, Octo-

ber—DeERHAM GIULIANI, 170 Flower Alley, Big Pine, California 93513.

VoL. 52, No. 1, January 1976 83

PACIFIC COAST ENTOMOLOGICAL SOCIETY

F. G. ANDREWS Ronatp E. STECKER F. ENNIK P. H. ArNaAup, Jr.

President President-Elect Secretary Treasurer

PROCEEDINGS

THREE HUNDRED AND SIXTy-THIRD MEETING

The 363rd meeting was held Friday, 21 February 1975, in the Morrison Auditorium

of the California Academy of Sciences, Golden Gate Park, San Francisco, President

Daly presiding, with 40 members and 24 visitors present.

The following persons were elected to membership. Student membership:

Patricia Bushnell, Albert Green, Rhonda D. Oliver, Norman J. Smith, Gordon

Sweeny, Charles H. Dill. Regular membership: Steven Anderson, Dr. Janice

Gillespie, Scott McCleve, John W. Armstrong.

The following notes were presented:

The Activity of Several Geometridae in the San Francisco Bay Area in

January (Lepidoptera) .—January of 1975 in the Bay Area was mostly sunny with

relatively warm mid-day temperatures. Adult flights of some diurnal Geometridae

occurred before heavy winter storms commenced the last few days, and extended

through most of February. Three univoltine species are of note.

Brephos infans oregonesis Swett was flying in Stonybrook Canyon (Palomeres

Rd., n. of Niles Canyon), Alameda County, California on January 24, 1975. I

first found this moth here on February 19, 1967, and all four stages were collected,

the dates were mid to late February. The adults were attracted to willow catkins

on the February dates, but in January of this year the trees were bare, with no

apparent nectar source.

Hydriomena nubilofasciata Packard was collected in Alum Rock Park, Santa

Clara County, California, on January 18, and was in good numbers in Stonybrook

Canyon on January 24, 1975. Though not strictly diurnal (it freely comes to

light), this widespread species (with several named forms), often flies in the

daytime. It can become rather common in moist, shady canyons in February and

early March.

Epirrhoe plebeculata Guenée was extremely abundant at Alum Rock Park on

January 18, and was already showing signs of wear by January 24 at Stonybrook

Canyon. This diurnal species usually has its peak flight in February or March

in northern California. I had observed this moth on December 27, 1972, in the

Santa Barbara area, and there is no doubt other “winter” records exist for it in

southern California. January records for previous seasons include other geometrids

(mostly Hydriomena spp.), noctuids, plus the Lycaenids Plebejus acmon “cottlei”

(Grinnell) and Philotes sonorensis (Felder and Felder).—Rosert L. LANcsTon,

Kensington, California.

The Range Extension of the Fruit Pest Stelidota geminata (Say) in

California (Coleoptera: Nitidulidae).—Both male and female specimens of

Stelidota geminata (Say) were collected by John W. Armstrong and Edwin L.

Soderstrom from samples of decaying oranges and fig bait traps in Fresno and

Madera Counties, California. Larvae of S. geminata were found in decaying oranges.

These specimens were collected from March through September, 1974.

S. geminata is known to occur from Massachusetts (Tyngsboro, Springfield) to

84, THE Pan-Pactric ENTOMOLOGIST

Florida (Capron, Haulover, Edgewater, Miami, Paradise Key), west to Iowa

(Mt. Pleasant), Missouri (St. Louis) and Texas (Columbus), south through

Central America to Colombia and Brazil. In the western United States, S. geminata

was previously known to occur only in southern California (San Diego to Los

Angeles), south of the Tehachapi Mountains—JoHn W. Armstrone, Fresno,

California.

The Effects of Gregarines on the Metabolic Rate of Tenebrio molitor

Larvae (Coleoptera: Tenebrionidae).—Most populations of Tenebrio molitor

harbor large numbers of gregarine protozoans in the alimentary canal. Larvae may

contain up to 6,000 gregarines completely blocking the alimentary canal. The

respiration rate of infected and uninfected larvae were compared by use of a

modified Wenneshind-Scholander single stage respirometer. Each larva was

placed in a separate respirometer and readings were taken for a period of four

hours, allowing half an hour for the larvae to adjust. Two trials were run, each

consisting of 24 infected and uninfected larvae. The trials were performed at

night between the hours of 9 p.m. and 2 a.m. The temperature ranged from

22.5 to 24.0 degrees centigrade. Upon the elapse of the four hours, the larvae

were removed from the respirometers, weighed, mildly anesthetized with a few drops

of 70% alcohol on the integument, and dissected. The alimentary canal was

removed and teased to certify that infection or uninfection of the larvae. The

results showed that there is a highly significant difference in the metabolic rate of

the infected and uninfected larvae (P < 0.01), with the infected larvae respiring

at a much higher rate.

