Vol. 53 APRIL 1977 No. 2

THE

PAN-PACIFIC ENTOMOLOGIST

POINAR — Observations on the kelp fly, Coelopa vanduzeei Cresson in Southern

Californiatbipteras Goeloprdae).. 20... 2. ele ae ee oh oe 81

CHENG — The Elusive Seabug Hermatobates (Heteroptera).....................-. 87

TIDWELL and PHILIP — A New Bolbodimyia from Mexico’s Central Plateau (Diptera:

MabaniGae)esarcryvs stews, eee sc eee epee Ocha, fier cee. Seles, lRtshai's oslo. oeeee cas caches 98

MAYER and JOHANSEN — Cautharidin from Meloe niger Kirby (Coleoptera:

Metoidac) gigi: Moa dir a wet CR ee ee Pn. le ee 2 101

YOUNG — Notes on the Biology of Hypothyris euclea in Costa Rica (Lepidoptera:

Nymphalidae). Seek ele SE cae a POR, Sem cee yl chi Ly cake 104

LANGSTON and MILLER — Expanded distribution of earwigs in California

(Denniapte ra) memes eee. eee tee cos ain Me ON oe os scala sheces oo yn tees ees 114

HARTMAN and HYNES — Biology of the range crane fly, Tipula simplex Doane

(Diptera aU GAS) Meera Pk cts evar epee re, cca Mineo e := geen Meeebens’ « ailebayegeceust’s 118

CHEMSAK — New Neotropical Tillomorphini in the genus Jetranodus Linell

(ColeopteraGerambycidae)k t..c . 4 54 oe retin atte ee ae ince Ss are 124

SIEGFRIED et al. — The Adults of Oroperla barbara (Needham) (Plecoptera:

REMOGIGGe)Y sic. ....gtcee ie vec siis, ome sheers AEN tay S of ores GARE Sue "TOSS aid arehe gy Saree 126

POTTS — Revision of the Scarabaeidae: Anomalinae 3. A Key to the Species of

Anomala of America North of Mexico (Coleoptera)................00000eee 129

PINTO — Descriptions of the First Instar Larvae of Three Species of Epicautine

Blister Beetles (Coleoptera: Meloidae) ............ 0... cece eee eee eee eee 135

SMETANA — A New Species of the Genus Cryptop/eurum Muls. from Central America

(Coleopteray Hydrophilidae)23 5.5.0: Se hes 2 ee ae) ne oe ne ee 142

DAILEY — Elevation of Loxaul/us brunneus variety atrior (Kinsey) to full species status

(ymenopteravGynipidaeyc, 2ecn8 Natalee ean Sa toaee i: Spee, uals scree 145

PAULSON and GARRISON — A List and New Distributional Records of Pacific

CoastOdoniatar. tr. Ay: Gets o-.choncsapehbehe se cae ierece of pMllue foe Wand oles Gaal 147

ZOO LO GIGAIENOMENGEATURES® «01 sic eels on ns cee enti oe en eels aus es 86

SCIENMEIGIN@MES wre. cceet ek. eee ac yan aves MBAS: Selec, eae 103, 113, 123, 144

ERD UNO REA Ua NO UG Eee, lm ee Sik ahh, Meteora vers cet, -, «oe MN Lae, SAW. an. Choa. aes 146

PISMO FUG ARUN Oily een ge ce wis cet eet eRe ence UNM crates eee aay oo -otc UBM ea nS 117

CORRECT O Nig soe ben cpa smears Ct ariefots in. ake AN BEC EE tere Dirge cab ang Mee «Swe lade 97

SAN FRANCISCO, CALIFORNIA ¢ 1977

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

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Publication #419440

The Pan-Pacific Entomologist

Vol. 53 APRIL 1977 No. 2

Observations on the kelp fly, Coelopa vanduzeei Cresson

in Southern California

(Coelopidae: Diptera)

George O. Poinar, ur.

Division of Entomology and Parasitology, University of California, Berkeley 94720

The kelp fly, Coelopa vanduzeei Cresson, occurs along the Pacific

coast from Baja California to Kodiak, Alaska (Cole, 1967). This fly

becomes numerous on beaches in southern California in late summer

and causes general annoyance to bathers and others frequenting the

seashore. Adult flies swarm over stranded kelp on the beach and

literally darken the sand with their numbers. Observations made ona

beach in southern California during the summer and winter months

show this fly to have a behavioral pattern different from that reported

in other parts of California (Kompfner, 1974).

Materials and Methods

Field observations were made during the months of August,

December and January (1974 and 1975) on a San Diego County beach

located in the town of Solana Beach, California. The beach is com-

posed almost entirely of sand which abuts against steep sandstone

cliffs. Large kelp beds which occur just offshore supply a continual

source of wrack on the beach. The kelp forming the majority of the

wrack beds are the giant kelp, Macrocystis pyrifera; the elk kelp,

Pelagophycus porra; and the feather-boa kelp, Egregia sp.

Mature larvae and puparia of C. vanduzeei were observed directly on

the beach and collected for laboratory studies by washing infested

beach sand through window screen. Adults were collected with a net

and maintained in cages with washed beach sand.

Results

All stages of C. vanduzeei were found in or under the kelp during

the study period. The adults probably occur on the beach throughout

The Pan-Pacific Entomologist 53:81-86. April 1977.

82 THE PAN-PACIFIC ENTOMOLOGIST

Fig.1. Small clump of kelp harboring developing larvae of C. vanduzee/. Note adults of

C. vanduzeei on the sand adjacent to the seaweed.

Fig.2. Adults of C. vanduzeei clustering on seaweed.

VOL. 53, NO. 2, APRIL 1977 83

the year, although their populations in winter are much lower than in

late summer. Whereas larvae and puparia are common in December,

many adult flies appear to be in a state of diapause. During the day,

they can be found clustering under seaweed at the high tide level,

and during the night many occur in crevices or protected areas on the

sandstone cliffs. During the month of August, the flies emerge overa

1-2 week period and literally cover the beach, not only clustering on

washed-up seaweed,: but alighting indiscriminately on sand and

bathers (Figs. 1 & 2). They were especially attracted to bird droppings

and marine invertebrates that were washed ashore.

The white, elliptical eggs were deposited singly or in small groups

in the kelp piles (Fig. 3). The females generally sought out the moist

seaweed at the bottom of the wrack piles. Eggs were usually only

deposited on seaweed at the high tide level and rarely on kelp

stranded in the middle or lower tide zone. This may be because the

eggs are only lightly attached to the seaweed and are easily washed

off. However, they hatch very quickly, often within 24 hours at temp-

eratures reached at the study site (90°F). The newly hatched larvae

may tunnel through the fleshy parts of the seaweed or simply grove

the outside portion of the plant, as is typical of the older larvalinstars.

In August, the larvae prefer to feed on kelp covered with 1-2 inches

of sand, probably because the exposed portions quickly desiccate in

the hot, dry climate. This also serves to protect them from being

washed out to sea. During December and January, the larvae can be

found feeding within the exposed kelp mass. The mature third instar

larvae collect together in clusters just under the surface of the sand

near the kelp piles. When the latter are turned over and the first few

inches of sand removed, the sand literally moves with maggots. Their

presence is indicated by the activities of shore birds such as the

western willet, Catopthrophorus semipalmatus and marbled godwit,

Limosa fedoa, which will come to the high tide level or even above to

feed on larvae of C. vanduzeei. Other enemies of larval kelp flies were

mainly staphylinid beetles, especially Cafius seminitens. This predator

was extremely abundant and could be collected in large numbers by

digging acylindrical hole (830 cm deep) in the sand near the fly larvae.

Amphipods were also found associated with larvae of C. vanduzeei.

The larvae pupate in the upper few centimeters of sand and the

adult flies emerge in 5-6 days. Most mature iarvae collected in August

ranged from 8-10 mm in length. The puparium is usually dark brown

and about 5 mm long. The waves wash them out of the sand where

they collect conspicuously on the beach (Fig. 4). The adult flies range

from 3.5 - 5.0 mm in length, and in the fall began mating soon after

emergence. At least a portion of the adult winter population appears

to be in diapause. The adults were restricted to the immediate beach

area and were never recovered from refuse near adjacent homes:

84

Fig. 3.

Fig. 4.

THE PAN-PACIFIC ENTOMOLOGIST

Eggs (arrow) of C. vanduzeei attached to seaweed.

Mature third-stage larvae and puparia of C. vanduzeei.

VOL. 53, NO. 2, APRIL 1977 85

Discussion

It is interesting that Egglishaw (1960) reported largest numbers of

Coelopa frigida and C. pilipes in England during the autumn and winter

months. This was associated with the comparatively larger masses of

wrack washed up on shore in the fall. The situation is reversed in

southern California. Aside from_the increased temperature, there are

two other factors which probably play a role in the summer’s produc-

tion of large numbers of flies. Large kelp beds just off shore are com-

mercially harvested throughout the summer. This results in masses

of kelp fragments which are washed on the beach and provide food

for the flies. Also, from time to time during the summer months, the

wrack is pushed aside by bulldozers to maintain a clean swimming

area. In so doing, large piles of seaweed which become half buried in

the sand above or at the high tide level provide an ideal breeding

habitat for C. vanduzeei and other flies. The adults then become a

general nuisance to people, alighting on their bodies and are some-

times seen around the eyes of children. It is known that adult kelp

flies from several coast sites in Australia carry a virus. Scotti et al

(1976) comment that this virus appears most common in winter when

the adult flies are gregarious. The host range of this virus has not

completely been determined.

The importance of C. vanduzeei and other kelp flies in ridding the

beaches of seaweed should also be mentioned. Egglishaw (1960)

mentioned that stalks of the seaweed, Laminaria that were not at-

tacked by larvae of C. frigida, withered and dried up, but stalks with

feeding larvae quickly decomposed, indicating that larval activity

greatly helped decomposition of the wrack. Thompson in Egglishaw

(1960) found that larvae of C. frigida did not survive well if only a few

were present, and suggested that numerous larvae may be necessary

to provide an attractive feeding environment. This may explain, in

part, why Coelopid flies are usually seen in large numbers.

Kompfner (1974) reported that in Monterey Bay, California, C.

vanduzeei occupied the lower beach wrack banks. The present

studies showed that during August, most activity occurred in the kelp

piles at the upper beach banks. This variation could result from the

different physical environment of the two beaches, however it would

be interesting to know if behaviorial races of this kelp fly species

occur.

Acknowledgments

The author thanks Mr. and Mrs. H. Baltz for providing living facil-

ities during these investigations. Thanks are also extended to W. N.

Mathis, of Oregon State University for identification of the flies and to

W. G. Evans, of the University of Alberta for identification of the

staphylinid beetles.

86 THE PAN-PACIFIG ENTOMOLOGIST

Literature Cited

Cole, F.R. 1969. The Flies of Western North America. Univ. Calif. Pres, Berkeley. 693 pp.

Egglishaw, H. 1960. Studies on the family Coelopidae (Diptera). Trans. Roy. Entomol.

Soc. London, 112:109-140.

Kompfner, H. 1974. Larvae and pupae of some wrack dipterans on a California beach.

Pan-Pac. Entomol., 50:44-52.

Scotti, P.D., A.J. Gibbs and N.G. Wrigley 1976. Kelp fly virus. J. Gen. Virol., 30:1-9.

ZOOLOGICAL NOMENCLATURE

ANNOUNCEMENT A.N. (S) 101

The required six months’ notice is given of the possible use of

plenary powers by the International Commission on Zoological No-

menclature in connection with the following names listed by case

number: (see Bul. Zool. Nom. 33,parts 3 & 4, 31 March 1977).

Z.N.(S.)2157 Goniurellia Hendel, 1927 (Insecta, Dip-

tera, TEPHRITIDAE): designation of

type-species.

Z.N.(S.)2168 Siphonophora Fischer, 1823 (Bryozoa),

status of: Siphonophora Brandt, 1837

(Diplopoda, Polyzoniida), validation of.

Z.N.(S.)2170 Pieris castoria Reakirt, 1867 (Insecta,

LEPIDOPTERA): proposed suppression.

Z.N.(S.)2173 Culex loewi Giebel, 1862 (Insecta, Dip-

tera, CULICIDAE): request for suppress-

ion so as to conserve Toxorhynchites

brevipalpis Theobald, 1901.

Comments should be sent in duplicate (if possible within 6 months

of the date of publication of this notice), citing case number to:

R.V. Melville,

The Secretary

International Commission on Zoological

Nomenclature

c/o British Museum (Natural History)

Cromwell Road,

LONDON SW7 5BD,

England.

Those received early enough will be published in the Bulletin of

Zoological Nomenclature.

The Elusive Sea Bug Hermatobates'

(Heteroptera)

Lanna Cheng

Scripps Institution of Oceanography, University of California, La Jolla, 92093

Although the genus Hermatobates was first established in 1892,

the nine known species have hitherto been represented by only 14

type specimens and the females of only two species have been des-

cribed (Table 1). In no case has a species been described from more

than three specimens. Several sea-going entomologists have sought

these elusive sea-bugs, but few have been successful and it is quite

apparent that these insects are rare. Although clearly distinct from

other known marine hemipterans as a generic entity, Hermatobates

was included in the family Gerridae by all the earlier authors, al-

though, its affinities to the other genera are unclear. Matsuda (1960),

in his monograph of the World Gerridae, excluded Hermatobates

from the family on the basis of several important structural differ-

ences, notably the completely fused meso- and meta-thorax in the

male, the highly modified meso-notum with lateral lobes in the

female, the three-segmented tarsus of all the legs, the granulated

appearance of the eyes, and the presence of a scent gland opening

on the dorsal surface.

The insects are certainly very different from the Gerridae and

should be treated as a separate family (Andersen and Polhemus,

1976). However, until we can establish the systematic importance of

various characteristics it is difficult to decide the phylogenetic rela-

tionships of Hermatobates to other aquatic Heteroptera.

Systematics and Review of Literature

Carpenter (1892), who discovered these insects, described the

genus Hermatobates on the basis of a single male specimen, which

he named H. haddoni, collected from coral reefs off the Australian

coast. Since his specimen was very different from all other known

gerrids, he suggested assigning it to a special subfamily (Hydro-

metridae in his paper). A second species, H. djiboutensis, was added

by Coutiére and Martin (1901a), who in the same year (1901b) des-

cribed a third, Hermatobatodes marchei, and created a new subfamily,

Hermatobatinae, to include these two genera (1901c). Bergroth (1906)

found that these two genera were distinguished merely by sexual

differences, since the two earlier species were only known from the

males, and therefore synonymised Hermatobatodes with Hermato-

bates. H. marchei was synonymised with H. haddoni (Esaki, 1947)

but the earlier name was later resurrected by China (1957), who des-

‘This paper was originally accepted for publication in Pacific Insects (see Cheng, 1976); it has been with-

drawn owing to printer’s problems at that journal.

The Pan-Pacific Entomologist 53:87-97. April 1977.

88 THE PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Hermatobates hawaiiensis male, dorsal view. Fig. 2. Hermatobates hawailiensis fe-

male, dorsal view (scale bar = 1 mm).

cribed three more species, H. hawaijensis, H. walkeri, and H. weddi

(1956, 1957),and constructed a key to the six known species (1956).

Two more species, H. breddini and H. tiarae, were added by Herring

(1965) and yet another was described by Cheng (1966, 1969), bringing

the total number of Known species in the genus to nine. Since most

of the species were described from one or two type specimens (see

Table 1), and substantial intraspecific variations were found among

specimens examined in this study, it is impossible to construct a

meaningful key to the species at present.

Distribution

The recorded localities of all nine Hermatobates species are shown

on Map 1. They are widely distributed, with representatives collected

from the Indian, Pacific, and the Atlantic Oceans. However, six of the

nine described species are known from only the type localities. Of

the remaining three, H. dj/iboutensis has been collected from Djibouti

and the Maldive Islands (Phillips, 1959), H. hawaiiensis appears to be

confined to the Hawaiian Islands, whereas H. haddoni has been re-

ported from Troughton Island (14° 45’S, 125° 10’E, Arafura Sea;

Walker, 1893; Carpenter, 1901), Monte Bello Island (China, 1957), the

Ryukyu Islands (Esaki, 1947), and Tahuata in the Marquesas Islands

(China & Usinger, 1950) in addition to its type locality (Mabuiag

Island). The Tahuata record was based on only one nymph, of which

the specific identity has been questioned by Herring (1965), as has

that of the specimens from New Caledonia and the Ryukyu Islands.

In an earlier paper, Esaski (1935) himself expressed uncertainty as to

the identity of the specimens, but he later decided that they belonged

to H. haddoni (1947). Since at that time no other Pacific species had

VOL. 53, NO. 2, APRIL 1977 89

been described, and since all other known Hermatobates species

are rather restricted in their distribution, we cannot verify the

presence of this species in the Ryukyus and New Caledonia until

Esaki’s specimens can be reexamined or until other collections from

these localities become available. The other records of this species,

from the Marquesas, Palmyra and Christmas Islands (Herring, 1965;

China, 1956),were all nymphs, hence their specific identity could not

be ascertained either.

More recently some specimens of Hermatobates have been col-

lected from Low Isles, Australia, but they have not been identified

to species (Marks, 1971). During several recent collecting expeditions

specimens of Hermatobates were collected from localities where it

has not been previously reported: Enewetak Atoll, Fiji, Tonga, New

Caledonia, Heron Island and Magnetic Island off the coast of Queens-

land, Australia, and Pulau Salu off Singapore. In Enewetak, Fiji and

Tonga adults attracted to lamps at night were netted as they skated

towards the light. They were caught at or shortly before low tide.

Collections at Enewetak were made at Japtan and Medren Islands

from dinghies anchored near wooden piers several hundred meters

offshore, in the vicinity of live corals. In Fiji, the insects were caught

near the Marine Science Institute, University of South Pacific, Suva,

beside a pier encrusted with barnacles and surrounded by coral

rubble. In the main harbour of Nuku Alofa, Tonga, some specimens

were caught when attracted to a light hung over a stone pier 3 meters

wide, several hundred meters long, about 100 meters from shore and

level with the reef flat; others were caught by random sweeps under

an overhanging shelf near the light on the leeward side of the pier;

none were caught from the windward side. These collections

probably represent yet other new species which will be described at

a later date.