To establish whether there is a correlation between the number of gregarines

present and the metabolic rate of the larvae, the respiration rate of twenty infected

larvae was studied. The concentration of gregarines was recorded as number of

gregarines per larva. Regression analysis showed that there is a definite positive

correlation between the number of gregarines and the larval metabolic rate

(P < 0.01). The mean metabolic rate of uninfected larvae was determined to be

0.9275 ml 02/g. wet wt./hr., whereas the mean metabolic rate of infected larvae

was 1.2173 ml 02/g. wet wt./hr. The increase in metabolic rate was calculated to

be 0.00554 ml 0:/g. wet wt./hr./gregarine—Grorce N. ZuHovresorr, California

State University, San Francisco.

Dr. Zhovreboff also reported on the apparent absence of extra-cellular protozoan

or helminth symbionts in the alimentary canal of the scorpion Uroctonus mordax

(Vaejovidae), based on a detailed examination of 13 males and 7 females collected

from Marin County, California.

The main speaker of the evening was Dr. C. B. Philip, California Academy of

Sciences, who presented the following talk as the presidential address for 1974:

Horse-flies, too, Take Some Victims in Cold-blood, as on Galapagos Isles.

—It is not new that tabanid flies have been occasionally reported as attacking

reptiles in various parts of the world, mostly subtropical. Such attacks, by

host-seeking female flies to obtain ovigenous blood, relate to the availability of

usually warm-blooded hosts, but recent evidence suggests occasional development

of even preferential feeding on cold-blooded ones as well.

My interest in cold-blooded entomophagy was enhanced on coming to the

California Academy of Sciences (CAS) in 1970, by finding a label, “biting turtle,”

cited later, on a pinned Galapagos horse fly. My speculative curiosity was aroused

on how frequently horse flies might be attacking other reptiles on the Archipelago.

VoL. 52, No. 1, JAnuARy 1976 85

At one time, the islands, though of volcanic origin, were popularly believed to

have had land connections to the mainland in the geologic past. Now, with the

concept of continental drift widely accepted, the movement of the mostly submerged

Pacific plate is considered; to be slowly toward the continental plates some 600

miles to the east (Cox, 1966 The Galapagos, Univ. Calif. Press, pp. 78-86), not

away from them. The obvious evolutionary derivation of the one Galapagos horse

fly species from recent ancestors of a large group of mainland species indicates

insular invasion by this horse fly in the relatively recent past. It is still a matter

of opinion among taxonomists whether this is a specifically or only subspecifically

distinct population, but reptile feeding would be an acquired secondary habit.

Historical. Interest in feeding of biting flies on reptiles was accelerated by

discovery of natural and experimental feeding of Culex mosquitoes on hibernating

and caged garter snakes, and impingement on a potential overwintering mechanism

of western equine encephalitis virus (see Thomas and Eklund, 1962, Proc.

Soc. Expt. Biol. Med., 109: 421-424). The recent first discovery by DeGiusti

and colleagues at Wayne State University, of biological transmission of a blood

protozoan Haemoproteus, among Michigan turtles by Chrysops callidus Osten

Sacken, added interest to this subject. C. callidus is a well-known deer fly pest

of man and animals, sometimes locally numerous from British Columbia to Maine

and south from Arizona to Florida; it is likely that other species of tabanids

with suitable habits can also carry the turtle blood-parasite.

Tabanid association with reptiles has been noted sporadically in the past. The

original series in the British Museum of the rare Tabanus crocodilinus Austen from

Nyassaland was labelled “on crocodile.” Though actual attack on these reptiles

was questioned later in 1954 by Oldroyd (The Horse-flies of the Ethiopian Region,

Vol. II.), credibility is strengthened by observations, specimens and photographs

from Professor Federico Medem of Instituto “Roberto Franco,” Villavicencio,

Colombia, during unpublished conservation studies of caimans in upper tributaries

of the Amazon River. He observed 4 species in 3 tabanine genera to feed exclusively

or preferentially on indigenous crocodilian hosts: Phaeotabanus fervens (Linn.),

P. obscuripilis (Krb.) (= nigriflavus Krb.), Stenotabanus incipiens (Walk.),

and Tabanus dorsiger ssp. modestus Wied. He also forwarded for identification

another 6 species in 4 genera which he took occasionally on these reptilian hosts,

namely, Diachlorus curvipes (Fabr.), Catachlorops (Psarochlorops) testaceus

(Macq.), P. cajennensis (Fabr.), P. innotescens (Walk.), P. ?prasiniventris

(Krb.), and JT. cicur Fehld. (= ?dorsiger ssp. stenocephalus Hine.). One of

these, P. fervens, has also been reported biting South American boas. A boa is

also listed as bitten in Old Panama Zoo by Fidena isthmiae (= flavipennis Krb.).

Austen (cited by Oldroyd) also notes that “Tabanus” (= Neavella) albipectus

Bigot, which is precinctive in the Malagasy Archipelago, was observed biting sea

turtles on Aldabra Is., and apparently also at sea, probably an old-established

feeding habit there. Aldabra Is., be it noted, is the only place outside the

Galapagos where giant tortoises have barely survived extinction by man.

Bequaert (1940, Rey. Entomol., 11: 253-269) comments on “the poverty of the

tabanid fauna” in the Antilles as well as of the usual mammalian hosts with no

native ungulates there, the preferred source of blood elsewhere. He further states

“it is quite possible that some of the horse flies which we now observe biting man

and domestic animals, occasionally fed on cold-blooded vertebrates,” since certain

86 THE Pan-Paciric ENTOMOLOGIST

species have elsewhere been observed “feeding on reptiles (crocodiles, monitors,

and turtles).”” The reference to monitors is probably the one reviewed by Leclercq

(cited by Oldroyd) of T. guyonae Surc. on a large, sun-basking, Varanus monitor

in the Algerian desert country. Inquiries of biologists studying the Komodo

dragon in the East Indies have not revealed tabanid attacks.

Observations on Galapagos Horse Flies. A long series, Tabanus vittiger Thomson

in CAS from various Galapagos islands, includes a female collected by F. X.

Williams, Entomologist on a 1906 Academy expedition, with his pencilled label

“biting turtle.” A published photo (see Slevin, 1911, Calif. Acad. Occas. Pap. 17:

1-162) shows him in a lagoon in a row-boat with overturned sea turtles; since

an objective of the expedition was land tortoises, his label would have so indicated

had a tortoise been the host. On recent inquiry of first-hand observers, I have

confirmed that this, the only species of tabanid on Darwin’s “Enchanted Isles,”

has been seen biting not only bathers and sea turtles, basking in the lagoons, but

also land tortoises in pens at the Darwin Research Station, Santa Cruz Island.

I suspect that in suitable locations land iguanas could have been also sources

of blood. Since their numbers have become so depleted and isolated locally,

they are not now the important hosts that I hypothesize various reptillians once were

on the Archipelago before man introduced now-common goats, pigs, dogs, and

other warm-blooded animals.

However, subsequent, admittedly rather sketchy information, makes me revise

my earlier theory that land reptiles on the Galapagos could have supplied the

past major source of ovigenous blood for the indigenous horse flies. An analysis

of casual horse-fly collection records points to habitats near sea-level lagoons and

beaches, with none of these flies among insects collected from the rainy

cloud-moistened uplands.

While blood-seeking female Tabanus are strong fliers, circumstantial evidence in

general indicates reluctance to stray far from semi-aquatic (here probably brackish

water) breeding places, depending on availability of hosts. It is likely also

that non-bloodsucking males would not wander very far from such breeding sites.

Surprisingly, only 1 male vittiger has been reported. It was described by me in

1965 (Ann. Entomol. Soc. Amer., 58: 876-880) from Post Office Bay, Floreana,

taken in April with 22 females by a Munich Museum expedition.

Data which I have compiled from various collections and reports on 250 females

additional to 2 other locations (locs.) on Floreana (or in some early ship’s logs.

Charles) for April and July, are also from coastal areas, probably in vicinity of

mangrove thickets: SANTA CRUZ (INDEFATIGABLE) —5 locs., Febr., Mar.,

Apr., May, July, and Nov.; SAN CRISTOBAL (CHATHAM) —2 locs., Apr., Oct.;

ISABELLA (ALBERMARLE)—Apr.; FERNANDINA (MARBOROUGH)—June;

BALTRA (SEYMOUR)—Mar.; SAN SALVADOR (JAMES)—Febr.; and SAN

BARTOLOME—2 locs., Febr.

These casual samples by various collectors since 1906 suggest not only that

the species has been established in the Archipelago long enough to invade most,

if not all, the major islands, but that breeding may be less seasonal in these

equatorial situations near sea level, than the breeding of mainland relatives. The

above sporadic records include all months except December, and tabanids have

been taken on occasion in considerable numbers together in one place. Word from

actual collectors of several series, located the prolific sites as on beaches near

mangrove thickets. The only record that appears to be at a distance from the

VoL. 52, No. 1, JANuARY 1976 87

beaches, is Finca Castro at “200 m. alt.,” which is an hacienda in the hills a

short distance above Academy Bay on Santa Cruz.