General Structure

All Known specimens of Hermatobates, like those of Halobates,

are wingless. The bodies of adults are dark brown or black, and mea-

sure 2.5 to 4mm in length and about 1 to 2 mm in width. The nymphs

are brown, but otherwise are similar to the adults in general structure.

The body and all the legs are covered with fine hairs. The head is ex-

tremely short and more rounded anteriorly than that of Ha/obates,

giving the insect a somewhat oval shape (Figures 1 & 2). The eyes

have a granulated appearance owing to-the pronounced convexity

of the individual ommatidia, whereas in Ha/obates the outer surfaces

of the lenses are rather flat and form a continuous surface. The Her-

matobates eyes further differ from those of Ha/obates in posessing

long, thin inter-ommatidial hairs or setae, similar to those on other

areas of the head of the insect (Figure 3). Such eye setae have been

observed in some terrestrial insects by Hinton (1970), who suggested

90 THE PAN-PACIFIC ENTOMOLOGIST

Fig. 3. Surface of eye showing inter-ommatidial hairs (stereoscan electron

micrograph scale bar = 100um).

that they are simply extensions of the cephalic receptor system,

and do not obstruct incident light normal to the ommatidia. In some

beetles such setae may help to make the eyes less conspicuous to |

predators (Hinton, 1970), but what function these setae may serve in

the sea bug is unknown.

The pronota of these insects are very short. In the male, the meso-

and meta-nota are completely fused and extend posteriorly to cover

several of the fused anterior abdominal segments (Figure 1). In the

female, some of the anterior abdominal segments are exposed be-

tween the lateral lobes of the mesonotum (Figure 2).

The front legs of the males of several species of Hermatobates

are evidently modified for grasping, and bear one or more large, tooth-

like tubercles in addition to smaller teeth. The tarsi of all the legs

are three-segmented, although the first segment is very short. This

character further distinguishes adult Hermatobates from other

Gerridae, although during the nymphal stages the tarsi are not seg-

mented. All distal tarsal segments bear strong claws, which are sub-

apical on the front tarsus but apical on the mid and hind tarsi (Figure

4). Such claws, on both sexes, suggest that these bugs could easily

cling to rocks or other objects. The middle legs are about the same

length as the hind legs which are held almost at the tip of the

abdomen, while in Halobates the middle legs are about 1.5 times as

long as the hind legs, which are held halfway up the abdomen.

VOL. 53, NO. 2, APRIL 1977 91

Fig.4. Stereoscan electron micrograph of mid-tarsal claws (scale bar = 50 um).

Biology

Our knowledge on the biology of these sea-bugs has hitherto been

based on only three field observations, by Walker (1893), Esaki (1947)

and Usinger & Herring (1957). Both Walker and Esaki found them

together with other marine insects on coral reefs which are sub-

merged at high tide. Walker found his specimens of Hermatobates

under a dead TJridacna shell, where there was also a spider; some

specimens of the ocean-strider Ha/obates were also found, in nearby

salt pools left by the receding tide. Esaki found his Hermatobates

at low tide ‘walking’ on the coral reefs or moving on the surface of

tide pools in a manner rather different from that of other gerrids. He

did not see them at high tide, and assumed that they hide in crevices

or under shells. They were found together with many Halovelia septen-

trionalis (Veliidae), an unidentified collembolan and an unidentified

staphylinid beetle. He also noted that marine midges (Clunio paci-

ficus and C. setoensis) were abundant nearby. The types of H. wedd/

and H., walkeri were both collected from coral reefs; those of H.

; a | 40°

Hermatobates sp % A .

ingapore Apishigaki I.

Qe oy ERED

H. marche! SE 2 N oot | A2oe

Philippines Palmyra I.

Christmas I. 0°

Marquesas I.

New A

H weddi \, Galedonia H tiarae 0

Monte Bello I, vy p Tamall Arch. a

H. wa/ker/ Ao?

Arafura Sea Lpeeant

Mabuiag I.

20° 60° 100° 140° 180° 140° |OQO0° 60° 20°

Map 1 Distribution of Hermatobates spp. showing type localities (0) and other collecting records for H. haddoni(A). -

c6

LSIDOTOWOLNA DlSIDVd-NVd SHL

VOL. 53, NO. 2, APRIL 1977 93

breddini and H. tiarae were collected at sea under lamps, as were the

specimens collected off the Singapore coast.

Our first daylight glimpse of Hermatobates was in Noumea, New

Caledonia, where these insects appeared as small silvery objects

moving extremely fast over the surface of the water amongst Sar-

gassum weed half exposed at low tide. Individual specimens ap-

peared very suddenly and disappeared with equal suddenness. The

first was caught after much patient waiting and fast chasing, but,

once we discovered what to look for, several additional specimens

were captured on two subsequent days at low tide, when the water

was calm. However, no insects came to the light of lanterns hung

over the water at high tide in the same locality at night.

On Heron Island we found adults as well as nymphs of Hermato-

bates at low tide during daylight hours in the afternoons, but we en-

countered none at our lights at night (when the winds were higher

and the sea surface choppy). Adults were found skating over fast-

flowing rivulets of ebbing seawater between large boulders at the

ridge of the reef-flat, i.e., at the outer edge of the reef where the

boulders are first exposed at receding tides. To our great surprise,

we also found nymphs of various stages clinging to the undersides

of such boulders, associated with Ha/ovelia, Collembola, staphylinid

beetles and mites. As the water ran off the overturned rocks many of

the insects began to move, while others remained motionless, en-

closed in air-bubbles. It is possible that normally, as the tide recedes

and the boulders became exposed, such nymphs will crawl out from

their hiding places to feed, and indeed we found several of them

moving over the surface of small rock pools when the tide was at its

lowest level, though they became rarer and disappeared as the tide

began to rise. None were seen at high tide, when presumably the

insects would have crawled under boulders and survived enclosed

in air bubbles until the next low tide. We could not directly observe

them to do so, since such small insects (less than 2mm in size) can-

not be watched in the splashing waters of an incoming tide. When

living specimens were set in a laboratory aquarium supplied with

dried pieces of corals and then artificially flooded, they simply rose

to the surface and swam or rested with bunched up legs on the water.

When bits of dead Jubipora musica (organ-pipe coral) were supplied

as substrate, the insects readily crawled into the tubes even without

flooding. On Magnetic Island, specimens of Hermatobates were col-

lected in conditions more or less similar to those on Heron Island.

All the collections were made at low tide during the day: here, too we

were unable to do any night collecting.

From such observations we deduced that the Hermatobates col-

lected previously at night lights on the open ocean were ‘strays’, the

natural habitats of these sea-bugs being coral rubble, as indicated

by Esaki (1947), and rocks in the low intertidal. At low tide they come

out to feed on small animals among intertidal rocks and algae, and at

94 THE PAN-PACIFIC ENTOMOLOGIST

Fig.5. Portion of thorax, showing microtrichia (stereoscan electron micrograph, scale

bar = 10um).

high tide they hide in crevices or under boulders, enclosing them-

selves in air bubbles much in the same way as do intertidal veliids

and saldids (Andersen and Polhemus, 1976; Polhemus, 1976). If,

however, they cannot find a resting place under water before the tide

rises, they can swim or rest on the water surface for many hours.

We were able to keep specimens in the laboratory on seawater for

more than 5 days without their showing any ill effects. It is possible

that in some areas, and at times when weather conditions are rough,

the incidence of ‘straying’ on the sea surface can be quite high.

Among surface plankton tows made in open waters off Hawaiian

shores, in the course of a study of larval fish, 23 samples contained

Hermatobates hawaiiensis: in one sample, 8 insects had been netted.

in less than 10 minutes (Cheng, unpublished data.).

Discussions

Hermatobates appears ideally suited to an intertidal coral reef

existence. It has large eyes, presumably adapted for vision in air

rather than in water (Figure 3); strong claws for clinging to algae-

covered rocks (Figure 4); and a well developed plastron (Figures

VOL. 53, NO. 2, APRIL 1977 95

Fig. 6. Microtrichia at higher magnification (stereoscan electron micrograph, scale

bar = 5yum).

5 & 6) similar to that of freshwater gerrids (Cheng, 1973). The strongly

developed front femora, found in males of several described species,

are probably adaptations for clinging to the females. Their legs, in

contrast to those of Halobates, are more adapted for walking over

rocks than for skating over water. Although they can skim extremely

quickly over water, their movements are mainly confined to short

dashes as they skate from boulder to boulder; presumably they do

not travel over large expanses of water as do Halobates (Cheng, 1974).

When chased they often make for the nearest rock, cling to it and

remain stationary; under such circumstances they are very hard to

see, Halobates is unable to do so. They can also jump well; adults

leaping from a water surface reach a height of 5-6 cm.

Since these insects may have to come out and feed at low tide

during rain storms, experiments were carried out on the effects of

reduced salinity on their activities. We detected no difference in the

behavior or viability of insects kept on seawater or on freshwater over

a 24-hour period.

Their ability to withstand submersion was tested by keeping

insects fully immersed in boiled (deoxygenated) and unboiled sea-

water. In the former, they ceased to move within 10-15 minutes; if

96 THE PAN-PACIFIC ENTOMOLOGIST

Table 1. Type specimens of Hermatobates spp. with type localities.

Species Author, Year No., Type Type Locality

breddini Herring, 1965 1 male Woodbridge Bay, Dominica

British West Indies

djiboutensis Coutiére & Martin, 1901a 1 male Djibouti, Red Sea

haddoni Carpenter, 1892 1 male Mabuiag Island, North Australia

hawailiensis China, 1956 1 male, Coconut Island, Hawaii

2 females

marchei Coutiére & Martin, 1901b 1 male Honda Bay, Philippines

singaporensis Cheng, 1976 1male, Singapore

2 females

tiarae Herring, 1965 1 male Tuamoto Archipelago,

French Oceania

walkeri China, 1957 2males Arafura Sea, N.W. Australia

weddi China, 1957 1 male Monte Bello Islands

then released and blotted dry, they regained their mobility in 10-15

minutes. If left in boiled seawater for more than 30 minutes, however,

they died. Insects submerged in ordinary seawater remained mobile

for at least 6 hours; when then released and dried, they became fully

-active in 10-15 minutes. Nymphs appeared to withstand submersion

much better than adults, remaining mobile under water for longer

periods of time and, when released, regaining mobility within shorter

periods of time.

Although our knowledge of the biology of these elusive sea bugs

is still rather fragmentary, now that their normal habitats are known

(i.e., coral rubble or reefs of tropical island shores) and the best times

and methods for capturing them have been established, they may not

remain so apparently rare or elusive for long.

Acknowledgments

! wish to thank the Mid-Pacific Marine Laboratory, University of

Hawaii, for providing travel funds and living and research facilities

at Enewetak; the Roche Institute of Marine Pharmacology, Australia,

for providing accommodation and research facilities on Heron Island;

the Department of Marine Science, James Cook University, for

laboratory facilities; and Ralph A. Lewin for assistance in the field.

‘Literature Cited

Andersen, N. M. & J. T. Polhemus. 1976. Water-striders (Hemiptera: Gerridae, Veliidae,

etc.). In Marine Insects (ed.) L. Cheng, pp. 187-224. North-Holland, Amsterdam.

Bergroth, E. 1906. Systematische und synonymische Bemerkungen Uber Hemipteren.

Wiener Ent. Z. Vienna. 25: 1-12. ;

Carpenter, G. H. 1892. Reports on the zoological collections made in Torres Straits

by Professor A.C. Haddon, 1888-1889: Rhyncota from Murray Island and

Mabuiag. Proc. R. Dublin Soc. (new ser.) 7: 137-146.

VOL. 53, NO. 2, APRIL 1977 97

Carpenter,G.H. 1901. Theinsects of the sea — VI. Knowledge 24: 245-248.

Cheng,L. 1966. Acurious marine insect, Hermatobates. Malay. Nat. J. 19(5) 283-285.

Cheng, L. 1973. Marine and freshwater skaters: differences in surface fine structures.

Nature 242(5393) 132-133.

Cheng, L. 1974. Notes on the ecology of the oceanic insect Ha/obates. Mar. Fish. Rev.

36(2): 1-7.

Cheng, L. 1976. A new species of Hermatobates (Hemiptera: Heteroptera). Pan-Pacific

Entomologist 52: 209-212.

China, W. E. 1956. A new species of the genus Hermatobates from the Hawaiian Is-

lands (Hemiptera-Heteroptera, Gerridae, Halobatinae), Ann. Mag. Nat. Hist.

ser. 12, 9(101): 353-357.

China, W.E. 1957. The marine Hempitera of the Monte Bello Islands, with descriptions

of some allied species. J. Linn. Soc. Lond., Zool. 40(291): 342-357.

China, W. E. & R. L. Usinger. 1950. Hermatobates haddoni Carpenter from the Mar-

quesas Islands (Hemiptera: Gerridae). Proc. Haw. Ent. Soc. 14: 53.

Coutiére, H. & J. Martin. 1901a. Sur un nouvel Hémiptére.halophile, Bull. Mus. Nat.

Hist. Paris. 4(s6r. 1): 172-177.

Coutiére, H. & J. Martin. 1901b. Sur un nouvel Hémiptére halophile, Hermatobatodes

marchei, n.gen.,n. sp. Bull. Mus. Nat. Hist. Paris. 5: 214-226.

Coutiére, H. & J. Martin. 1901c. Sur une nouvelle sous-famille d’Hémiptéres marins,

les Hermatobatinae. C. R. Acad. Sci. Paris. 132: 1066-1068.

Esaki, T. 1935. Insect fauna on the coral reefs in the Yaeyama Islands. Zool. Mag.

(Dobutugaku Zassi) 47: 140-141 (in Japanese).

Esaki, T. 1947. Notes on Hermatobates haddoni Carpenter. Mushi 18(7): 49-51.

Herring, J. L. 1965. Hermatobates, a new generic record for the Atlantic Ocean, with

descriptions of new species (Hemiptera: Gerridae). Proc. U.S. Nat. Mus. 117:

123-129.

Hinton, H. E. 1970. Some little known surface structures. [In] Insect Ultra-structure

(ed.) A. C. Neville, pp. 41-58. Blackwell Sci. Publ., Oxford.

Marks,E.N. 1971. Australian marine insects. Aust. Nat. Hist. 17: 134-138.

Matsuda, R. 1960. Morphology, evolution and a classification of the Gerridae (Hemip-

tera: Heteroptera). Univ. Kansas Sci. Bull. 41(2): 25-632.

Phillips, W. W. A. 1959. Notes on three species of marine Hemiptera taken in Addu

Atoll, Maldive Islands, between October, 1958, and April, 1959. Ent. Mon.

Mag. 95: 246-247.

Polhemus, J. T. 1976. Shore bugs (Hemiptera: Saldidae, etc.) [In] Marine Insects (ed.)

L. Cheng, pp. 225-262. North-Holland, Amsterdam.

Usinger, R. L. & J. L. Herring. 1957. Notes on marine water striders of the Hawaiian

Islands (Hemiptera: Gerridae). Proc. Haw. Ent. Soc. 16(2): 281-283.

Walker, J. J. 1893. On the genus Halobates, Esch., and other marine Hemiptera. Ent.

Mon. Mag. 29: 225-232.

Correction — Hippomelas Plant Associations

In a recent article on this subject (Pan-Pacific Entomologist,

52:272-285), the authors quote G.H. Nelson (in /itt.) (p. 279) as confirm-

ing that Hippomelas planicauda Casey has been consistently taken on

“Acacia.’’ This should have read ‘“‘Mimosa’’ as the data in the preced-

ing paragraph indicate. — E.G. Linsley.

A NEW BOLBODIMYIA FROM MEXICO’S CENTRAL PLATEAU

(Diptera: Tabanidae)

M.A. Tidwell

Apartado Aereo 5390 Cali, Colombia

and

Cornelius B. Philip

- California Academy of Sciences San Francisco 94118

Three species of the New World genus Bolbodimyia are known

presently to occur in Mexico including B. atrata (Hine) from west

central Mexico and Arizona, B. dampfi Philip from southern Mexico

and Guatemala and the more recently described B. /Jampros Philip

and Floyd (1974) from Chihuahua.

An adult female of a fourth species, distinct from those included in

Stone’s (1954) and Fairchild’s (1964) papers, was reared from a larva

collected in El Chico National Park on Mexico’s Central Plateau

approximately 100 Km northeast of Mexico City. The description of

this female and the pupais given below.

We are pleased to name this species for the collector, Luis

Bermudez, who with his wife Ema, was of special assistance to the

senior author during his stay in Mexico.

Bolbodimyia bermudezi, new species

A robust, entirely black-bodied species with black vestiture except for conspicuous

pre-alar patches of silky pale yellow hairs, face and genae golden orange, and wing

subhyaline with a sharply contrasting black costal border nearly to apex. Scapes shining

black, expanded ventrally.

Holotype female, 15mm in length. Head flattened, a little wider than thorax. Eyes

pilose, background color dark green with lower 2/3 containing numerous shining reddish

irregular spots. Frons (Fig. 1a) nearly bare, height slightly more than 2 times basal width,

blackish, expanded below, with a shining black, swollen basal callosity filling slightly

less than the lower third and scarecely connected on the lateral margins to a broad black

median callus above. Lateral margins of frons, area between basal and median calli and

upper margin of median callus with fine gray pollinosity. Lateral margins of frons from

upper portion of basal callus to vertex with reclining balck hairs directed mesally. No

ocelli. Subcallus bare, shining black, inflated. Face and genae golden orange pollinose

with concolorous hairs. Antennae (Fig. 1b) blackish with basal portion of plate somewhat

lighter in colo#; scape shining black and swollen below with numerous stout bristles;

pedicel with a dorsal tooth; plate approximately 1/4 longer than style. Palpi (Fig. 1c) black

with concolorous hairs.

Body entirely subshining black with black hairs except for pre-alar patches of pale

yellow hairs. Legs black including tarsi, tibiae not noticeably swollen, hind tibial fringes

black, not accentuated. Wing (Fig. 1d) subhyaline with contrasting black costal border a

little more expanded posteriorly than in related B dampfi, reaching vein R 4 +5, and apical

hyaline crescent smaller. Vein R4 strongly curved forward. Basicostas bare. Halteres dark

brown.

Type locality. — MEXICO: Hidalgo, El Chico National Park, approximately 20 Km north

of Pachuca, 8 June 1974. Luis and Ema Bermudex. Holotype in the California Academy of

Sciences. No. 12785. Collected as larva.