Such coastal breeding habitats are also substantiated by Dr. Peter Cramer,

former Director of the Darwin Research Station, Santa Cruz (in litt., May 1971)

who observed horse fly prevalence along the coasts, especially during the “hot

season” (Jan. to Apr.) when often several flies at a time fed on penned tortoises

at the Station, biting between the horn plates.

During questioning of Tui De Roy, who spent 18 years of her youth on the islands,

I learned that she and her brother had stirred up submerged, torpedo-shaped

“maggots which surfaced swimming” in rain pools above a lagoon; these might

have been tabanid larvae. The biting adults with prominently-lined abdomens,

were quite familiar to the De Roys, especially while bathing, and they also

confirmed biting of sun-basking sea turtles. In her most recent letter, she confirms

also seeing flies at least attempting to bite young sea lion pups at play on a

mangrove-bordered beach. The pups seemed to pay no attention to the flies, but

were too active to determine at a distance if any flies actually obtained blood.

It seems not improbable that flies ranging from such habitats, might also invade

marine iguana rookeries on not too distant rocky points, though wind would likely

be more of a deterrant on such exposures.

The present wide inter-island horse fly distribution can hardly have resulted

either from the advent of man’s introduced animals or from some unknown migrant

mammal that has disappeared. On the other hand, the here-postulated considerable,

primitive resort of this horse fly to reptiles as a source of blood would be a

secondary development by the flies after early “invasion” from the mainland. This

tabanid has been isolated on the Archipelago long enough to develop its own

biological identity, but is still considered only a recent offshoot from a vigorous,

modern, highly specialized group of mainland Tabanus the females of which are

primarily dependent on warm-blooded vertebrates for their ovigenous blood-meals.

“Invasion from the mainland” is a glib way of avoiding speculation on how this

one horse fly species could have reached the islands. Flying heavy-bodied insects

are not believed to have made the long, overseas journey by wing. Conversely,

at least some, like cerambycid beetles, most likely were carried by the prevailing

currents, though seasonal, in floating material, which seems the most likely

pathway for T. vittiger in the larval stage to have reached the islands. To become

established, however, there would be the further extraordinary circumstance of

both incipient sexes of these probably cannibalistic tabanid larvae surviving

successfully cross at or about the same time! They obviously arrived long before

the advent of man as discussed above. No other explanation for their insular

occurrence at present seems plausible, but it is not unlikely the progenitors were

also brackish water, sea-level breeders. This may even provide an ecological clue for

seeking a close, modern relative on the west coasts of the mainland from which

the Humboldt current takes its well-known swing westward to envelop on occasion

the Galapagos Archipelago.

The closest presently recognized relative, based on adult structural similarities in

both sexes, 7. guatemalanus Hine, is not known southwest of Panama. Fairchild

(1942, Ann. Entomol. Soc. Amer., 35: 153-183) makes the pertinent statement

that guatemalanus (as subspecies of vittiger) in Panama occurs uncommonly

“where it seems to be confined to the drier areas of the Pacific Coast.” I have

88 THE Pan-PactFic ENTOMOLOGIST

seen specimens from the Mexican west coasts of Nayarit, Sinaloa, and Sonora

(including a confirmatory male from “nr. San Jose beach, 40 mi. SW Obregon,

16-23.V.64, Howden & Martin” with females in the Canadian National Coll.).

The species is more common from around the eastern Mexican Gulf Coast, southern

Florida, and some Caribbean islands, to Surinam; the breeding habitats, when

known, could also include brackish water situations. Recent capture in April,

1974 by J. T. Doyen of 4 guatemalanus females in a Malaise trap set in a sea-level

mangrove swamp on Isla San Jose in the Gulf of California, strengthens the

postulated similarity in breeding sites to the Galapagos relative. Of possible further

related zoogeographic significance is that certain other components of the Galapagos

insect fauna and flora are known to have some early Caribbean relationships. It is

interesting that another of the “lined horse fly” group, T. dorsiger var. modestus

Wied., widespread in northern South America, is one of the species reported by

Dr. Medem to preferentially attack crocodilian hosts in the Colombian Amazon

Basin.

The speculative question naturally arises, could the progenitors of modern,

bloodsucking Tabanidae ever have fed on dinosaurs when those large reptiles

were a dominant form of terrestrial life. Though the paleontologic evidence is

meager, the answer appears to be “yes.”

Mackerras (1954, I. Aust. J. Zool., 2: 431-454) summarizes the evolution of the

Tabanidae as a specialized derivation of the Diptera which Tillyard believed

originated in the Permian just before the Mesozoic era, often referred to as the

“Age of Reptiles.” Generalized, non-bloodsucking ancestral tabanids were sug-

gested to have originated about the beginnings of the Mesozoic; they probably fed

on “juices of plants,” but an “adaptation to blood-sucking” was considered by

Mackerras to be one of the early trends in their evolution. Certain groups of

tabanids with the popular “Gondwanaland distribution must have evolved not

later than the middle of the Mesozoic”; this would have been when reptiles were

flourishing, and would furnish probably the most available source of blood for

those specializing tabanid ancestors that had “developed the habit.”

No need to carry this speculation further than to contrast it with the present-day

situation when, as Bequaert (referred to above) states in discussing food habits of

the adults: “The females of most species suck vertebrate blood, but they often

prefer to bite certain types of animals. Mammals, and particularly ungulates, are

the preferred hosts of the majority of species.”

It is also speculative to decide which, if any, present-day species attacking

cold-blooded vertebrates, are exhibiting atavistic, “cold-blood-feeding” tendencies,

or simply the pursuit of vertebrate blood from the most available local hosts. At

least in the early restricted Galapagos ecosystem, any habitual feeding on reptiles

must have developed secondarily.

Finally, an amusing bit of speculation relates to the oft-cited inclusion of

“dinosaur juices” in the fossil fuel with which we propel our cars—if ancient

tabanids occurred in pest numbers, I’ve been intrigued by the possibility I might

be burning fossil tabanids in my auto travels as well.

THREE HuNDRED AND Srxty-FourtH MEETING

The 364th meeting was held Friday, 21 March, 1975, in the Morrison Auditorium

of the California Academy of Sciences, Golden Gate Park, San Francisco, President

Daly presiding, with 16 members and 14 visitors present.

VoL. 52, No. 1, January 1976 89

The following persons were elected to membership. Student membership:

Stephen L. Buchman, Terry A. Klein, Raymond T. Ross, Jeffrey F. Holzman,

John Hafernik, George N. Zhovreboff, Mike Phelan, Scott E. Miller. Regular

membership: Jan Brocksen.

The main speaker of the evening was Dr. Hugh F. Rowell, Department of

Zoology, University of California, Berkeley. His illustrated talk was titled

“Control of coloration by acridid grasshoppers”.

Coffee and refreshments were served in the Trustees Room following the

meeting —F. Ennik, Secretary.

THREE HUNDRED AND Sixty-FirtH MEETING

The 365th meeting was the annual field and picnic held on Saturday, 10 May

1975, at Castle Rock Park, near Mount Diablo, Walnut Creek, California, with

11 members and 13 visitors present.

Insect collecting in riparian, chaparral, and oak-savanna woodland _ habitats

were the main activities. Swimming and other recreational activities were also

available——F. Ennik, Secretary.

THREE HUNDRED AND SIxTy-SixtH MEETING

The 366th meeting was held Friday, 17 October 1975, in the Morrison Auditorium

of the California Academy of Sciences, Golden Gate Park, San Francisco, President

Daly presiding, with 40 members and 42 visitors present.

The following persons were elected to membership. Student membership:

Rolf L. Aalbu, Philip Haney, Mark A. Deyrup, Kirk I. Hays, David J. Minnesang,

Merton Price, Jr., Kathleen Richardson, James J. Tokarz, Bruce Walsh, Marvin

Dommer, and Robert Horton. Regular membership: John K. Bouseman, Glenn D.

Campbell, Dr. Clifford D. Ferris, Dr. Richard D. Goeden, Terry Griswold, Randy

Hansen, Dr. J. C. Lien, Robin McMaster, Tom Parker, Durward D. Skiles, Michael

F. Tharp, Philip F. Torchio.

President Daly reported on behalf of the Executive Committee that Dr. Melville

H. Hatch and Mr. Hugh B. Leech were elected to Honored Member status for

their many years of dedicated service to entomology and to the Society.

Doctor David H. Kavanaugh was appointed Historical Committee Chairman by

President Daly to replace Hugh B. Leech.

Bruce Walsh reported on several butterflies of the genus Pseudophilotes,

found in isolated areas along the coast range of California, which are becoming

rarer due to human encroachment upon their native habitat and host plants.