The Pan-Pacific Entomologist 53:98-100. April 1977.

VOL. 53, NO. 2, APRIL 1977 99

Fig. 1. Bolbodimyia bermudezi, female. a. Frons. b. Antenna. c. Palp.d. Wing. Fig. 2.Pupa

of B. bermudezi. a. Frontal plate. b. Terminal aster.

The adult female is readily differentiated from females of B. dampfi by the distinctly

pilose eyes, presence of the golden-orange pilosity of the face, the lighter colored pale

yellow pre-alar patches not extending on to the pleura, tarsi basally and antennal plates

darker; the first basal cell (cell R) of wing mostly subhyaline and lacking the complete

infuscation as found in B. dampfi.

100 THE PAN-PACIFIC ENTOMOLOGIST

Critical differences preclude thisbeing the unknown, possibly dichromatic female of B.

lampros including black not reddish flagella, prealar lobes but not upper pleura yellow,

and wing infuscation restricted to costal areas in contrast to remainder of wing.

The larva of B. bermudezi was collected in moss on rocks in running water. The

collectors reported that the substrate was made up of an association of mosses,

Polytrichum sp.; liverworts, Marchantia sp.; and Cieraceae, Cyperus spp. The principal

trees and plants in the area includd fir, Abies religiosa; Umblliferae including Hydrocotyle

ranunculoides; and various Compositae.

The adult emerged 2 March 1975. The pupal case was preserved in excellent condition;

however, the larval exuvium was not recovered.

Pupa. — Length 21 mm, frontal plate (Fig. 2a) with area between antennal sheaths

strongly inflated; antennal ridges prominent, separated by a median cleft; height of ridge

at cleft approximately 0.5 mm; each ridge subdivided by an indentation, the median

portion larger with a heavily sclerotized, unusually pointed crest; frontal ridges distinct;

callus tubercles prominent, 0.4 mm high on lateral margins tapering mesally; antennal

sheaths reaching well beyond epicranial suture; anterior and posterior orbital tubercles

prominent, the posterior tubercles more so. Thoracic spiracles 0.4 mm, elongate C-

shaped. Fringes of abdominal segments 2-7 with well-developed spines, progressively

longer on posterior segments, length of spines on dorsum generally about 0.4 mm long

on segment 2, grading to about 1.0 mm on segment 7; shorter spines present on all

segments usually situated slightly anterior to longer spines. Terminal segment (Fig. 2b)

with dorsal and lateral combs continuous totaling 18 and 20 spines including minute

spines; ventral combs with 7 and 10 spines.

The pupal case was compared with 2 female pupal cases of B. astrata from Arizona

provided through the courtesy of Dr. John F. Burger, University of New Hampshire. B.

bermudezi is easily separated by having the area between the antennal sheaths more

inflated and antennal ridges at cleft nearly twice the height of those in B. atrata. Frontal

ridges present and well developed; callus and orbital tubercles prominent with longer

spines.

The immature stages of B. atrata will be presented in a paper by Burger in press at this

writing.

Literature Cited

Fairchild G.B. 1964. Notes on Neotropical Tabanidae (Diptera) IV. Further new species

and new records for Panama. J. Med. Entomol., 1:169-185.

Philip C.B. and L. Floyd. 1974. New North American Tabanidae XXI. Another new Bo/bodi-

myia from Mexico. Pan-Pacific Entomol., 50: 145-147.

Stone, A. 1954. The genus Bo/bodimyia Bigot (Tabanidae, Diptera). Ann. Entomol. Soc.

Amer., 47: 248-254.

Cantharidin from Meloe niger Kirby

(Coleoptera: Meloidae)

D. F. Mayer and C. A. Johansen!’

Department of Entomology, Washington State University, Pullman 99163

Meloidae are the source of the medicinal drug cantharidin, com-

monly called Spanish fly. It has been generally assumed that all

Meloidae with the possible exception of the tribe Horiini contain

cantharidin (Selander, 1960; Pinto and Selander, 1970). However,

much of the early literature is inaccessible and some of the early

identifications are ambiguous. Actual analytic data are available

for only about 33 of about 2,000 species, most of them belonging to

the genera Epicauta and Mylabris (Dixon et a/., 1963; Carrel and Eisner,

1974.)

During a 4-year study of Meloe niger Kirby, a predator of the alkali

bee, Nomia melanderi Cockerell, we tested for cantharidin in adult

beeties. We believe this is the first positive analysis for cantharidin

from a North American species of Meloe.

Methods and Procedures

Adult M. niger were collected from the field during March and April,

1976 and transported to the laboratory alive.

Blood samples obtained from reflex bleeding were treated accord-

ing to Carrel and Eisner (1974), and tested for cantharidin. Whole

adults were ground in dry ice with a mortar and pestle and extracted

with chlorform according to a method modified from Dixon et. al.

(1963).

Samples were injected into a Hewlett-Packard 5700A gas chro-

matograph. Two columns were used: a 1.83 m (6 ft) coil of 10% sili-

cone rubber SE-30 on chromsorb W Aw 80-100 mesh, and 2) a 1.52 m

(5 ft) coil of 5% OV1 gaschrom Q. A oven temperature of 200°C. was

used.

Results and Discussion

The presence of cantharidin in M. niger was confirmed by gas

chromatography-mass spectrometry from both blood samples and

whole beetles with peaks corresponding to authentic pure cantha-

ridin (Supplied by Inland Alkaloid Co.). Blood contained 57 mM and

whole beetles an average of 0.88 mg (0.12% of body weight) cantha-

ridin.

The only previous, authenticated record of cantharidin in Meloe is

Dixon ef. a/. (1963) in the European Meloe proscarabeus L. at 2mg

(0.187% of body weight). Interestingly, Shimano et. a/. (1953) tested

' Work conducted under Washington Agricultural Research Center Project No. 0147, Scientific Paper

No. 4680.

The Pan-Pacific Entomologist 53:101-103. April 1977.

102 THE PAN-PACIFIC ENTOMOLOGIST

for cantharidin in the Asiatic spp. Epicauta gorhami Marseul and

Meloe auriculatus Marseul, finding it in the former but not the latter.

Westwood (1839) also states that some Mel/oe are not servicable

in medicine (/.e. do not contain cantharidin). Apparently, all Meloe

do not contain cantharidin.

When disturbed, Meloidae respond by reflex bleeding, which con-

sists of the emission of blood from the leg joints while the beetle

remains immobile (death feigining). Male M. niger emit a maximum of

1 to 2 ul and females 8 to 10 ul. Male blood is a distinct red color and

female blood a light yellow. Reflex bleeding is thought to be a non-

glandular chemical defense mechanism with cantharidin, which is

toxic to vertebrates and insects, functioning as a feeding deterrent.

We have never seen reflex bleeding or death feigning by M. niger

under natural conditions, but have observed it in the laboratory by

hitting adult Deeties a sharp rap with a probe and in the field when

collecting and handling the beetles.

The general assumption is that vertebrates associate the delayed,

noxious effects of cantharidin with the insect eaten and learn to dis-

criminate against it (Carrel and Eisner, 1974). There is very little

specific evidence of predation of Meloidae (Selander, 1960; Pinto and

Selander, 1970) and we have not observed any predation on adult

M. niger. In feeding tests, Meloidae have been refused as food with-

out first becoming sick by baboons and falcons (Marshall, 1902);

monkeys (Carpenter, 1921); lizards (Pritchell, 1902) and green sunfish

(Tafanelli and Bass, 1968). This would suggest the ‘“‘becoming sick-

learning to discriminate’ mechanism is not operating. A possible

mechanism, suggested by Eisner (1970), is a chronic debilitating

effect on predators into a species from which the tendency to cap-

ture the lethal prey is entirely selected out. This would call for a high

order of selection pressure. .

When placed near an ant colony (Formica obscuripes Forel), most

adult beetles were able to escape without: feigning death, though

they were attacked by the ants. In one instance, we held the beetle

among the ants until they punctured the beetle’s abdomen. Ants that

came in contact with blood from the puncture immediately backed

away and began cleansing activities. The ants eventually succeeded

in killing the beetle, but they did not use it for food. Carrel and Eisner

(1974) found this same type of cleansing response by Pogonomyrmex

occidentalis Cresson from contact with the blood of Epicauta brunnea

Werner. They also established that cantharidin is a feeding deterrent

to some insect predators and not others.

Meloid blood, due to the cantharidin, causes redness, pain, and

blistering when applied to the human skin. We have not found this

reaction in 10 humans that have contacted the blood of M. niger.

Cantharidin in M. niger blood may be tightly bound to lipoproteins

and therefore may not enter the human skin. The concentration of

VOL. 53, NO. 2, APRIL 1977 103

cantharidin in M. niger blood is 57 mM, and this greatly exceeds the

solubility of cantharidin in water. Alternatively, M. niger blood may

contain a pharmacologically active principle which counteracts the

effects of cantharidin.

Literature Cited

Carpenter, C.D. H. 1921. Experiments on the relative edibility of insects with special

reference to their coloration. Trans. R. Entomol. Soc. Lond., 69: 1-106.

Carrel, J. E., and T. Eisner. 1974. Cantharidin: potent feeding deterrent to insects.

Science, 183: 755-757.

Dixon, A. F. G., M. Martin-Smith, and S. J. Smith. 1963. Isolation of cantharidin from

Meloe proscarabeus. Can. Pharm. J., Sci. Sect., 96: 501-503.

Eisner, T. 1970. Chemical defense against predation in arthropods, p. 157-217. /N

Sondheimer, E., and J. B. Simeone (ed.), Chemical Ecology. Academic Press,

New York.

Marshall, G. A. K. 1902. Five years’ observations and experiments (1896-1901) on the

bionomics of South African insects chiefly directed to the investigation of .

mimcry and warning colours. Trans. R. Entomol. Soc. Lond., 50: 287-584.

Pinto, J. D., and R. B. Selander. 1970. The bionomics of blister beetles of the genus

Meloe and a classification of the new world species. Univ. Ill. Biol. Mono.

42, 222 pp.

Pritchell, A. H. 1903. Some experiments in feeding lizards with protectively colored

insects. Biol. Bull., 5: 271-287.

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

blister beetles (Coleoptera, Meloidae). Univ. Ill. Biol. Mono. 28, 295 pp.

Shimano, T. M., M. Mizujo, and T. Boto. 1953. Cantharidin and free amino acids in

Epicauta gorhami and similar insects. Ann. Proc. Gifu Coll. Pharm., 3: 44-45.

(Chem. Abstr. 59: 133089).

Tafanelli, R. J., and J.C. Bass. 1968. Feeding response of Lepomis cyanellus to blister

beetles (Meloidae). Southwest Natur., 13: 51-54.

Westwood, J. O. 1839. An introduction to the modern classification of insects,

vol. 1. London.

SCIENTIFIC NOTE

Note on the Distribution and Host Relationship of Idiomelissodes duplocincta (Cock-

erell) in Mexico (Hymenoptera: Apoidea). — On 9 and 10 September 1976 | collected bees

from blossoms of Ferocactus sp. (probably F. wis/lizenii (Engelm.) Britt. & Rose) growing

in ornamental plantings on the campus of the Unidad Noroeste del Instituto Tecnologico

y de Estudios Superiores de Monterrey. The site is located 14 km. south of Ciudad

Obreg6n (ca. 27° 29’ N, 109° 56’ W) in the irrigated and heavily farmed Yaqui Valley of

Sonora. | made collections between 1130 and 1200 on both days, and on September 10, |

took one female of the little Known eucerine bee, /diomelissodes duplocincta (Cockerell)

from an open blossom of one of the cacti.

Idiomelissodes duplocincta has been recorded in Mexico from Chihuahua and Coahuila

by LaBerge (1956, Univ. Kansas Sci. Bull. 37: 911-1194) and from Baja California Sur by

Zavortink (1975, Pan-Pacific Entomol. 51:236-242). Zavortink reported the ecological

relationship of this deserticolous bee to Ferocactus wislizenii on the basis of observations

made by him in Arizona and New Mexico.

My collection is the first record of /diomelissodes duplocincta in the Mexican state of

Sonora, and it helps to confirm the use of Ferocactus as a pollen source by this bee.

Thanks are due to Dr. Juan M. Mathieu, Director of the Unidad Noroeste at Ciudad

Obreg6n, for his kind hospitality during my visit to that institution and to Dr. Wallace E.

LaBerge of the Illinois Natural History Survey at Urbana for confirming the identity of the

bee. — JOHN K. BOUSEMAN, //linois Natural History Survey, Urbana, Illinois 61801.

The Pan-Pacific Entomologist 53:103. April 1977.

Notes on the Biology of Hypothyris euclea in Costa Rica

(Lepidoptera: Nymphalidae: Ithomiinae)

Allen M. Young

Department of Invertebrate Zoology, Milwaukee Public Museum, Milwaukee, Wisconsin 53233

Although the local species density of ithomiine butterflies in tropi-

cal forests can be high (Brown, 1972), it is generally unusual to find

very large adult populations of a single species. However, some

ithomiines, such as Hypothyris euclea euclea (Latreille) in Trinidad,

have seasonal bursts of synchronous eclosion (Barcant, 1970), which

could lead to the sudden appearance of large numbers of adults over

a short period of time at a locality. This paper reports the biology of

Hypothyris euclea leucania (Bates) (tribe Napeogenini) at one locality in

northeastern Costa Rica, and emphasizes that the larvae of this

butterfly are gregarious defoliators of So/anum rugosum Dund.

(Solanaceae) during the early dry season. The preliminary data on de-

foliation of this food plant suggests that heavily defoliated plants

produce fewer flowers and fruits.

Habitat and Methods

Clumps of mature S. rugosum create a canopy of about three

meters high in young secondary forest (2-10 years old) at ‘Finca La

Tirimbina’, near La Virgen (220 m elev.), Heredia Province, Costa

Rica. The locality is in the Premontane Tropical Wet Forest life zone

(Holdridge, 1967). Previous observations revealed that S. rugosum is

the food plant of H. euc/ea at this locality; other food plants have not

been found. Therefore, | began studies on the biology of H. euc/ea in

the field and laboratory, using methods of previous studies (e.g.,

Young, 1974). Most observations were made between January 12 and

February 12, 1976, and finally on March 31, 1976. Included in these

studies was the estimation of defoliation by counting heavily fed

upon) in four large clumps (Patches 1-4). | also counted the number

of individuals in each clump bearing flowers or fruits (Solanum

rugosum flowers and sets fruit during January and February, months

of erratic dry spells which precede a period of more uniform dryness

(March) ). The four clumps of S. rugosum were visited a total of eleven

days during the period of January 12 to February 12. As little is known

about the life cycle of this species, oviposition and early stages were

also observed.

Results

General Biology: Oviposition activity is high during January, and

rafts of oblong white eggs are placed on the ventral sides of mature

leaves throughout the day (Fig. 1). From January 12 to February 12,

The Pan-Pacific Entomologist 53:104-113. April 1977.

VOL. 53, NO. 2, APRIL 1977 105

Fig. 1. Life cycle of Hypothyris euclea /leucania. First column, top to bottom: female in

oviposition, raft of eggs, first instar caterpillars. Second column: second instar

caterpillars and a third instar caterpillar.

more than 50 oviposition acts were observed in Patch 2, and of about

5,000 eggs counted in different clumps at various times (4 days)

during this period, 70% were found in Patch 2. A raft of eggs has from

40 to 90 eggs, and an egg is deposited at five-second intervals until a

raft is completed. During January and February, 73 egg rafts were dis-

covered in Patch 2. Oviposition by other ithomiines on this plant was

not observed during the study period. Four to five days after eggs are

106 THE PAN-PACIFIC ENTOMOLOGIST

Fig. 2. Life cycle of H. euc/ea. Left: prepupa and lateral view of pupa; Right: dorsal

view of pupa, and freshly-eclosed adult in the field.

laid, they produce greenish-yellow translucent larvae with shiny black

heads; second instar larvae are dark green, and third instars possess

a striped color pattern of light blue, dark gray, and yellow (Fig. 1). The

latter color pattern also occurs in the fourth and fifth instars, and a

translucent light green prepupa produces a golden pupa (Fig. 2). The

adults appear about 22 days later, bearing the familiar orange, yellow,

and black ‘‘tiger stripe’ pattern characteristic of some species in the

tribe Napeogenini (Fig. 2). Adults produced from a single cluster of

VOL. 53, NO. 2, APRIL 1977 107

\

eggs may exhibit some color variation within sex (Fig. 3). In north-

eastern Costa Rica, as in eastern Brazil (Brown and D’Almeida, 1970),

Hypothyris is sympatric with several other ithomiine genera at forest

edge habitats.

Adults are slow fliers of shady forest edge and light gap habitats,

where they feed on a variety of resources, including the partly eaten

remains of insects, presumably attacked by birds and other small

vertebrates (Fig. 3).

A distinctive behavioral feature of the larvae is their gregarious

habit. Young larvae, recently hatched from an egg raft, stay together

on a leaf, but as they grow, they split up into smaller groups on dif-

ferent mature leaves. Survival from predators and parasites seems

high, although short periods of heavy rain often result in large num-

bers of larvae dying.

Gregarious Defoliation: The cluster oviposition habit of H. euclea,

in which the number of eggs per raft is much higher than for

Mechanitis (another ithomiine with gregarious larvae), results in the

gregarious larvae becoming severe defoliators (Fig. 4 and 5). The de-

foliation often results in a ‘skeleton canopy” (Fig. 6).

Food plant clumps with little or no defoliation bear large numbers

of flowers and fruits. In fact, in unattacked shrubs, 30 to 60 clusters of

healthy fruit occur on each shrub: of a total of 23 shrubs examined,

the mean number (and standard deviation) of fruit clusters is 41 +

19.5, and the mean number of fruits per cluster is 32 + 11.2. Inflores-

cences are abundant during late January, and green fruits are abun-

dant by February. Thus, the infestation of H. euclea is intense

(January and perhaps earlier) when the S. rugosum population is

entering a period of reproduction. In the four widely separated

clumps studied, although canopy height and branch density are very

similar, clumps of shrubs not heavily defoliated have far more flowers

or fruits than individuals in clumps of severe defoliation by H. euclea

larvae (Table 1). Although the sample size is small, the four clumps

examined are sites where this species is very abundant. While leaf

damage is severe in these clumps, by March 31 the H. euclea popula-

tion had either experienced a severe reduction in size or else

dispersed, as very few adults, eggs, and caterpillars were present,

and previously attacked shrubs had fresh leaves. During January and

February there are probably only two or three overlapping generations

present on S. rugosum.

Discussion

Although it is reported that Mechanitis is the only ithomiine genus

exhibiting gregarious behavior of the immature stages (Rathcke and

Poole, 1975), it is apparent that Hypothyris also possesses this be-

108 THE PAN-PACIFIC ENTOMOLOGIST

Fig. 3. Top: a series of laboratory-reared H. eucilea (left column — 2 females; right

column — male above, female below). Bottom: Hypothyris eucilea feeding on the leg of a

grasshopper in forest understory, Finca La Tirimbina, Costa Rica, February 1976.

havioral trait. In fact, an interesting comparison of the early stages

can be made between Hypothyris as a representative of the

Napeogenini and Mechanitis as a representative of the Mechanitini.

VOL. 53, NO. 2, APRIL 1977 109

Fig. 4. Heavy defoliation of mature leaves of Solanum rugosum by second instar cater-

pillars of H. euciea at Finca La Tirimbina (January 1976).

Fox (1967) has reported the early stages and gregarious larvae of

Mechanitis. Several differences are noted: (1) the larvae of Hypothyris

lack the lateral tubercles and uniform coloration of Mechanitis larvae,

(2) egg size and size of egg rafts are larger in Hypothyris than

Mechanitis, (8) the pupa of Hypothyris is short and thick in profile,

while that of Mechanitis is elongate and thin, and (4) groups of gre-

garious larvae are considerably larger in Hypothyris than in Mechanitis.

Furthermore, Mechanitis lays eggs on the small, heavily-spined

species of Solanum (Fox, 1967), differing from S. rugosum, in terms

of size, profile, leaf texture, and lack of spines (in S. rugosum).

Examinations of about 100 plants of So/anum spp. attacked by

Mechanitis isthmia at Finca La Tirimbina during February 1976 (A.M.

Young, pers. obs.) revealed that in no instances were individual

plants (usually widely scattered) heavily defoliated, and mortality of

egg rafts was very high. These biological traits are different from the

interaction of H. euclea with S. rugosum at the same locality. It shares

with Mechanitis the characteristic of being very abundant locally,

unlike most other ithomiines. In addition to biological traits such as

cluster oviposition, gregariousness of larvae, and an abundant food

110 THE PAN-PACIFIC ENTOMOLOGIST

Fig.5. Defoliation by fourth instar caterpillars of H. eucilea.

plant, Hypothyris may be experiencing local population expansions,

perhaps similar to that noted for Mechanitis in Brazil (Brown and

D’Almeida, 1970).

The gregarious behavior of the larvae, and the bright coloration of

caterpillars and adults, suggest unpalatable properties in this butter-

fly. The ‘“‘tiger-stripe” ithomiines are classical unpalatable models for

mimicry complexes (Bates, 1862; Miller, 1878; Brown and Neto, 1976),

including some species of Hypothyris.

Perhaps, in addition to intensifying the advertisement of distaste-

ful properties to vertebrate predators, cluster oviposition and the

gregarious behavior of larvae have important consequences for the

food plant. Barcant (1970) reports that H. euc/ea adults are abundant

at certain localities in Trinidad at the wet season’s onset, but that

numbers dwindle subsequently. Birch (1957) argues that local

climatic factors play major roles in determining the abundance of

some insects. A period of prolonged and intense rainfall in the

tropics promotes the growth of leaves, while the dry season

promotes the flowering and fruiting of some tree species (Janzen,

1967). Thus, by the end of the wet season, the local plant community

is characterized by a large leaf biomass, providing a reliable and

VOL. 53, NO. 2, APRIL 1977 111

abundant food base for some herbivorous insects. Given other suit-

able environmental conditions, populations of herbivorous insects

might become high by the end of the wet season and early dry season

in seasonal tropical habitats. The Premontane Wet Forest life zone in

Costa Rica is characterized by alternating bouts of dryness and wet-

ness, but the longest succession of wet days occurs between

October and December, and the longest period of dry days occurs in

March. Thus, by early January, many plants may have allocated most

of their energy to vegetative growth. For H. euclea on S. rugosum, a

wet period resulting in a large potential food base for larvae followed

by several short series of dry days that optimize courtship and ovi-

position promotes rapid growth of the adult population. The last two

weeks of December (1975) were exceptionally dry (Dr. Robert Hunter,

personal communication), and these conditions promoted reproduc-

tive activity in H. euc/ea and other butterflies. But, during January and

February, although there are many days of complete dryness, there

are 3-5 day wet periods that promote mass larval mortality and re-

duced oviposition. Some studies have shown that adult butterflies

may even be killed by bursts of intense rainfall (e.g., Cook et al., 1971).

Although adults of H. euc/ea appear to be slow fliers, it seems likely

that, if necessary, breeding should extend to all clumps of S. rugosum

if the population is resource-limited at this time of the year. However,

this does not happen and some clumps escape from heavy defolia-

tion. Those clumps that are intensely fed upon show a reduction in

fruit set. Solanum rugosum flowers and sets fruit during the short,

erratic dry season, as do trees in more seasonal regions of Costa Rica

(Janzen, 1967; Frankie et al., 1974). Prior to, and during this period

there is intense reproductive activity by the butterfly. The progressive

reduction of host leaf biomass due to larval feeding during January

Tabie 1. Patterns of heavy leaf attack by caterpiilars of Hypothyris

euci/ea and flowering and fruiting in four patches* of Solanum rugosum

in premontane tropical wet forest near La Virgen, Heredia Province,

Costa Rica, February 12, 1976.

No. trees with No. trees with No. trees with No. trees with

Patch fiowers/ fruits flowers/ fruit no flowers/ fruit no flowers/ fruit

No. and attacked and not attacked and attacked and not attacked

1 0 21 0 7

2 1 15 0

3 0 2 4 4

4 0 16 0 5

*The four patches are widely separated along a dirt road that connects Finca La Tirimbina to the Penal

Colony at Magasay. Only trees 2-3 meters tall were included in the census, although there were trees less

than 2 meters tall that experienced heavy defoliation: Patch No. 1-0, Patch No. 2-8, Patch No. 3-0, Patch

No. 4-4. Patches were of similar size, each occupying about 5-6 meters of roadside secondary forest.

112 THE PAN-PACIFIC ENTOMOLOGIST

Fig.6. Defoliated bushes of S. rugosum.

and February, the specificity to a single host plant, the shift in alloca-

tion of energy to flowers and fruit, and the mortality of larvae are

factors promoting a reduced breeding population of H. euc/ea by late

March. 7

eh

Acknowledgments

This study is a by-product of National Science Foundation Grant

GB-33060, and a travel grant from the Milwaukee Public Museum. The

assistance of Dr. Kenneth Starr of the Milwaukee Public Museum is

appreciated. | thank Dr. Lee D. Miller (Allyn Museum of Entomology)

for identifying the butterfly, and Sr. Luis Poveda (Museo Nacional de

Costa Rica) for identifying the host plant. The cooperation of Dr. J.

Robert Hunter for logistical Support and facilities at Finca La

Tirimbina is appreciated. | thank Cheryl Castelli for typing the manu-

script.

Literature Cited

Barcant, M. 1970. Butterflies of Trinidad and Tobago. London: Collins Press.

Bates, H. W. 1862. Contributions to an insect fauna of the Amazon Valley, Lepidoptera:

Heliconidae. Trans. Linn. Sci. London 23:495-566.

VOL. 53, NO. 2, APRIL 1977 113

Birch, L. C. 1957. The role of weather in determining the distribution and abundance of

animals. Cold Spring Harbor Symp. Quant. Biol., 22:203-218.

Brown, K. S., Jr. 1972. Maximizing daily butterfly counts. J. Lepid. Soc. 26:183-195.

Brown, K.A., Jr., and R.F. D’Almeida. 1970. The Ithomiinae of Brazil (Lepidoptera:

Nymphalidae). Il. A new genus and species of Ithomiinae with comments on

the tribe Dircennini D’Almeida. Trans. Amer. Ent. Soc. 96:1-17.

Brown, K. S., Jr., and J. V. Neto. 1976. Predation on aposematic ithomiine butterflies by

tanagers (Pipraeidea melanonota). Biotropica 8:136-141.

Cook, L. M., K. Frank, and L. P. Brower. 1971. Experiments on the demography oftropical

butterflies. 1. Survival rates and density in two species of Parides. Biotropica

3:17-20.

Fox, R. M. 1967. A monograph of the Ithomiidae (Lepidoptera), Part Ill, the tribe

Mechanitini Fox. Mem. Amer. Ent. Soc., No. 22, 190 pp.

Frankie, G. W., H. G. Baker, and P. A. Opler. 1974. Comparative phenological studies of

trees in tropical lowland wet and dry forest sites in Costa Rica. J. Ecol. 62:881-

929.

Holdridge, L. R. 1967. Life zone ecology. Tropical Sci. Center, San Jose, Costa Rica.

Janzen, D. H. 1967. Synchronization of sexual reproduction of trees within the dry

season in Central America. Evolution 21:620-637.

Miller, F.1878. Uber die Vortheile der Mimicry bei Schmetterlinge. Zool. Orz. 1:54-55.

Rathcke, B. J. and R. W. Poole. 1975. Coevolutionary race continues: butterfly larval

adaptation to plant trichomes. Science 187:175-176.

Young, A. M. 1974. A natural historical account of Oleria zelica pagasa (Lepidoptera:

Nymphalidae: Ithomiinae) in a Costa Rican mountain rain forest. Studies

Neotrop. Fauna 9:123-140.

SCIENTIFIC NOTE

Biological and Distributional Data for Evergestis angustalis (Lepidoptera: Pyralidae) Ever-

gestis angustalis (Barnes & McDunnough) is a primarily desert species which flies in winter

and early spring and is easily recognized by its exceptionally narrow forewings. The

typical subspecies is known from the western Colorado Desert area of southern Cali-

fornia. Munroe (1973, The Moths of America North of Mexico. Fasc. 13.1C Pyraloidea,

Pyralidae (part). Evergestiinae. pp. 253-304.) indicated that he had seen specimens only

from three localities in western Imperial and eastern San Diego counties in the low desert,

but, in contradiction, stated that angustalis also occurs in the Mojave-Desert. He also

characterized new subspecies from central Arizona and Santa Catalina Island, California.

Records in the Essig Museum of Entomology, University of California, Berkeley, provide

life history data and show that the species is considerably more widespread, occurring

in Baja California Norte and northward into central California.

The flight occurs in January and February in southern California and in Arizona, but

there is one record each for May and July in Arizona (Munroe, 1973). Adults have been

taken at lights elsewhere during early spring: in Baja California Norte, near Santo

Domingo and 5 miles east of El Rosario, Mar. 18, 19, 1972 (Doyen & Powell); and in the

San Francisco Bay area of California, at Alum Rock Park (near San Jose), Santa Clara

Co., Mar. 8, 1960 (S. D. Smith) and at Walnut Creek, Contra Costa Co., Feb. 9, 1972 (J.

Powell).

Larvae were found feeding in hollow stems of Caulanthus inflatus Wats. (Cruciferae)

at Big Panoche Gorge, San Benito-Fresno Co. line, April 21, 1967, and one moth

emerged Jan. 23, 1968 (J. Powell no. 67D88). This plant is a desert species which reaches

its northern limit in western Fresno County and its southern limit in the Mojave Desert

(Munz, 1963, California Flora. U. Calif. Press, Berkeley; 1681 pp). All other collection re-

cords are outside the known range of Caulanthus inflatus and presumably represent popu-

lations of E. angustalis associated with other Cruciferae. — J. A. POWELL, University of

California, Berkeley 94720.

The Pan-Pacific Entomologist 53:113. April 1977.

Expanded distribution of earwigs in California

(Dermaptera)

Robert L. Langston

31 Windsor Ave., Kensington, California 94708

and

Scott E. Miller

Santa Barbara Museum of Natural History, Santa Barbara, California 93105

In continuing studies on the earwigs of California, recent records

of Dermaptera are presented, plus older data, either missed or some-

how not included in current literature.

The earwig fauna of California was recently reviewed by Langston

and Powell (1975). However, due to paucity of museum specimens

and time lag in publication, their distribution in Santa Barbara County

(particularly the Channel Islands), and in parts of the San Joaquin

Valley were not well represented. The detailed California records

indicate the museums and collections where these specimens are

on deposit: California Academy of Sciences, San Francisco (CAS);

California Department of Food and Agriculture, Sacramento (CDA);

California Insect Survey, University of California, Berkeley (CIS);

Los Angeles County Museum of Natural History, Los Angeles

(LACM); Merced County Department of Agriculture, Merced (MDA);

National Museum of Natural History, Washington, D.C. (NMNH);

San Joaquin County Department of Agriculture, Stockton (SJDA);

and Santa Barbara Museum of Natural History, Santa Barbara,

(SBMNH).

Anisolabis maritima (Gén6), the maritime earwig

Recorded in Orange County in 1921, and there are quarantine

records from Los Angeles and San Diego Counties in the 1930’s

and 40’s. Searches were made by both authors in southern California

with no success. Hence, A maritima should remain as only adventive,

or by quarantine south of the San Francisco Bay Area. Additional

specimens have been taken at established colonies in Alameda,

Contra Costa, Marin and Solano Counties, 1973-1976, but no new

locations were found since the detailed records of Langston (1974).

Euborellia annulipes (Lucas), the ring-legged earwig

The records below are numerous, but detailed data were not in-

cluded in Langston & Powell (1975). Therefore, old records are given

for the offshore islands and for some of those more recent than

mapped (op. cit., Map 2). Only one county (Tuolumne) can be con-

sidered new.

California records. — IMPERIAL CO.: Winterhaven, IX-10-70 (R. A. Flock, CDA). LOS

The Pan-Pacific Entomologist 53:114-117. April 1977.

VOL. 53, NO. 2, APRIL 1977 115

ANGELES CO.: San Clemente Island, Barracks, 1d, VI-24-1971 (D. C. Rentz & D. B. Weiss-

man, CAS); Santa Catalina Island, 3 specimens, X-28-31-1908 (O. Marsh, NMNH); Granada

Hills, 19, IIl-27-1975 (R. Sherman, LACM). MERCED CO.: Atwater, 299, VIII-7-1975 (R.

Langston, CAS); Merced, Applegate Park & County Courts Park, several dd & 29: XI-3-

1975 (F. Carl, MDA). SAN JOAQUIN CO.: Lathrop, VI-4-1972, Quarantine from Vietnam

(L. S. Hawkins, CDA); Stockton, 19, VIl-2-1975 (R. Langston, CAS), 19, 4 juv., VII-8-1975

(R. Langston, SJDA), 19, VIl-14-1976 (R. Langston, CAS). SANTA BARBARA CoO.: Santa

Rosa Island, 192, VII-12-1939 (LACM). SOLANO CoO.: Vallejo, shore of Carquinez Strait,

1d, 229, XI-21-1965, 1d, VI-2-1966 (R. Langston, CIS). TUHOLUMNE CO.: Sonora, 19, 2 juv.,

VIIN-25-1975 (R. Langston, CAS).

Euborellia cincticollis (Gerstaecker), the African earwig

Merced and San Joaquin Counties are recorded as new in the data

below, representing naturally occurring established populations.

California records. — IMPERIAL CO.: 3 mi. NW. of Glamis, 2d, 299, IV-1972 (A. Hardy

& M. Wasbauer, CDA). MERCED CO.: Merced, 10, 399, VIII-4-1975, 19, VIII-7-1975 (R. Lang-

ston, CAS). SAN JOAQUIN CO.: Holt, 1d, XII-20-1971 (L. S. Hawkins, SJDA); 1 mi. E. of

Peters, 19, 1 juv., I-7-1972 (L. S. Hawkins, SJDA).

Labidura riparia (Pallas), the shore earwig

Kern and Santa Barbara Counties are recorded as new. The Bakers-

field example is a northerly extension of over 100 miles from the

closest Los Angeles County locale. The Montecito specimen is a

northwesterly range extension from Malibu in Los Angeles County

of about 75 miles.

California records. — IMPERIAL CO.: 3 mi. NW. of Glamis, 100+ dod, 99° & juv. at black-

light, IX-16-1972 (A. Hardy & M. Wasbauer, CDA), 1d, IX-10-1974 (M. Wasbauer & R.

McMaster, CDA). KERN CO.: Bakersfield, 1 adult, VIl-14-1971 (H. Knipp, P. Martin & T.

Tandrow, CDA). LOS ANGELES CO.: Granada Hills, 1d, VI-12-1973, 1d, IIl-18-1975 (R.

Sherman, LACM). RIVERSIDE CO.: Carvina Beach, 12 mi. SE. of Mecca, VII-12-1974 (J.

Doyen, CIS). SANTA BARBARA CO.: Montecito, 19, VII-1968 (W.S. Hull, SBMNH).

Doru taeniatum (Dohrn)

In Langston & Powell (1975) specimens of this species were iden-

tified as Doru lineare (Eschscholtz, 1822). However, the revision by

Brindle (1971) of the genus Doru Burr indicates that the name D.

lineare applies to a species which occurs in Brazil, Paraguay and

Argentina. The species of Doru which occurs commonly from Bolivia

through Central America, and into Mexico and the United States is

D. taeniatum (Dohrn, 1862). Gurney (1972) reviews this and the two

other North American species of Doru. All records of D. /ineare from

Central and North America should be referred to D. taeniatum.

Despite its abundance in southeastern Arizona and Mexico, this

species does not appear to be established in California. Only two

additional records were found, one apparently new for San Bernar-

dino County.

California records. — RIVERSIDE CO.: Blythe, Border Quarantine Station, II-18-1966,

Quarantine from Vera Cruz, Mexico (J. M. Donovan, CDA). SAN BERNARDINO CoO.: Fon-

tana, VI-9-1972, Quarantine on Zea mays (Shurtliff & Bengston, CDA).

116 THE PAN-PACIFIC ENTOMOLOGIST

Forficula auricularia Linnaeus, the European earwig

During recent fieldwork on Santa Rosa Island by the SBMNH, F.

auricularia has been found to be well established. Localities on the

mainland in the Santa Barbara area were mapped as either ‘‘quaran-

tine” or ‘‘adventive” (Langston & Powell, 1975, Map 10). Currently it

is common on the Santa Barbara-Goleta coastal shelf. Likewise,

Hogue (1974) considers the European earwig established in the Los

Angeles Basin. However, there are no established records from

further south along the coast, and the European earwig is unknown

from the Colorado or Mojave Deserts. It is now considered estab-

lished in Merced County and Santa Rosa Island which were not pre-

viously recorded.