The following note was presented:

Sarcophaga tibialis Macquart Intercepted in California (Diptera: Sarco-

phagidae).—In February and March, 1975, The California Academy of Sciences

sponsored a “Coelacanth Expedition” to the Archipel des Comores under the

direction of Dr. John E. McCosker, Superintendent of Steinhart Aquarium, in

pursuit of live specimens of Latimeria chalumnae. From associated research

projects, dry invertebrate materials collected by Mr. Les Gunther were packed at

Moroni on the island of Grande Comore and air shipped to the Academy. On

March 26, 1975, in the process of unpacking this material, Mr. Dustin Chivers, of the

Department of Invertebrate Zoology, found several live Diptera larvae. These

were placed in a jar with sand, the larvae pupated, and on 19 April 1975 two

90 THE PAn-Paciric ENTOMOLOGIST

male sarcophagids emerged. These were forwarded to Dr. F. Zumpt, of the South

African Institute for Medical Research, and J. Reed, of his staff, identified them

as Sarcophaga tibialis Macquart. This species is reported to be common in the

Madagascan region.—P. H. Arnaup, Jr., California Academy of Sciences, San

Francisco.

The principal speaker of the evening was Mr. Ron Russo, East Bay Regional

Park Naturalist, whose illustrated talk was entitled, “Natural History of Galls and

Gall-Forming Insects of California”.

Coffee and refreshments were served in the adjacent Bufano exhibit room

following the meeting.—F. Ennik, Secretary.

THREE HUNDRED AND SIXTyY-SEVENTH MEETING

The 367th meeting was held Friday, 14 November 1975, in the Morrison

Auditorium of the California Academy of Sciences, Golden Gate Park, San

Francisco, President Daly presiding, with 35 members and 39 visitors present.

The following persons were elected to membership. Student membership:

Robert W. Brooks, Natalie Vandenberg, Herbert C. Field. Regular membership:

Brent L. Prothero, Dr. Alice Hunter.

Professor Charles H. Martin, retired Professor of Entomology, Oregon State

University and Entomologist, Oregon Agricultural Experiment Station, died on

October 21, 1975, at Tucson, Arizona. Professor Martin was a member of the

Pacific Coast Entomological Society for 43 years, having been elected to member-

ship in 1933,

Charles H. Martin was born in Aline, Oklahoma, on March 23, 1901. He

received his B.A. and M.A. degrees from the University of Kansas, and he reported

in his “Membership Record” that Dr. H. B. Hungerford was responsible for his

choosing entomology for a career. He received his Ph.D. (Entomology) from

Cornell University in 1939. He was appointed Associate Entomologist in 1946 and

Associate Professor of Entomology in 1947 at Oregon State University, and retired

from these positions in September 1967.

Professor Martin was a specialist of the dipterous family Asilidae and made

important contributions to the knowledge of this family. His last major paper

entitled “The generic and specific characters of four old and six new Asilini

genera in the Western United States, Mexico, and Central America” was

published on September 11, 1975, by the California Academy of Sciences as

Occasional Paper No. 119 (with 107 pages and 78 figures). His private collection

of Asilidae, which numbered approximately 50,000 specimens (in 1965) with

1250 species, was recently sold to the Florida State Collection of Arthropods at

Gainesville.

Professor Martin is survived by his wife Luretta B. Martin, and a sister,

Irene Martin—P. H. Arnaup, Jr., California Academy of Sciences, San Francisco.

Alan Kaplan reported a new host record for the trephritid, Paroxyna americana,

in the flower heads of the composite, Grindelia humilis, which grows along the

salt marshes of Richardson’s Bay.

The following note was presented:

Compsilura concinnata (Meigen) (Diptera: Tachinidae) Reared from

Halisidota (Lepidoptera: Arctiidae) in California.—Compsilura concinnata

(Meigen) is considered the most important parasite in the United States against

the Gypsy Moth larva. In North America it has been reared from about 150

VoL. 52, No. 1, JANuARY 1976 91

different species representing 22 families of Lepidoptera (Arnaud, A host-parasite

catalog of North American Tachinidae, in press). Compsilura concinnata was

first introduced in New England from several European countries in 1906

(Clausen, 1956, U.S. Dept. Agric. Tech. Bull. No. 1139:97) and subsequently also

colonized in other states including California.

Compsilura concinnata has been rarely encountered in California. Two records

have been published on the basis of host rearings in the San Francisco Bay Area

in the Pan-Pacific Entomologist (in 1969 by Arnaud, vol. 45, p. 77, from

Agraulis vanillae incarnata (Riley) in San Leandro; in 1973 by David J. Horn,

vol. 49, pp. 402-403, from Malacosma spp. (constrictum (Hy. Edwards) and/or

californicum (Packard) in Hayward).