California records. — MERCED CO.: Merced, Applegate Park & County Courts Park,

several dd & 29, XI-3-1975 (F. Carl, MDA); Merced, 16th & V Sts., 19, VII-6-1976 (R. Langs-

ton, CAS). SANTA BARBARA CO.: Santa Rosa Island, Arlington Canyon, 2dd, 299, Il-

25-1976 (W. G. Abbott & P. W. Collins, SBMNH), 19, 7 juv., IV-22-1976 (S. E. Miller,

SBMNH); S. Rosa I., ranch headquarters area, 2dd, 499, II-24-1976 (Abbott & Collins,

SBMNH), 399, IV-23-1976 (Miller, SBMNH); S. Rosa I., Wreck Canyon, 14, VII-3-1971 (D. C.

Rentz & D. B. Weissman, CAS).

Quarantine records of Dermaptera in California

In addition to the six foregoing species, four others have been

taken in quarantine by the California Department of Food and Agri-

culture. These have all been determined by Dr. A. B. Gurney, National

Museum of Natural History, Washington, D.C.

Labia curvicauda (Motschulsky) [Labiidae: Labiinae]

This circumtropical species is essentially cosmopolitan. Published records as cited

in Sakai (1970) include the major Ethiopian, Neotropical and Oriental Regions, Africa and

most of the Pacific Ocean Islands. Despite its commonly being found worldwide, it has

been taken in California only once previously — in 1934 in San Diego by quarantine from

Guam.

SAN JOAQUIN CO.: Lathrop, V-4-1972, Quarantine from Vietnam (L. S. Hawkins & K.

Wright, CDA).

Paracosmia toltecus (Scudder) [Forficulidae: Opisthocosmiinae]

This species is Known from the state of Vera Cruz, Mexico and also Guatemala (Sakai,

1973).

SAN DIEGO CO.: Port of San Diego, XI-23-1966, Quarantine from Mexico (C. S. Badman

& R. M. Ireland, CDA).

Skalistes inopinatus (Burr) [Forficulidae: Forficulinae]

Published distribution includes Costa Rica (types), Antigua, Ecuador, Guatemala,

Mexico and Peru (Sakai, 1973).

LOS ANGELES CO.: West Los Angeles, 1 adult on leaf of bromeliad, III-28-1972, Quaran-

tine from Guatemala (F. Cunningham, CDA).

Skalistes vara (Scudder) [Forficulidae: Forficulinae]

The type series is from Puebla, Mexico, with the distribution as cited by Sakai (1973)

including the states of Mexico, Morelos and many from Puebla.

SAN DIEGO CO.: Port of San Diego, XI-23-1966, Quarantine from Mexico (C. S. Badman

& R. M. Ireland, CDA).

VOL. 53, NO. 2, APRIL 1977 Bed

Acknowledgments

The authors thank Waldo G. Abbot, Paul W. Collins, and the Vail

& Vickers Ranch Company for making possible the collection of the

Santa Rosa Island specimens, plus Stephen Newswanger (Santa

Barbara City College Life Science Museum) for providing a significant

Labidura riparia specimen. The help of Alan Hardy (CDA) is appre-

ciated for use of the department files on additional quarantine inter-

ceptions, and for examination of specimens. Thanks are extended

to the following for studying the material under their care and/or

collecting specimens on their own with our encouragement: Kirby

W. Brown (SJDA); Frank Carl (MDA); Charles L. Hogue (LACM); and,

David C. Rentz (CAS).

Literature Cited

Brindle, A. 1971. A revision of the genus Doru Burr (Dermaptera, Forficulidae). Papeis

Avulsos de Zoologia [Sao Paulo, Brazil], 23: 173-196.

Gurney, A. B. 1972. Important recent name changes among earwigs of the genus

Doru (Dermaptera, Forficulidae). U.S. Dept. Agr. Coop. Econ. Ins. Rpt., 22:

182-185.

Hogue, C. L. 1974. The insects of the Los Angeles Basin. Published by the Los

Angeles Co. Museum. : 27.

Langston, R. L. 1974. The maritime earwig in California (Dermaptera: Carcino-

phoridae). Pan-Pac. Entomol., 50: 28-34.

Langston, R. L. and Powell, J. A. 1975. The earwigs of California (Order Dermaptera).

Bull. Calif. Insect Survey, 20: 1-25.

Sakai, S. 1970. Dermapterorum catalogus praeliminaris. Il: Labiidae. Daito Bunka

Univ., Tokyo, 2: 1-177.

Sakai,S. 1973. Dermapterorum catalogus praeliminaris. VIl. Forficulidae. Daito Bunka

Univ., Tokyo, 6: 1-357.

100 Years Ago In Entomology — 1877

ADOLPHE BOUCARD, Coleopterist — was collecting birds and Coleoptera in Costa Rica.

THOMAS LINCOLN CASEY, Coleopterist — was a cadet at West Point.

HENRY EDWARDS, Lepidopterist — was an actor in San Francisco, associated with the

“California Theatre’.

C. R. OSTEN SACKEN, Dipterist — returned to Germany from New York. He returned a

large collection of North American Diptera to the Museum of Comparative

Zoology.

S. W. WILLISTON, Dipterist — was leading an expedition to western Kansas in search of

fossil dinosaurs.

H. G. HUBBARD and E. A. SCHWARZ, Coleopterists — were collecting on Lake Superior.

H. C. FALL, Coleopterist — at age 15 was in public school in Dover New Hampshire. He

collected his first beetle this year.

W.S. BLATCHLEY, Orthopterist, Hemipterist, Coleopterist — at age 18 was a door to door

salesman in Putnam Co., Indiana.

W. J. HOLLAND, Lepidopterist — was in Pittsburgh, where he was a pastor of the Belle-

field Presbyterian Church, Professor of Ancient Languages and Trustee of the

Pennsylvania College for Women.

AUGUST BUSCK, Lepidopterist — was a seven year old child in Randers Denmark.

J.H. MCcDUNNOUGH, Lepidopterist — was born this year.

Biology of the range crane fly, Tipula simplex Doane

(Diptera: Tipulidae)

Margaret J. Hartman

California State University, Los Angeles

and

C. Dennis Hynes

California Polytechnic State University, San Luis Obispo

The range crane fly, 7ipula simplex, described by Doane (1901),

is a univoltine pest of valued rangeland in the San Joaquin Valley.

The larvae live in the grass of unirrigated pastures and when at high

density, will eat the grass roots. Tulare County reported outbreaks

of the species in 1961, 1967, and 1973. This latest outbreak affected

800 hectares on one rach alone, with acrane fly density of 3000 larvae

per square meter. At this high density, the crane fly larvae denude

the hills of all grass and other forage, with an adverse effect on the

watershed. However, such high densities of crane flies occur only

occasionally and ranchers detect no signs of the larvae in intervening

years. Since little information on the biology of the species has been

reported, this study was undertaken to determine the species’ life

history and habits, as necessary prerequisites to understanding

the causes of the population fluctuations. This study was conducted

on unirrigated pasturelands in the foothills of the Sierra Nevada

mountains in northern Tulare County from October, 1974 through

March, 1976, and was supplemented with laboratory rearings and

experiments.

Habitat

In May the soil starts to dry out and by the end of June the soil

contains only 0.3% water by weight. By mid summer the soil mois-

ture has dropped to 0.2% with the temperature one inch below the

surface of the soil exceeding 50° Celsius during the day. The first

rain usually comes in September (about ¥2 inch), stimulating the

growth of turkey mullein or dove weed (Eremocarpus setigeras Benth.),

vinegar weed or bluecurls (Trichostema lanceolatum Benth.), and

tarweed (Hemizonia congesta DC.), but the soil dries out almost com-

pletely before the second rains come (usually in October). After these

second rains the forage crops appear, and the soil remains mois-

tened by periodic rain until April when the rains end, and the soil

starts drying out again.

Adults and Oviposition

The males usually emerge before the females in February or March.

When the female adult emerges the male grasps her and sometimes

The Pan-Pacific Entomologist 53:118-123. April 1977.

VOL. 53, NO. 2, APRIL 1977 119

pulls her out of her pupal case. Copulation occurs immediately, arid

in the laboratory will continue for 24 hours. When the female is re-

leased, she walks and pulls her way through grasses in the field

until she falls into a depression in the ground. These depressions

may be holes caused by loosened clods of soil and rocks, or imprints

of cattle hooves. The female will oviposit all her eggs (average 96,

S = 28.6) in the soil together, approximately % and % inches below

the surface. Neither male nor female have been seen feeding. All

adults in any one field emerge within a three week period.

Eggs

The egg is the stage which must survive the harsh summer condi-

tions of high temperature and low moisture.

In the laboratory we were able to induce hatching by drying the

eggs for six months at a 16:8 photoperiod, then transferring them to

a 10:14 photoperiod. They were kept moist for two weeks, then dry

for one week, and then moist again. The hatching rate was 5.2%

within one week. Hatching could be induced by this procedure any-

time in the fall, but not if the alternate wettings and dryings were

started in January or later. However, an additional 5% would hatch at

age 18 to 21 months old, if the above mentioned wetting and drying

regimen was repeated in September of the second year after ovi-

position.

Eggs are laid in clumps, which makes sampling difficult. Lang

(unpublished data) collected 13 samples (11.5 diameter circle, to a

depth of % inch). From each sample he took 4 samples of 5 grams

each for testing. The number of eggs per 5 gram sample ranged from

0 to 2641, with a mean of 68.48 (S = 174.43). Obviously, sampling

for eggs would require a large number of samples to insure that the

sample mean is similar to the population mean. Hanson’s equation

(1967) allows us to calculate the number of samples necessary to

approximate the population mean (n = t?se?/(x-y)?, where n is the

number of samples, t is the student T statistics for whatever confi-

dence level, s.e. is the standard error, (y-) is the amount of deviation

from the true population mean that the investigator will accept. Using

Lang's data, and calculating 95% confidence limits, we can calculate

that we must take 48 samples to be 95% sure that the sample mean

is within 10% of the true population mean, or 195 samples to be 95%

sure that the sample mean is within 5% of the true population mean.

Larvae

The first instar has never been found in the field. Laboratory rearing

of eggs to hatching indicate that the first instar is morphologically

different from the other three instars. The second instar larvae show

a clumped distribution due to the oviposition habits of the females.

By the time the crane flies are third instar larvae, a different pattern of

120 THE PAN-PACIFIC ENTOMOLOGIST

aggregation is observed. The crane flies are spread evenly through-

out the grass, but are highly aggregated under cowpads that are one

or more seasons old, under pieces of wood, or under any other debris

in the field. For example, in one field, cowpads had 1104 individuals/

meter? (S = 513.3) while in the grass the density was 193 individuals/

meter? (S = 32.9). The degree of aggregation in part depends upon

the dryness of the field. In the very dry winter of 1976 virtually all the

crane flies in the field were concentrated under cowpads (2200 larvae/

meter’).

The fourth instar larvae retain the aggregated pattern. However,

during the late fourth instar (January and February) predation by birds

takes a heavy toll. The birds that are most often seen eating the crane

fly larvae are blackbirds and starlings, although curlews and meadow-

larks will also feed on the larvae in years of high crane fly density.

The most typical mode of feeding is for the birds to flip over one year

old cowpads and feed on the larvae underneath (curlews) or poke

their beaks through the cowpad and eat both the larvae which have

burrowed into the manure and those which live between the cowpad

and the soil (starlings and blackbirds). This predation drastically

changes the distribution pattern. The remaining aggregations are

located under and around older cowpads which have grasses growing

through them, making them less vulnerable to predation by birds.

Predation ceases abruptly with pupation of the larvae. When birds

turn over a cowpad which has both larvae and pupae under it, they

feed only on the larvae.

Pupae

In one week, the percentage of crane flies in the pupal stage rises

dramatically (12% to 82%). Males and females can be easily distin-

guished at this stage (Hynes and Hartman, in preparation). The pupae

make movements in response to light and pressure, but have limited

locomotion. The pupal stage is fairly short. The first adults appeared

in the field 12 days after the first pupae.

Aggregation

The aggregation pattern of early second instar larvae is believed

to be due to the habit of adult females laying all their eggs in the

same place.

The second, third, and fourth instar larvae are much more motile

than the first instar, and it is not Surprising that a change in the distri-

bution pattern occurs. Laboratory tests were performed in 1974 and

1975 to determine the factors influencing the habitat choice of the

older larvae.

The test apparatus was a straight tube (80 x 25 x 4 cm) which was

divided into two equal halves. Different conditions could be main-

tained in each side of the tube. Parameters tested were light (0 versus

VOL. 53, NO. 2, APRIL 1977

11

5.6 lux) and moisture (0 versus 2 gm water on 12.5 cm diameter filter

paper). Tests, when both sides of the cage had identical conditions,

indicated that the larvae showed no preference for either side of the

cage. When animals were given a choice between a moist and a dry

environment, they always spent a significantly greater amount of

time in the moist environment. The response to light was more

variable. If both sides of the cage were moist the animals spent a sig-

nificantly greater period of time in the dark. If both sides of the cage

were dry, the animals exhibited no preference for light or dark, and

moved continually (Table 1).

This indicated a kinesis in response to water. Further tests on

speed of movement indicated that a larva in a moist artificial environ-

ment moved at arate of 0.08 cm/sec and in adry artificial environment

it moved at a rate of 0.16 cm/sec. This was significant using the

paired difference test. In a moist environment the number of head-

turns that a larva made in a given time was not significantly different

than the number of headturns for the same period of time when the

animal was in a dry environment. By definition (Denny and Ratner,

1970), the larvae show an orthohydrokinetic response, that is they

slow down, but do not increase turning, in the presence of soil mois-

ture.

The response to light was also tested by putting the crane fly

larvae into a more natural condition of soil or manure in a cage. Their

response to light above and below them was tested. All larvae moved

down as light was shined from above (6 tests, 4 larvae/test) and 27 out

of 32 larvae moved upward as light was shined from below (8 tests,

4 larvae/test). The difference between the numbers of larvae moving

up to escape light versus the numbers of larvae moving down to

escape light was not significant (Student’s t test). These results

indicate that the crane fly larvae are negatively phototactic, and that

their response to light overrides any response to gravity.

Table 1. Preference for Conditions of Moisture and Light

Test # of larvae Conditions in Conditions in Mean difference in

tested preferred side of non-preferred side time spent intwo

maze (A) and mean of maze (B) and mean sides A-B

minutes spent mintues spent

5 5 dry/dark (19) dry/light (11) 8 min N.S.

6 5 wet/dark (25.2) wet/light (4.8) 20.4**

7 5 wet/dark (27.4) dry/dark (2.6) 24.8**

8 5 wet/light (26.2) dry/light (3.8) 22.4**

9 5 wet/light (24.6) dry/dark (5.4) 19.2**

10 5 wet/dark (25.8) dry/light (4.2) 21.6**

**p>.01

122 THE PAN-PACIFIC ENTOMOLOGIST

Finally, we performed a test to determine if a substance produced

by the crane fly larvae affected aggregation. Ten larvae were placed in

a cage containing moist filter paper (9 cm diameter) for 24 hours, on

which they defecated and left skin traces. This paper containing the

excretions of the larvae and moist filter paper were placed in a large

cage (9 x 10 x 100 cm) containing a layer of moist sand. A number of

fourth instar larvae were then introduced into the cage. After 24

hours, it was found that 94% of the larvae were collected under the

filter paper with crane fly extract and 6% were found under the

control paper. The aggregation index (Roth and Cohen, 1973) for this

crane fly substance is 0.875. (1 is a perfect aggregation).

We hypothesize that aggregation of third and fourth instar larvae

in the field under cowpads or rotten wood is due to the following

factors. The area under the cowpads provides a temperate, moist,

dark habitat. When larvae wander into the area, they slow down

(orthohydrokinesis). The aggregation pheromone <acts either to

attract other larvae to this favorable environment, or by decreasing

the rate of locomotion helps to retard emigration by larvae that have

moved into the area by chance.

Distribution

Tipula simplex has a known distribution from northern Santa Cruz

County north to Marin County, and across to Sacramento (Alexander,

1967). Another population occurs in Tulare County. The area between

Sacramento and Tulare counties may contain the fly, but it has not

been recorded in the literature or found by us in this area. In southern

Tulare County the proportion of Tipula simplex decreases and the pro-

portion of Tipula acuta Doane increases. In Kern County (south of

Tulare), Tiou/a acuta replaces Tipula simplex.

The distribution in fields varies from year to year. In three years we

have mapped the areas of crane fly populations on one ranch. In 1974,

populations of medium density (100-300 larvae/m?) were found in

three fields, LB, SC, and WM and in 1975, very light densities (<50

larvae/m?) were found in the north facing slopes of LCA. In 1976, light

densities were found in WM, and medium densities were found in the

south facing slopes of LCA, SC, and LC. As a general rule, the same

area did not have measureable populations for two consecutive years

(an extensive search did not reveal any Tipula simplex larvae). One

exception to this was field LCA, but the same area was not infested

in the two consecutive years.

Density Measurements

We have made several attempts to accurately measure the crane

fly larval density. The most successful, to date (our estimated mean is

calculated within 6.8% of the true population mean at the 95% confi-

dence limits), involves measuring the number of larvae under 25 cow-

VOL. 53, NO. 2, APRIL 1977 123

pads of measured size, the number of larvae in 25 samples of grass

of similar size, and then estimating the cowpad coverage (which was

determined by the point-centered quarter method) (Cottam & Curtis,

1956). In field LC in 1976, for example, the mean of 25 samples was

equivalent to 159.7 larvae/meter? cowpads (accuracy 6.8%), 273.3

larvae/meter’ grass (accuracy 6.8%) and the cowpad coverage was

1.05% (accuracy 6.2%). The estimated average number of larvae/

meter? was 272.1, and we are 95% confident that the actual popula-

tion mean was between 253.6 and 290.6 larvae/meter? (based on

method of Hanson, 1967). Because this sampling was done after bird

predation, the mean density of crane flies under cowpads is lower

than the mean density of crane flies in grass.

Acknowledgments

We gratefully acknowledge the cooperation of the Agricultural

Commission of Tulare County and the financial assistance of the J.

G. Boswell Company.

Literature Cited

Alexander, C. P. 1967. The crane flies of California. Bull. Calif. Insect Surv. Vol. 8.

Univ. Calif. Press, Berkeley & Los Angeles. 269 pp.

Cottam, G. and J. T. Curtis. 1956. The use of distance measures in phytosociological

sampling. Ecol. 37: 451-460.

Denny, M. R. and S. C. Ratner. 1970. Comparative psychology. Research in animal

behavior. The Dorsey Press, Homewood, Illinois. 869 pp. »

Doane,R.W. 1901. Descriptions of new Tipulidae. J. Entomol. Soc. 9: 97-127.

Hanson, W. R. 1967. Estimating the density of an animal population. J. Res. on

Lepidoptera. 6(3): 203-247.

Roth, L. M. and S. Cohen. 1973. Aggregation in Blattaria. Ann. Entomol. Soc. Amer.

66: 1315-1323.

SCIENTIFIC NOTE

Aphilanthops hispidus as a Predator on Bees (Hymenoptera: Sphecidae).—The genus

Aphilanthops includes only four species, two of them known to prey upon queen bees of

the Formica fusca group (Evans, 1962, Behaviour, 19: 239-260). It has been assumed that

specificity for queen Formica ants distinguished Aphilanthops ethologically from the

related genera Clypeadon (prey: worker Pogonomyrmex ants) and Philanthus (prey: bees and

wasps). However, Aphilanthops hispidus Fox, a deserticolus species of southwestern U.S.

and northwestern Mexico, is a predator on bees. | located a nest of this species on 12

June 1975, 16 km W of LaPaz, Baja California Sur. The female was seen bringing bees into

a burrow in coarse, flat sand in an arroyo, and the nest was excavated. | failed to find any

cells, but | did find 7 paralyzed bees stored in the burrow 30 cm from the entrance, 12 cm

below the surface. They belonged to 4 different families: Colletes daleae Cockerell (80d)

(Colletidae), Agapostemon melliventris Cresson (1d)), A. mexicanus Robertson (1d)

(Halictidae), Ashmeadiella meliloti Cockerell (12) (Megachilidae), and Epeolus sp. (19)

(Anthophoridae). | am indebted to Dr. R. M. Bohart for identifying the wasp and to Dr.

G. C. Eickwort for identifying the bees. — HOWARD E. EVANS, Department of Zoology and

Entomology, Colorado State University, Fort Collins, CO 80523.

The Pan-Pacific Entomologist 53:123. April 1977.

New Neotropical Tillomorphini in the genus Tetranodus Linell

(Coleoptera: Cerambycidae)

John A. Chemsak

University of California, Berkeley

The members of the genus Jetranodus are unique among the New

World Tillomorphini by having the males with antennal segments

three to six incrassate. Additionally, the small size (8-6 mm in length)

and transverse ivory fasciae which extend completely in a single line

across the elytra make the species distinctive.

Since the genus was reviewed by Chemsak in 1969, additional

material representing two new species has become available. Of

added significance is the extension of the range of the genus to

South America. Previously the group was known only from Texas to

Honduras.

The authorities of the American Museum of Natural History, New

York, Field Museum of Natural History, Chicago, and the University

of Michigan, Ann Arbor, are gratefully acknowledged for the loan of

specimens. This study was supported by the National Science Foun-

dation through Grant GB-BM574.

Key to the species of Jetranodus

1. Elytra gibbose at base or with basal punctures coarse, dense, without broad

longitudinal spaces between rows of punctures .......... 00. c eee eee eee 2

Elytra with basal punctures fine, sparse, arranged in rows separated by broad

longitudinal glabrous spaces; color shining black, antennae paler. Length,

4-5 mm. Chiapas, Mexico to Guatemala ...............0000000- reticeps (Bates)

2(1). Elytra gibbose at base, punctures absent orindistinct .................0000 eee 3

Elytra not gibbose at base, basal punctures coarse, dense, without broad longi-

tudinal spaces between rows of punctureS........ 0... cc eee eee eee 4

3(2). Elytra with basal gibbosities acutely angulate at apices, median eburneous fasciae

oblique; color shining dark reddish brown. Length, 5mm. Panama.............

eh Be Oe RO SP A Bi WD Pag” oy ell fete, Je Be, Sens Calm geal tropipennis, n. sp.

Elytra with basal gibbosities rounded at apices, median eburneous fasciae trans-

verse; head, pronotum and at least part of antennae orange, elytra shining, dark

brownish. Length, 5-6 mm. Colombia ..............0000 eee xanthocollis, n. sp.

4(2). Elytra behind fasciae separately punctate, not scabrous .................0008s 5

Elytra behind fasciae deeply, closely punctate, scabrous appearing; color reddish,

elytra dark behind ivory fasciae. Length, 4.5 mm. Sinaloa, Mexico..............

ECAR Dic ti a dota RE ae Ae SE To RN 1 TS Pd en eee carne rugipennis Chemsak

5(4). Pronotum cylindrical, sides not angulate at middle; elytra with basal punctures

dense, not linearly arranged; males with segments three to six of antennae

strongly incrassate. Length, 3.5-5 mm. Texas to Oxaca, Mexico .. niveicollis Linell

Pronotum with sides somewhat expanded, angulate; elytra with basal punctures

linearly arranged, lines separated; males with segments three to six slightly

expanded. Length, 3-4 mm. Honduras ...............005. angulicollis Chemsak

Tetranodus tropipennis, new species

Male: Form small, cylindrical; integument shining, dark reddish brown. Head reticulate

punctate, micro-sculptured, sparsely clothed with long erect hairs; palpi with apices

The Pan-Pacific Entomologist 53:124-125. April 1977.

VOL. 53, NO. 2, APRIL 1977 125

strongly dilated; eyes large, pointed behind; antennae about as long as body, segments

three to six strongly incrassate, third segment longer than scape, fourth shorter than first,

outer segments finely pubescent. Pronotum longer than broad, cylindrical, base broadly

constricted; disk longitudinally striate; long erect hairs sparse; basal margin with a band

of appressed pubescence; prosternum deeply impressed, transversely rugulose, long

erect hairs sparse; mesosternum with a patch of pale appressed pubescence on epi-

meron; metasternum sparsely pubescent. Elytra about 2% times as long as broad,

broader behind middle; base with two elevated, acutely angulate gibbosities; median

eburneous fasciae slightly oblique; punctures on basal half very sparse, each bearing a

long erect seta, punctures behind middle sparse and setose, much finer and denser on

apical one-fourth with setae numerous; apices rounded. Legs sparsely pubescent, tibiae

carinate. Abdomen subglabrous, very sparsely punctate and pubescent. Length, 5mm.

Holotype male (University of Michigan) from Boquete, Chiriqui Province, Panama, 8

March, 1923 (F. M. Gaige).

The acutely angulate basal gibbosities of the elytra will separate this species from the

others.

Tetranodus xanthocollis, new species

Male: Form small, cylindrical; integument shining, brownish, head, pronotum and most

of antennae yellowish-orange. Head reticulate punctate, long, erect hairs sparse; palpi

with apices strongly dilated; eyes moderate sized, not acute behind; antennae shorter

than body, segments three to six moderately strongly incrassate, third segment longer

than scape, fourth shorter than first, basal segments with a few, long, erect hairs, outer

segments finely pubescent. Pronotum longer than broad, sides subangulate; disk shal-

lowly, longitudinally striate, sparsely asperate, each asperite bearing a long erect seta;

base constricted; basal margin moderately clothed with short appressed pubescence;

prosternum impressed, glabrous, with a few long, erect hairs; mesosternum with

epimeron densely white pubescent; metasternum with a patch of appressed pubescence

posteriorly at sides. Elytra about 2% times as long as broad, slightly broader behind

middle; base with two rounded elevated gibbosities; median eburneous fasciae trans-

verse; basal one-half with a few, seta-bearing punctures, apical one-fourth with numerous

small punctures, setae shorter than basal ones; apices rounded. Legs sparsely pube-

scent, tibiae carinate. Abodmen glabrous. Lengtyh, 5mm.

Female: Form similar. Antennae slender, only first two segments orange, extending

to apical one-fourth of elytra. Length, 5-6 mm.

Holotype male and allotype (Field Museum of Natural History) and two female para-

types from Pozo Colorado, 11 km SW Santa Marta, Magdalena Province, Colombia, 25-30

April, 1968, 1-15 May, 1968 (B. Malkin).

This species may be recognized by the orange head and pronotum, rounded basal

gibbosities of the elytra and the transverse eburneous fasciae.

Literature Cited

Chemsak, John A. 1969. Records and descriptions of Mexican and Central American

Tillomorphini. Pan-Pacific Entomol., 45: 303-317, 8 figs.

The Adults of Oroperla barbara (Needham)

(Plecoptera: Perlodidae)

C.A. Siegfried, P.J. Sheehan and A.W. Knight

Hydrobiology Laboratory, Department of Land, Air and Water Resources

University of California, Davis 95616

It has been over 40 years since the appearance of the description

of the nymph of this unique stonefly with lateral abdominal gills

(Needham, 1933). In that time only the adult male has been collected

and described (Jewett, 1966). Below is a description of the adult

female and the egg of Oroperla barbara. The drawings of the nymphs

were made by Penelope Kellar and Adele Hipps, and that of the egg

by Kathleen Jevons.

In March 1974 live Oroperla nymphs were collected from two N.

California locations on the Yuba River (T17N:R12E:S24). The nymphs

were maintained in a constant-temperature water bath at ambient

temperature ( 6°C) for the first two weeks after capture, after which

the temperature was increased about two degrees each week. The

nymphs were fed tubifex worms while in captivity.

As usual among the Pelcoptera, males preceded females in emer-

gence. The first male emerged 10 April and the first females at-

tempted to emerge 26 April although none emerged successfully

until 5 May. Emergence was complete by 15 May. We succeeded in

rearing 10 males and 4 females, while 2 additional females died dur-

ing emergence. No other deaths occurred during the rearing process.

Fig. 1. Lateral view of male terminal abdominal segments.

The Pan-Pacific Entomologist 53:126-128. April 1977.

VOL. 53, NO. 2, APRIL 1977 127

Fig. 2. Ventral view of female terminal abdominal segments.

The adults mated readily in the laboratory.

Our observations of the male coloration agree with the description

by Jewett (1966). Observations of 10 males reared in our laboratory,

however, did not reveal the sharp bend in the distal third of the

epiproct described in Jewett (1966). Instead, the eiprocts of all males

examined in our laboratory were smoothly curved anteriorly in the

distal third rather than sharply bent posteriorly (Fig. 1). The difference

between our specimens and that of Jewett (1966) in shape of the

epiprocts cannot be accounted for unless our specimens were some-

what teneral or Jewett’s was abnormal in the direction of the tip of

this structure (S. G. Jewett, personal communication). The latter

explanation probably is the case since our specimens were observed

mating and some were kept alive as long as two weeks after emerg-

ing, more than ample time to develop mature genitalia.

FEMALE — Length to wing tip, 27mm. Length of body, 220mm.

Length of antennae, 16mm. Length of cerci, 14mm. Head capsule

width, 5mm. Coloration and gill location as in male (Jewett, 1966).

Subgenital plate approaches tip of tergite of ninth abdominal seg-

ment (Fig. 2). Proximal 24 of genital plate triangular, narrowing to

about 1 its basal width. Distal % slightly longer than wide, with deep

U-shaped notch medially. Notch about 1/5 length of genital plate in

depth. Genital plate coloration pale, dull yellow, somewhat darker in

distal third.

128 THE PAN-PACIFIC ENTOMOLOGIST

Fig. 3. Lateral view of mature egg.

The dark honey-colored eggs were generally oval in shape, about

630u long and 430u wide, with a collar (Fig. 3). The surface was

covered with a series of oval indentations. One female mated and laid

209 eggs. The two mature female nymphs that died at emergence

contained 492 and 284 eggs.

Although all the above individuals were from the Yuba River,

nymphs of Oroperla barbara also have been collected from Miller

Creek (Jewett, 1966), a tributary of the American River, from the

American River itself, and from Indian Creek, Plumas County, Cali-

fornia (T25N:R11E:S8) by the authors.

Literature Cited

Jewett, S. G. 1966. Notes on Arcynopteryx (Oroperla) barbara Needham (Plecoptera).

Pan. Pac. Ento. 42:175-177.

Needham, J. G. 1933. A stonefly with paired lateral abdominal appendages. Jour.

Ento. and Zool. 25:17-19.

Revision of the Scarabaeidae:Anomalinae

3. A Key to the Species of Anomala of America North of Mexico

(Coleoptera)

Robert W. L. Potts

California Academy of Sciences, San Francisco, California

Because of a generally close relationship, combined with a remark-

able variability, Anoma/a species offer few morphological characters

so stable as to furnish easy differentiation between species. Many

of the key characters used in the past proved questionable when we

examined hundreds of specimens. A species that, in the main, has

only one broad elytral interval may have two or three in variant

specimens. Species with tridentate protibiae not uncommonly pro-

duce individuals with no sign of a third tooth. Or, as in the case of a

basal tooth on the tarsal claw joint, while the difference does

separate the species, the character is impossible to see in most

specimens without microscopic dissection. Variation between trans-

lucent thinly sclerotized and opaque heavily sclerotized specimens

of aspecies is more common than in most coleoptera, the two super-

ficially appearing as distinct species. Indeed, in one case of partially

opaqued sclerotization, the resultant false pattern was the principal

character cited for a proposed new species. Nonetheless, species

exist and are identifiable, but not often on single, simple characters.

Nor are the genitalia helpful. In the Anomalinae many species

share a generalized type whose only differences are in size and pro-

portion, and these are as variable as the external characters. Even in

those species where differentiation is possible there is a sufficient

variation for a drawing of a single specimen to be quite misleading,

and difficult to match.

Working with large numbers of specimens of such variable popu-

lations literally forced a number of compromises in working out the

key characters. However, it is believed that while the key may not be

altogether conventional, that that is not a drawback.

Distribution is sometimes the most positive distinction, as be-

tween species of the Southeastern, and the Southwestern States,

and it also may be positive in the case of relictual species confined to

a sharply limited habitat. Size is sometimes a close second as a dis-

tinguishing feature, but sometimes holds only for average speci-

mens, so it is normally used only in combination with one or more

additional characters. However, size is noted for all species, but more

as an easy check against error in the use of the key. In a number of

species the color pattern is most distinctive. Description of pattern is

not always simple, and this may appear confusing at first, but if com-

pared character against character with actual specimens, it is usually

clearly evident and positive. Finally, in one case, the time of year the

two species occur is the simplest and best differentiation.

The Pan-Pacific Entomologist 53:129-134. April 1977.

130 THE PAN-PACIFIC ENTOMOLOGIST

The key has been as carefully written and tested as possible, with

considerable effort being made to describe characters so they may

be checked negatively as well as positively, refusing to fit specimens

where they do not belong, but, in the aggregate, fitting well enough

where they do. However, a certain number of variants will refuse to

key at all except, perhaps, on an intuitive basis, or by association

with more normal specimens collected at the same time.

Key to the Species of Anomala of America North of Mexico

1. Larger protarsal claws simple or very obscurely cleft, with minor ramus extremely

fine, closely appressed; mesotarsal claws simple or sometimes finely cleft....2

Larger pro- and mesotarsal claws obviously cleft, minor ramus sometimes quite

small but always distinct; uncommonly the mesotarsal claws simple........ 11

2( 1). Clypeus subquadrate, sides parallel, subparallel or convergent basally, less than

tWIGB aS Wide as |ON Gs a. ese ek Seek e Wap ete te Mane toe scat ee lage te ve 3

Clypeus subsemicircular or trapezoidal, sides subparallel to divergent basally,

more than twice as Wide ASIONG...... keene eens 5

3( 2). Pale straw-colored to medium brown, unicolorous or pronotum with dark central

macula, elytra narrowly margined darker brown .............00+- eee eee eens 4

Dark brown; subsutural interval rugose, with few punctures; 11.5 mm; Arizona,

DragooniMtnS rer sek sete se eee ee obi dcey cbinaicn ges diel butleri (Howden)

4( 3). Clypeal suture more or less carinate; subsutural interval irregularly uniseriately

punctate or impunctate, rarely weakly biseriately punctate; 8-13mm, moderately

slender; Colorado and Kansas to Texas andArizona......... cavifrons LeConte

Clypeal suture flat or depressed in d, but distinct angle between planes of frons

and clypeus may give cariniform appearance, carinate in 2; subsutural interval

confusedly multipunctate, rarely strongly biseriate; 13-21mm, heavy-bodied;

western Texas to southern Arizona. ........ 0.000 ce eee eee carinifrons Bates

5( 2). Entirely dark brown to black above, sometimes paler beneath; pronotum may

become gradually paler toward lateral margins, but never with distinct light

brown band lateral. Ag kceccteea cide te etre sis. arate ecole ‘oleh Bibs Seg) a. eae eae Mia eI a a 6

Pale straw to largely brown-black; pronotum with at least lateral margins light

Drowns IN @GiStiMOt BAM sce wie eee Vere. a aprebesrwesew u erevenwta win WHO wa ae ae le ae 7

6( 5). North Carolina to Florida; 6-8mm; entirely shining black or rarely brown-black;

elytral punctation coarse with striae moderately grooved ....... mendica Casey

New Mexico and Arizona; 9-14mm; brown-black; elytra often partly pruinose,

especially lateroposteriorly; elytral striae shallow, finely punctate; a narrow

band of close-set short setae along carinate edges of abdomen and across

propygidium, but this hidden by margins of elytra when properly postioned

WM, Sits Satie cus ang ye eel le onlin t,o, RU PO lel otal te ical Ie Url een! eb castaniceps Bates

7( 5). Pronotum with single central macula or pairof maculae...................00e 8

Pronotum immaculate; elytra often with moderately broad dark margins but vary-

ing to nearly immaculate; 8.5-11.5mm; lower Colorado River basin, California,

ATAZOM Aire cogs % Mee aesazanal cc eine erueioe Mae Saadeh a: x acetate eae ae imperialae Potts

8( 7). North Carolinato Florida, Alabama. ......... 0.0... ccc cee eee eee nee 9

Texas, middle Rio Grande basin, near Del Rio and Eagle Pass; about9mm.......