A mature larva of Halisidota maculata (Harris) or possibly H. argentata Packard

(determined by Dr. T. D. Eichlin) was collected on a house in the Ingleside

District, San Francisco, California, in October or November 1974, by Ms. Roxi

Berlin. The host larva did not feed and pupated thereafter. The fly larva

emerged from the posterior end of the Halisidota puparium and pupated within

the moth cocoon. A female specimen of Compsilura concinnata emerged on

February 20, 1975—P. H. Arnaup, Jr., California Academy of Sciences, San

Francisco.

The principal speaker of the evening was Dr. William E. Ferguson, Department of

Biology, California State University, San Jose, whose illustrated talk was entitled,

“Entomological observations in Asia, Australia, and some Pacific Islands”.

Coffee and refreshments were served in the Trustees Room following the

meeting.—F. Ennik, Secretary.

THREE HUNDRED AND Sixty-E1igHtH MEETING

The 368th meeting was held Friday, 19 December 1975 in the Morrison

Auditorium of the California Academy of Sciences, Golden Gate Park, San

Francisco, President Daly presiding, with 33 members and 12 visitors present.

The following persons were elected to membership. Student membership: Dennis

Morihara, David Wahl, Steven R. Scott, Herbert C. Field. Regular membership:

James Saulnier, Stanley R. Nichols, Steven L. Jensen.

President Daly appointed Dr. John Pinto and Dr. T. D. Eichlin to the Publication

Committee to replace Marius Wausbauer and E. Gordon Linsley. On behalf of

the Nominating Committee, composed of William Ferguson, Robin Thorp and F.

Louis Blanc, Dr. Ferguson announced the following slate of candidates for 1976:

President, Dr. Fred G. Andrews; President-Elect, Dr. Ronald Stecker; Treasurer,

Dr. Paul Arnaud, Jr.; and Secretary, Franklin Ennik. There were no nominations

from the floor and the candidates were elected by unanimous vote.

D. H. Kavanaugh showed color slides of the habitat of Nebria desolata Kavanaugh

in south central Utah, and discussed the adaption of this species of a cold-adapted

group to survival in a warm climate. He proposed that the subelytral space,

which is enlarged in this species through vaulting and the reduction in size of the

hind wing, serves as a humidifying chamber in contact with the abdominal

spiracles and documented this proposal with a slide showing subelytral condensate.

The following notes were presented:

Oviposition Behavior of Templemania (Tortricidae).—J. A. Powell

exhibited kodachrome slides and specimens of T. apertana (Walsingham) and

T. sarothrura (Felder), illustrating their bizarre oviposition habits. The moths

92 THE PAn-PAcIFIC ENTOMOLOGIST

were observed near Iturbide, Nuevo Leon, Mexico, during September, 1975. The

eggs are deposited in an imbricate patch, as in Archipini and Sparganothidini,

and are then surrounded with a ring of multiple rows of large scales placed on

end, like a miniature picket fence. The scales originate from the huge caudal

scale tufts on the female abdomen that are characteristic of members of the

Tribe Anacrusiini. This behavior was reported for “Aesiocopa” patulana (Walker)

by Beebe (1947, Zoologica, 32:147) in Venezuela, and was observed for patulana

again by Powell at Fortin de las Flores, Veracruz, this year. Thus it appears that,

like other tribes of Tortricinae, a special mode of egg laying will prove to be

a good biological characteristic for Anacrusiini as a whole—J. A. Powe 1,

University of California, Berkeley.

Antennal Deformities in Collops (Coleoptera: Malachiidae).—Of approx-

imately 14,000 specimens of Collops spp. examined, seven specimens with antennal

deformities were found. The deformities were fusion of two antennal segments,

reduced number of antennal segments, and duplication and bifurcation of segments.

The deformities occurred in both males and females of Collops bipunctatus, a

female Collops blandus, and an unidentified female Collops. The altitudes these

specimens were collected at range from 5000 to 7000 feet. The greater exposure

to higher levels of ultra-violet radiation at high altitudes may be the cause of

these teratologies. The ratio of 1:2000 (deformed specimen to normal specimens)

is eight to twenty-five times higher than published estimates of the occurrence of

such deformities—A. I. Kapitan, University of California, Berkeley.

The main speaker of the evening was Dr. Howell V. Daly, Department of

Entomology, University of California, Berkeley, who presented his Presidential

address entitled, “The Africanization of South American Bees”.

Coffee and refreshments were served in the Trustees Room following the

meeting.—F. Ennik, Secretary.