Ee cl eR el Be eT Ps a ate meer Sr veeeeaee. Aabla Potts

9( 8). Pronotum with single macula, sometimes only margins pale................. 10

Pronotum with pair of small central maculae; 7-8mm (cf. couplet 36).............

BTL RPMS 2 Sa Sis aees OES ghhc re sha chte area Manoa eeemarut.at cvths cai 7 Rea Ae cama: parvula Burmeister

VOL. 53, NO. 2, APRIL 1977 131

10( 9). Elytra straw to pale brown, rarely 3rd and 5th intervals streaked darker brown;

pronotal macula variable in size, but if complete to base, enclosing pale area

mediobasally; clypeus deeply concave; body pilose beneath; 5-7mm

eh Ueno A RUM Lae neti So Ny Mes aia ncom ail Gc Meelis BY Ptoras ooo et sn ee semilivida LeConte

Elytra with dark brown margins to entirely brown-black; pronotal macula incom-

plete or complete to basal margin but never enclosing pale area; clypeus broad-

ly flat, strongly but narrowly reflexed to margins; not more than sparsely or

inconspicuously pilose beneath; 5-7mm (cf. couplet 37)...... minuta Burmeister

11( 1). Mesosternum between mesocoxae concave or flat anteriorly, never rising pos-

teriorly to more than AlIOW UMDO.... Lee eee eens 12

Mesocoxae separated by a distinctly convex to prominent raised mesosternal

PFOCESS; SOMELIMES POMTECE on. 6ee sees: dee ale ne Se ote eee ee ace aN es Oe ee 13

12(11). Mesocoxae moderately separated, the mesosternal plate between about as broad

or broader than a tarsal segment; pronotum often entirely red-brown to black;

pygidium- sometimes short pilose... -. 5 «saw ccn e ns ee ede eee ese 16

Mesocoxae narrowly separated, mesosternum between depressed, often concave;

pronotum pale brown or with central maculation, very rarely entirely dark;

pygidium with only fewlong hairs... ......... 0... e eee eee 24

13(11). Elytra subcostate or strongly striate but if with more than 10 or 11 apparent striae,

then 2 or 3 formed by large, rather than by small punctures; usually smaller,

6-12mm 14

Elytra with 13 to 15 subequal finely punctate grooved striae; often larger, 11-16mm;

pronotum with large dark macula and flacate margins, rarely entirely dark; Great

Lakes to southern New England, south to Texas and Floridamarginata (Fabricius)

14(13). Medium brown, varying to uncommonly entirely dull black; labrum visible before

CIV DOCUS Tes, ae, siaurs: se «. re Mt tice vars ectete. Neve megan 7 a Werle 2 Uae ce a OTe eatery tila oer 15

Entirely black, often with greenish metallic luster; labrum not visible before

clypeus; elytra laterally with 4 distinctly prominent subequal costate intervals;

LOPLI ARS. fsa vicemebavere reer ance Fare ey ed eheonase Aeree ch WO we A avec ated tie dato ene robinsoni Potts

15(14). Labrum moderately emarginate, extending only slightly beyond nearly vertical

| - anterior face of clypeus; elytra commonly with 7th and 9th intervals only

moderately convex and with a distinct 8th interval between, often for more than

half the length of intervals; Great Lakes to New England, south to Kansas and

FOIA eli ste teteage sus ters Rit he aret ase ihe ake erase seth ok Se an ee ees Aye lucicola (Fabricius)

Labrum deeply emarginate, porrect well beyond strongly oblique anterior face

of clypeus; elytra commonly with two strongly swollen costate intervals later-

ally, but 8th interval between often obsolescent, or if distinct, then uncommonly

extending for more than a short distance; Great Lakes to New England, south

to Tennessee and Florida. ..... 0... ee eee eee oblivia Horn

16(12). Pygidium largely glabrous; elytra with rather finely punctate or impunctate

grooved striae; antennal club of d often notably longer than stem........... 17

Pygidium pilose; elytra with strial series formed by brown or black colored punc-

tures, these sometimes coalesced into short foveae, most or all striae plane

or very slightly grooved; club of d subequal tostem ..............0e eee eee 19

17(16). Brown, or bicolorous pale and dark brown to black, if rarely slightly metallic, with

luster confined to head and pronotuM........ cee ee eens 18

Black, pronotum with green or coppery, elytra with green or bluish-green metallic

luster, rarely elytra dull; 11-13mm; southern Texas ........... insitiva Robinson

18(17). Head and pronotum pale straw or head a little darker; ciypeus about twice as wide

as long; labrum projecting shelf-like beneath broadly rounded-under anterior

face of clypeus; 10-13mm; Arizona, Coconino Co............ adscita (Robinson)

Head and pronotum red-brown, dark brown, greenish- or brownish-black or black;

clypeus more than twice as wide as long; labrum appressed to anterior face of

clypeus, not porrect; 8-13mm; Great Lakes and New England south to eastern

New Mexico and Florida. ........ 0.2.0. c cee eee flavipennis Burmeister

132

19(16).

20(19).

21(20).

22(21).

23(21).

24(12).

25(24).

26(25).

27(26).

28(27).

29(28).

THE PAN-PACIFIC ENTOMOLOGIST

Protibiae with external tooth... 2.2.0... ee eee 20

Protibiae without external tooth; 6.5-7.5mm; Florida, Highlands Co..............

South central Texas to North Dakota, east to New England and Florida ........ 21

Western New Mexico and Arizona; front of head punctorugulose, almost always

with scattered larger punctures as well; elytra immaculate, with fine binotate

spots or rarely with a little fine streaking, strial punctures discrete or uncom-

monly somewhat foveate; 9-12MM........ 0.00 cee eee eee ellipsis Casey

Elytra with strial series 2 and 3, 4 and 5 geminate, formed in part by short foveae,

sutural, 6 and sometimes 7 similarly formed; binotate spots rarely developed . 22

Elytra with striae not clearly geminate, colored strial punctures almost always

discrete; binotate spots rarely wanting .......... 00. e cece eee eee 23

Lateral margins of elytra with only short dark brown or blackish band from humeral

angle, extending only briefly beyond umbone; 8.5-13mm; southern Texas ......

eRe LAR tend oN ets soe ete igutiece gests wie bsyapanetye a doce cate Oraminosa Bates

Lateral margins of elytra more or less broadly dark brown or blackish from

humerus almost to suture; 6-9mm; North Dakota to Indiana, south to Texas and

LZQUISTAING secre oS tien os eaverrat aon chee cites eae Ieee 6 eee otaTae ee ludoviciana Schaeffer

Elytra dark red-brown to blackish, concolorous with pronotum or nearly, but often

with pale basal area; colored strial punctures and binotate spots more or less

hidden by dark coloring; 8-10mm; New Jersey to Florida.......... umbra Casey

Elytral disc often lighter than pronotum but suture and outer margin brown to

black, sometimes very broadly so; colored strial punctures sometimes large

and dark, almost always with a strong central dark spot on each elytron; 8-12mm;

Atlantic States to Great Plains and central Texas........... binotata(Gyllenhal)

Protibiae with external tooth .... 0... . ccc eee eee eee eees 25

Protibiae lacking external tooth; extremely small, 3.9-4.5mm; Florida, Lake

AlitapO Pp haar charset cus gesct fete tetieactes wih ncendey ce nlagalekseet woes gieeer ss so8t exigua (Schwarz)

Basal bead of pronotum formed by distinct groove along line proximal to basal

margin, entire or briefly interrupted medially.............. 0. ce ecb eee eee 26

Basal bead entirely wanting; labrum porrect before clypeus, deeply emarginate;

TIATOMIMPNOWSCISCY elect ce Bh, dese a seedog ew hen NS ie eek Patel os dubia (Scopoli)

Elytra with 5 or 6 slightly grooved striae before humeral umbone, intervals not

all nearly equal, with at least subsutural interval much broader and multipunc-

tate at leastin' parti. Bl ercyatihe 4 tetecs team lds wee sia cetee a, eso eters ae yale lok 27

Elytra with 6 well grooved striae before umbone, intervals subequal, costate, sub-

sutural never much broader, never multipunctate; pronotum immaculate to

black but often with paired maculae; elytra variable but often with dark fasciate

spots or bands; 7-11mm; New Jersey to Connecticut .... orientalis (Waterhouse)

Protibiae tridentate or with some angulate indication of 3rd tooth along outer

marginal line; elytra often broadly dark margined to almost entirely dark, less

commonly varying to immaculate; New Mexico to southern California....... 28

Protibiae bidentate, outer margin smoothly curved or straight before external

tooth; elytra usually narrowly margined, but sometimes spotted or entirely

black, Southwestern, Midwestern and Eastern species................0005 30

Pronotum with single central macula to entirely dark. ..............-2.-0000- 29

Pronotum commonly with pair of small to large maculae, sometimes immaculate,

rarely as a single macula but then often a pale median line partway through;

elytra broadly margined, both laterally and at suture, but varying to rarely nar-

rowly dark at suture only; 6.5-10mm, slender, not much broader behind middle;

lower Colorado River basin, California, Arizona................4. flavilla Bates

Elytra with only sutural margin narrowly dark brown; 9-12mm, slender to moderate-

ly robust, subparallel to somewhat cuneate; southern Arizona... digressa Casey

Elytra largely dark brown or brown-black with pale streakings on narrow intervals,

rarely with only humeral and umbonal areas dark; 9-13mm, broadest behind

middle, robust; southern Arizona and southwestern New Mexico

30(27).

31(30).

32(31).

33(32).

34(33).

35(30).

36(35).

37(36).

38(37).

39(35).

VOL. 53, NO. 2, APRIL 1977 133

Clypeus subrectangular, sides parallel or subparallel, less than twice as wide as

long; metatibiae shorter than or subequal to femora, no longer than first 4 tarsal

SEGQMMOMUSH 20 ice ota gdiee aes Seat ete ce he toaa Aree ete Ps Seleee le atotare eae eae ss foe eRe 31

Clypeus trapezoidal or subsemicircular, sides divergent basally, rarely subparallel,

more than twice as wide as long; metatibiae longer than femora, longer than

HINSt-AtarsaliSOGMOENtS ess series ecg oti Festa a eat Sra nede ear leig once eae ne Te an He fopel ate 35

Metatibiae not notably obconical, obviously more than twice as long as wide at

ADDON Ree ac Ss nite Macrae cureyint a ance OT ze eter ances Tea ce MEMO RG Anan caer hice cee vlon pertain a eabatee! Sabo 32

Metatibiae strongly obconical, not more than twice as long as wide at apex; 10mm;

ROK AS ttcas ty wes eeerntant: Chote Ras 42s, 4/0 Itc ERE Da courasta RUE Os ca Coe e ee ORNS sb zed tibialis Schaeffer

TO: DAS@ ras, 2c tear Fs Wis esis o alin | Bok aeaittve aretaoe Wevas a Tarek mapteeapanacare rss ne woe, webs 33

Maculation dark brown, strongly contrasting to ground color; single large pronotal

macula complete to base; elytra with dark margins strong and complete; ante-

rior pronotal angles obtuse but distinctly angulate; 7mm; western Texas

estes egsrt a diga tees eM OME aie epewcMake icc: haldtoneee se yaatietn a at etee Ma ae opti eat ward d suavis Potts

Minor protibial claw subequal to larger, divided claw in length; pronotum usually

with central macula or pairof macula@..... 0.0... . cc eee 34

Minor protibial claw approximately half as long as larger claw; pronotum immacu-

late; 5.0-8.5mm; California, Imperial Co., Glamis Sand Dunes..... carlsoni Hardy

Northwestern Arizona and southwestern Utah; pronotum with pair of well-sepa-

rated small anteromedial maculae, or immaculate; 6.5-9mMm......... kanei Potts

Southeastern New Mexico and western Texas; pronotum usually with single tri-

angulate anteromedial macula; 6-9mMM................6-. antennata Schaeffer

Atlantic Coast to Great Plains and south-central Texas; pronotum with dark central

macula, pair of maculae, or entirely brown or black; elytra broadly margined,

immaculate, or with fasciate spots or bands to entirely black............... 36

Western Texas to southern California; pronotum never entirely dark; elytra never

with fasciate spots or bands or entirely black

APP eer Peas odin pag re Soe ere 39

Pronotum never with paired maculae. ..... 2... ee eee eee 37

Pronotum with pair of central maculae, sometimes reduced or faint; 7-8mm; North

Carolinato Florida... 0... 0... cee eens parvula Burmeister

Elytra narrowly margined, usually with blackish fasciate spots or bands but varying

from immaculate to entirely black; Minnesota to southern New England, to

southcentral Texas and Florida... ......... 0c cece eee eee eens 38

Elytra medium brown with broad darker brown margins, particularly outside

humeral umbone, sometimes posterolaterally as well, more uncommonly broad

margined to almost extirely dark brown; 5-7mm; Georgia, Florida, and Alabama

(ciucoupletel O) a. «scene lasts ce tinge cecalte sg, teers een egg ee te minuta Burmeister

Generally a summer species, June and July; elytra immaculate to spotted, banded

or black, nearly always with dark area over apical umbone; head often bicolorous

pale brown and blackish; pronotal macula triangulate with waist-like constric-

tion near anterior margin, uncommonly varying to entirely black; scutellum pale

or narrowly dark margined, rarely entirely dark except in black form; 6-9mm.....

Be gy Fei eee ES TIT o gs tye Ee i ee ee eR Be ee Ree innuba (Fabricius)

Generally a spring species, March to May; elytral maculation similar but never

entirely black, and rarely with apical umbonal area dark; head often entirely

red-brown, gradually darkening to vertex but not distinctly bicolorous; pronotal

macula commonly complete to basal margin, but if not, then broadly rectangu-

lar, not especially constricted anteriorly nor pointed posteriorly; scutellum

nearly always dark or broadly dark margined, with pale median area; 7.5-10.5mm

Se MTU hee! eeaseartn satedeat sap tmeg a its Ste le eyatites! Sequel Mtoe, ameiteiae t undulata Melsheimer

Pronotum with central maculation .......0.. 0.0... cc cece eee eee 40

Pronotum IMMaculateseecttee te cet es eric le ais Wl ete ete ove wept woe ages avec 45

134

40(39).

41(40).

42(41).

43(42).

44(43).

45(39).

THE PAN-PACIFIC ENTOMOLOGIST

Pronotum with single central macula, or rarely with macula somewhat divided

medially, the division narrow or not complete to anterior margin, or maculae

Kote | anaes RRC Anchok eed ree eR nS bem chat alert nt Sia, Avatnnc ie’ fy Pakepman yh petite erat 41

Pronotum with well-separated pair of usually quadrate strong maculae; elytra with

narrow to broad margins, uncommonly only the suture narrowly margined (cf.

COUDIEHZS)) sey ta cer eka iba a yee tes eel etd etdnce a aan verter or flavilla Bates

Elytra pale to medium brown, very rarely more than narrowly to moderately mar-

gined, but if broadly marked, then only subsutural interval broad and multipunc-

ESS ee cP a ca lee PP Re ten enc ete aS eA OE Al ER EP ort Piva ee el A 42

Elytra dark brown or brown-black, often with pale streaking on narrow intervals;

two or three intervals broad and multipunctate (cf. couplet 29) .. . nimbosa Casey

Arizona, Santa Catalina Mtns., Sabino Canyon; 13-17mm; pronotum with central

macula, often extended posteriorly as a short line or point but sometimes com-

plete to basal margin, then enclosing pale area medially; elytral suture narrowly

dark margined and sometimes a dark macula over humeral umbone; metatarsi

remarkably stout, twice as wide as othertarsi .................. sabinae Potts

NOTPaS:GESCHIDOO et cadet, a Serta te ten cay reete et tan ae, Ist aes tae ues vast tte apecetne th 43

Elytra without distinct, or with narrow dark lateral margins, sometimes a dark

macula over humeral umMbone........... 2... cee ee ees 44

Elytra laterally with short blackish band below humeral umbone, and often again

dark around apical angle to suture, or with dark lateral margins complete but

fading or narrowing at middle; central pronotal macula often complete to base,

enclosing a pale mediobasal area and sometimes a pale anteromedial area

also; 8-11mm; central and southern Arizona. ................0 000s arida Casey

Pronotum with red-brown central macula complete to base, only lateral margins

pale, but sometimes tonal difference not great; elytra with dark sutural margin

extending forward to enclose scutellum to base; sutural interval flattened, al-

most as broad apically as basally; 9-12mm; southern Arizona (cf. couplet 29)... .

Boe sas eS bacysua SWebkivan ty shee 2 Rekha Whe tata a, atl cig BANS ee, cote Mee Ge digressa Casey

Pronotum with 5-sided dark brown macula, often slightly incised laterally, rarely

enlarged and complete to base; elytra with dark sutural margin ending at apex

of scutellum or extending only a little around it; sutural interval costate, widest

at middle, markedly tapering to apex; metatarsi and maculation quite similar

to sabinae, except consistently smaller, 8.5-12.5mm; Arizona, Baboquivari

Mtns. to Texas, Davis Mtns... ......... 0.000. delicata Casey

California, Imperial Co., Glamis Sand Dunes; elytra with sutural interval only

moderately costate anteriorly, flattened posteriorly, dark margin as a thin line

at extreme inner edge of suture; head basally of same pale straw hue as pro-

notum, gradually becoming light red-brown anteriorly; 7-10mm. hardyorum Potts

California, Riverside Co., Coachella Valley; elytra with sutural interval distinctly

costate, more so posteriorly than anteriorly, dark marginal band occupying at

least half of width; head almost unicolorous red-brown, contrasting to straw

hue of pronotum; 6.5-10mm (cf. couplet 28) ............ flavilla coachellae Potts

Descriptions of the First Instar Larvae of Three Species of

Epicautine Blister Beetles

(Coleoptera: Meloidae)'

John D. Pinto

Department of Entomology, University of California, Riverside 92521

The first instar larvae of Pleuropompha costata (LeConte), Epicauta

insignis Horn, and E. corvina (LeConte) are described below. Descrip-

tions of the two Epicauta at this time will facilitate their comparison

with members of the Epicauta maculata Group in a forthcoming mono-

graph focusing on the latter taxon. Of the two species included in

Pleuropompha, only the larva of P. tricostata Werner has been

described (MacSwain, 1956). In addition to the description of the larva

of the second species, P. costata, a key to species and a discussion

of the traits separating Pleuropompha from Epicauta are included.