HONORED MEMBERS OF THE PACIFIC COAST

ENTOMOLOGICAL SOCIETY

M. H. Hatch A. E. Michelbacher

H. B. Leech R. C. Miller

C. P. Alexander E. G. Linsley

VoL. 52, No. 1, January 1976

PACIFIC COAST ENTOMOLOGICAL SOCIETY

STATEMENT OF INCOME, EXPENDITURES AND

CHANGES IN FuND BALANCES

Years Ended September 30, 1975 and 1974

Income

Dupes Garih Sierra GS a a et cease elmer

Reprints and miscellaneous —.

aE SSE Sderot te: Bes a ee oe A So ae Ee

Interest on savings accounts —_

Dividends, American Telephone & Telegraph Co. —.

Increase (decrease) in value of capital stock

of American Telephone & Telegraph Co. —

Expenditures

Publication costs—Pan Pacific Entomologist —

Derren ay 8 were We Taw 2 No

Increase (decrease) in fund balances

Fund balances October 1, 1974 and 1973

Fund balances September 30, 1974 and 1973

STATEMENT OF ASSETS

September 30, 1974 and 1975

Cash in bank

Clomitiiereierl eceowmiet eee

Savings accounts

Ee yotelg: | ra 714.5 Aika Ot OTL, ESE A) Ree, eee OT

Mignierr rine eT Ve oy i! Pee ol ie ees ee

aL Peete le eh ee a eS aS ee a Be

Investment in 80 shares of American Telephone

& Telegraph Co. common stock (Life Membership

and Fall Funds), at market value

See accompanying notes to the financial statements.

93

1975 1974

$ 5,728 $ 5,516

6,700 7,377

640 1,888

1,019 744

279 241

390 (880)

14,749 14,886

14,748 8,013

291 92

2,939 1,977

17,978 10,082

(3,229) 4,804

26,839 22,035

$23,610 $26,839

1975 1974

$ 770 $ 7,908

12,035 8,861

2,582 2,457

2,526 2,404.

2,027 1,929

19,940 23,559

3,670 3,280

$23,610 $26,839

94, THe Pan-Pactric ENTOMOLOGIST

NOTES TO THE FINANCIAL STATEMENTS

Year Ended September 30, 1975

SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES

Accounting Method

Income, expenditures and assets are recorded on the cash basis of accounting.

Marketable Securities

American Telephone & Telegraph Co. common stock is carried at market value.

Increases and decreases in value are reflected in income.

Income Tax

The Company is exempt from Federal income and California franchise tax.

Accounts Receivable

As of September 30, 1975 accounts receivable aggregates $871 as follows:

August and September, 1975 billings $800

June and July billiags 2 17

Pa RS oe oe tt Sr ts a em eee te o4

$871

As Chairman of the Auditing Committee, and in accordance with its by-laws, I

have reviewed the financial records of the Society.

During the course of this review nothing was noted which indicated any inaccuracy

in the foregoing statements.

H. Vannoy Davis

Chairman of the Auditing Committee

VoL. 52, No. 1, January 1976 95

SCIENTIFIC NOTE

Observation of Pleocoma hirticollis vandykei Linsley in Santa Clara

County, California (Coleoptera:Scarabaeidae).—On 29 October, 1974 two

Pleocoma hirticollis vandykei females were collected south of San Jose, Santa Clara

County, California. The soil was adobe clay with coarse ground gravel; the

dominant vegetation was Baccharis pilularis (Coyote Bush) and pasture grass.

The specimens were collected at 6:00 pm (just dark) on the evening following

the second rain of the season. The weather was cool, humid, and overcast.

At dawn 30 October four males were collected using one of the females as an

attractant. Both captured females copulated with one of the males. This male

copulated a total of four times, three times with the same female. The male

showed immediate signs of stimulation upon placement in the container with

one of the females. The male extended his aedeagus and approached the female.

The female buried her anterior half into the sand and opened her vaginal

orifice. The orifice was a distinct white contrasting color against her brown

abdomen and elytra. Copulation was complete in less than 60 seconds. One of the

females lived in sand at room temperature until 12 August 1975, without depositing

eggs.—R. E. Pace anv J. W. FisHer, San Jose State University, San Jose, California

95192.

THE PAN-PACIFIC ENTOMOLOGIST

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January 21, 1975. Price $3.00. (Plus 25¢ postage.) *

* (Add 6% sales tax on all California orders (residents of Alameda,

Contra Costa and San Francisco counties add 642%). Members of the

Society will receive a 20% discount.

Send orders to:

Pacific Coast Entomological Society

c/o California Academy of Sciences

Golden Gate Park

San Francisco, California 94118