To facilitate species comparisons, descriptions closely follow the

terminology and format employed by MacSwain (1956) in his exten-

sive study of the first instar larvae of the Meloidae. All quantitative

data represent means based on five (slide mounted) specimens that

emerged from the same egg mass. Where variation was substantial,

the range of measurements is given instead of the mean. Roman

numerals refer to segment number of the structure specified unless

otherwise stated. Exemplars of the species described here will be

deposited in the California Academy of Sciences.

Pleuropompha costata (LeConte)

(Fig. 1)

Color. Head, thorax and abdominal segments I-V yellow brown, abdominal segments

VI-IX dark brown. Head 0.92 as long as wide, as long as or only slightly shorter than pro-

and mesothorax combined; lateral margins gradually narrowing behind middle to dis-

tinctly emarginate; gula 2 as long as greatest head width, gular setae 2-24 as long as

greatest gular width. Antennae. II twice as long as Ill, two long and one short seta on apex;

sensory organ slightly shorter and wider than III; terminal seta short, only % the length of

Il. Mandibles very slender, with 20 very small, poorly delineated teeth; teeth slightly convex

apically; apical mandibular seta slightly longer than basal seta. Maxillary palpi. \ll twice

as long as wide, lateral margins curved, widest medially, narrowest apically; sensory area

of Ill extending 3/5 the length of segment; papillae of sensory area short, sparse, ca. 35

in number; two-segmented sensory appendix short, its length slightly less than

maximum width of Il of labial palpi. Lab/a/ palpi. | Y2 as long as Il; II slightly over twice as

long as greatest width; only a single seta on Il, this seta barely attaining apex of segment.

Thorax. Prothorax subequal in length to meso- and metathorax combined; line of

'This study was supported by grants GB-30907 and BMS75-17779 from the National Science Foundation.

The Pan-Pacific Entomologist 53:135-141. April 1977.

136 THE PAN-PACIFIC ENTOMOLOGIST

dehiscence absent from metanotum. Abdomen. Ten setae in posterior marginal rows on

terga }-VIII; spinelike evaginations at base of marginal setae on |-VII short, very poorly

sclerotized (Fig. 1), most strongly developed on II-V; posterior marginal setae on V as long

as or only slightly shorter than length of segment; setae of median transverse tergal rows

relatively long, those on V ¥3 as long as longest marginal seta; pleurites ventral, pleurite

V as long as wide, spiracle located in lateral half; abdominal spiracle | 4/5 the diameter

of mesothoracic spiracle; abdominal spiracles II-VIIl equal in diameter, all slightly smaller

than I; spiracle VIII smaller than Vil; sternum of segments I-VII weakly sclerotized, VII

with two medial sclerotized areas larger than those on preceeding segments, each

includiag two setae, areas occasionally joined; segments VIII and IX well sclerotized.

Legs with distance from articulation to apex of first coxa less than twice as long as

greatest coxal width; anterior femur with seven lanceolate setae; anterior claw with

longest seta reaching a point 9/10 the distance from base to apex of claw; claw long,

length % greater than maximum gular width. Body /ength 2.7 mm, caudal setae 0.8 mm.

Remarks. Pleuropompha, a genus quesiionably distinct from Epi-

cauta, contains only two species, both occurring in the southwestern

United States and northern Mexico. The larva of P. tricostata Werner

was described by MacSwain (1956). MacSwain believed that P/euro-

pompha could be distinguished from Epicauta by the rounded rather

than spinelike evaginations at the base of the posterior marginal rows

of setae on the abdominal terga. Although difficult to observe under

the light microscope, the scanning electron microscope reveals

typical spinelike evaginations on segments I-VIl in both P. costata and

P. tricostata (Figs. 1-3). At present then, we are left without a single

larval characteristic to separate these two genera. Only in the adult

stage are they easily distinguishable (Werner, 1943; Pinto, 1973).

Assuming that the spinelike evaginations are observed, both

species of Pleuropompha will run to couplet 16 in MacSwain’s key to

Epicauta. The following combination of characters should distinguish

them from all known Epijcauta: .

Head capsule emarginate; mandibles extremely slender, with 20

very small teeth; femur with seven lanceolate setae; abdominal seg-

ments not uniformly colored, VI-IX darker than II and IV; abdominal

sterna I-VIl incompletely sclerotized; abdominal terga with a posterior

marginal row of ten setae, terga I-VII with spinelike evaginations at

base of posterior marginal setae relatively short and very poorly

sclerotized.

The two species of Pleuropompha can be separated by the following

key.

1a Terminal seta of antenna distinctly longer than antennal segment

Il; Il of the labial palpi with two or three long setae; claws relative-

ly short, length of claw on anterior leg slightly less than great-

est gular width; setae of median transverse row of abdominal

terga short, those on V only 1/5-1/6 as long as longest marginal

seta; metathorax and abdominal segments | and II darker in color

APTANA IEA acess ect, afar xa get el t-te OOD Ok ran iak Pig! an ce, ol peu P. tricostata

VOL. 53, NO. 2, APRIL 1977 137

1b Terminal seta of antenna only ca. 3/4 the length of antennal seg-

ment II; Il of labial palpi with only a single long seta; claws rela-

tively long, length of claw on anterior leg ca. 1/4 greater than

maximum gular width; setae of median transverse row of abdom-

inal terga long, those on V ca. 1/3 as long as longest marginal

seta; metathorax and abdominal segments | and II similar in

GU GS Pe Cial WEN rol sactcscus) « alee WAGE A psieene Gea e arn. wlan arent om P. costata

Material studied. Larvae from a mass of 110 eggs; adults collected

7 mi. SE. Deming, New Mexico, VIII-2-1976. Larvae from a mass of

undetermined size; adults collected 6.8 mi. S. Apache, Cochise

County, Arizona, VIII-5-1972, J.D. Pinto. Quantitative data are based

on New Mexico material.

Epicauta insignis Horn

(Fig. 4)

Color. Yellow brown, head slightly darker. Head 0.98 as long as wide, 1/4 shorter than

pro- and mesothorax combined; laterial margins moderately to extremely emarginate;

gula 1/2 as long as greatest head width, gular setae slightly less than 1/2 as long as

greatest gular width. Antennae. II twice. the length of Ill, two long and one short seta on

apex, narrowed at base; sensory organ slightly wider and about as long as III; terminal

seta 1/2 longer than II. Mandibles moderately robust, with 11-15 apically flattened teeth,

9 teeth visible in outline; apical mandibular seta slightly longer and stouter than basal

seta. Maxillary palpi. \\| 3/4 longer than broad, evenly convex on outer margin, not swollen

basally; sensory area of Ill extending slightly over 1/2 length of segment; papillae of sen-

sory area short, sparse, ca. 40 in number; two-segmented sensory appendix relatively

long, its length 4/5 maximum width of II of labial palpi. Labja/ palpi. | 1/2 as long as Il; Il

twice as long as greatest width; single seta on II barely attaining apex of segment. Thorax.

Prothorax slightly longer than meso- and metathorax combined; line of dehiscence

extending full length of pro and mesonotum, indicated at apex of metanotum or not.

Abdomen. Ten setae in posterior marginal row on terga I-VIII; spinelike evaginations (Fig.

4) well developed at base of marginal setae on I-VIl; evaginations absent at base of

median transverse row; posterior marginal setae on V slightly less than 1/2 as long as

segment; setae of median transverse row slightly over 1/2 as long as longest marginal

seta; pleurites ventral, pleurite V wider than long, spiracle located in lateral half;

abdominal spiracle | 2/3 diameter of mesothoracic spiracle, slightly longer than ab-

dominal spiracle Il, remaining spiracles subequal to Il; sternum of segments I-VII weakly

ceeding segments, each with two setae; segments VIII and IX well sclerotized. Legs.

Seven lanceolate setae on anterior femur; anterior claw with longest seta almost reaching

apical 9/10 of claw; claw 3/20 longer than greatest gular width. Body /ength 1.6mm., caudal

setae 0.3mm.

Remarks. The larva of E. insignis is most similar to that of E. nigri-

tarsis (LeConte), a species placed in Group G by MacSwain (1956)

along with E. maculata (Say) and E. pardalis LeConte. A close relation-

ship between E. insignis and E. nigritarsis is also suggested by adult

anatomy (Werner, 1945; Werner et a/., 1966). The larvae of both

species run to couplet 14 in MacSwain’s key to Epicauta. Differences

between the two are as follows.

138 THE PAN-PACIFIC ENTOMOLOGIST

VOL. 53, NO. 2, APRIL 1977 139

In E. nigritarsis the sensory papillae on Ill of the maxillary palpi

are relatively dense with ca. 60 in number (not 100 as indicated by

MacSwain); in E. insignis only ca. 40 occur. Also, the two-segmented

sensory appendix on Ill of the maxillary palpi is long in E. insignis

with a length ca. 4/5 the maximum width of II of the labial palpi; in E.

nigritarsis its length is only 1/2 the width of this segment. The two

species can also be separated by the spinelike evaginations at the

base of the marginal setae on the abdominal terga. In E. nigritarsis

they are rather short (no more than 1/5 as long as their associated

setae) and only occur on segments I-V. In E. insignis they are longer

(up to 1/3 as long as their associated setae) and occur on I-VII.

In various species of Epicautina the cuticular reticulae comprising

the surface of the abdominal terga are produced into a short spine

apically. These are probably serially homologous to the evaginations

at the base of the marginal setae. In E. insignis the reticulae on the

apical half of abdominal segments I-VII in particular, are strongly

spinose (Fig. 4). These spines are easily visible under the light micro-

scope at 200X. In E. nigritarsis very few of the reticulae are spinose.

The spines in this species are also obsolescent and can only be

verified under the light microscope at magnifications approaching

500X.

Material studied. Larvae from amass of 257 eggs; adults from Pinery

Canyon, Chiricahua Mts., 5000 ft. elev.,.Cochise County, Arizona,

VII-25-1972, J.D. Pinto.

Epicauta corvina (LeConte)

(Figs. 5-7)

Color. Head golden brown; prothorax yellow with a broad, irregular, brown band across

middle; mesothorax and abdominal segments I-V yellow, metanotum and abdominal seg-

ments VI-IX dark brown. Head (Fig. 7). 0.94 as long as wide, slightly longer than pro- and

mesothorax combined; lateral margins broadly arcuate to subparallel, not emarginate;

gula slightly less than 1/2 greatest head width, gular setae 2/3 as long as maximum gular

width. Antennae. || twice the length of Ill, two long and one short seta at apex; sensory

organ slightly wider than III, subequal in length; terminal seta 2/3 longer than II. Mandibles

extremely robust, with 8-9 large, apically acute teeth; 4-5 teeth visible in outline; mandi-

bular setae well developed, apical seta considerably longer and stouter than basal seta.

Maxillary palpi. \ll 3/4 longer than broad, expanded at outer basal margin; sensory area ex-

tending 2/3 the length of segment; papillae of sensory area moderately long and dense;

length of two-segmented sensory appendix slightly more than 1/2 maximum width of II

of labial palpi. Lab/a/ palpi. | 1/3 as long as Il; Il twice as long as broad, with a single seta

barely attaining apex of segment. Thorax. Prothorax 1/10 longer than meso- and meta-

thorax combined, almost twice as broad as long; line of dehiscence extending full length

of pro-and mesonotum, absent from metanotum; meso- and metanotum with short, robust

—__—

Figs. 1-5. Scanning electron micrographs of sclerous evaginations at base of

posterior marginal setae on abdominal terga of epicautine first instar larvae. Micrographs

are of setae on abdominal segment V in E. insignis, and on IV for all other species. 1.

Pleuropompha costata, 6000X; 2. P. tricostata, 4000X (short variant); 3. P. tricostata, 4000X

(long variant); 4. Epicauta insignis, 1400X; 5. E. corvina, 2000X. Fig. 6. Scanning electron

micrograph of abdomen of E. corvina showing sclerous evaginations on both theposterior

and medial rows of tergal setae (140X).

140 THE PAN-PACIFIC ENTOMOLOGIST

0.2mm

Fig. 7. Head (ventral view) of first instar larva of Epicauta corvina.

integumental evaginations at base of setae along posterior margin; metanotum also with

evaginations at base of two central setae of transverse medial row. Abdomen. Ten setae

in posterior marginal row on terga I-VIII; integumental evaginations (Fig. 5) robust,

apically truncate, present at base of marginal setae on I-VIl; evaginations also well

developed at base of median transverse row of setae on I-IV or V (Fig. 6); posterior mar-

ginal setae on V subequal in length to that of segment; pleurites ventral, pleurite V wider

than long, spiracle located in lateral half; abdominal spiracle | 7/10 the diameter of meso-

thoracic spiracle, slightly larger than abdominal spiracle II, remaining spiracles subequal

to Il; sternum of segments I-VIl weakly sclerotized; VII slightly better sclerotized than pre-

ceeding segments, with two medial sclerotized areas each including three (rarely two)

setae; segments VIII and IX well sclerotized. Legs. Distance from articulation to apex of

first coxa 2 1/2 times as great as greatest coxal width; anterior femur with eight lanceolate

setae; anterior claw with longest seta reaching a point 9/10 distance from base to apex

of claw; claw length less thanwidth of gula. Body /ength 2.7 mm, caudal setae 0.7 mm.

Remarks. E. corvina is similar to E. pensylvanica (Degeer) and E.

funebris Horn as judged by adult characteristics (Werner, 1945; Wer-

VOL. 53, NO. 2, APRIL 1977 141

ner, et al., 1966). These two species as well as £. cinerea (Forster)

and £. pestifera Werner (as E. solani Werner) were placed in larval

Group B by MacSwain (1956). The larva of £. covina is not easily

placed in any of MacSwain’s groups but appears to most resemble B

and C. Like species in these groups, E. corvina has cuticular evagina-

tions at the base of the transverse median row of setae on theanterior

abdominal terga. Also, like E. funebris the head capsule is not emar-

ginate. A character shared with members of Group C is the more nor-

mal (= seven) number of lanceolate setae on each femur. The mandi-

bles, however, are even more robust than those found in members of

groups B and C, and are most similar to those of E. oblita (LeConte)

and E. callosa LeConte (Group J).

E. corvina is easily distinguished from all described North Ameri-

can Epicauta by the presence of the extremely robust, truncate eva-

ginations associated with the tergal setae (Figs. 5,6). In all other

Epicauta these evaginations are spinelike (e.g. Figs. 1-4). FE. corvina

runs to couplet 5 in MacSwain’s key to Ep/cauta. The distribution and

shape of the integumental evaginations, and the presence of eight

lanceolate setae on the femora will easily separate it from all species

keyed beyond that point.

Material studied. Larvae from a mass of 264 eggs; adults from 1.7

mi. E. Apache,Cochise County, Arizona, IX-5-1970, J.D. Pinto.

Acknowledgments

lam obliged to Dr. J.A. Chemsak (University of California, Berkeley)

for making Dr. J.W. MacSwain’s collection of larvalMeloidae available

for study. Figure 7 was prepared by Mr. Carl T. Conley. Technical

assistance on the scanning electron microscope was provided by

Mr. Jack L. Imbriani. Portions of this study were carried out at the

Southwestern Research Station of the American Museum of Natural

History, Portal, Arizona (Mr. V.D. Roth, Director).

Literature Cited

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

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

Pinto, J.D. 1973. Sexual behavior in the genus Pleuropompha LeConte: A new mating dis-

play in blister beetles (Coleoptera: Meloidae). Can. Entomol., 105: 957-969.

Werner, F.G. 1943. Revision of the genus Pleuropompha LeConte (Coleop., Meloidae).

Psyche, 50: 30-33.

Werner, F.G. 1945. A revision of the genus Epicauta in America north of Mexico (Coleop-

tera: Meloidae). Bull. Mus. Comp. Zool., 95: 421-517.

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

Univ. Arizona Tech. Bull. 175.

A new species of the genus Cryptopleurum Muls.

from Central America

(Coleoptera, Hydrophilidae)

Ales Smetana

Biosystematics Research Institute, Agriculture Canada, Ottawa, Canada

Cryptopleurum Mulsant is a rather small genus of the terrestrial

hydrophilid beetles belonging to the subfamily Sphaeridiinae, tribe

Megasternini. It is almost worldwide distributed (it seems to be miss-

ing in the Australian region) and includes about 20 species at present.

Only 1 species, C. impressum Sharp, has been known from the New

World, south of the United States border (Mexico).

During my work on the revision of the subfamily Sphaeridiinae of

America north of Mexico, | had the opportunbity to study a short

series of anew species of this genus from the Panama Canal Zone.

This is the first strictly neotropical species of Cryptop/eurum.

In the following, | present the description of this distinctive new

species and a key to the 2 New World species of Cryptopleurum

known to occur south of the United States border.

Cryptopleurum tenue, new species

Rather narrowly oval, convex, shiny; dark rufobrunneous with head darker (except

clypeus), elytra gradually becoming paler apically; an indistinct vague darkening of

lateral portions of elytra in about basal third present in most specimens; palpi and

antennae testaceous; legs darker testaceous with paler tarsi; undersides rufobrunneous

with more or less darker, sometimes almost piceous, sterna. Head with extremely fine

punctation, intervals between punctures on vertex distinctly larger than diameter of

punctures; surface without microsculpture; suture separating clypeus very fine, narrowly

interrupted in middle, punctation of clypeus slightly denser. Pronotum transverse, nar-

rowed in front, punctation sparser and slightly coarser than that on head, surface with-

out microsculpture. Each elytron with sutural stria and 9 striae; striae 1-5 not at all im-

pressed in basal half, rather indistinct, represented merely by serial punctures which are

only slightly coarser than punctures on intervals; striae gradually becoming slightly im-

pressed towards elytral apex; in general striae 6 and 7 weak and indistinct; 6th stria rudi-

mentary, very indistinct and formed only by few punctures, however, not fused with 7th

Stria; striae 8 and 9 developed as in other species of this genus, but again fine and incon-

spicuous; intervals with fine and rather sparse punctation, surface without microsculp-

ture; both strial and interval punctures bearing golden-yellow hairs which gradually be-

come more distinct towards elytral apex. No secondary sexual characters on 5th visible

abdominal sternite. Prosternum finely rugose, subopaque. Mesosternum coarsely and

rather densely punctate, no microsculpture between punctures. Elevated middle portion

of metasternum not impressed, finely and sparsely punctate; surface without microsculp-

ture. Aedoeagus and genital segment as in Figs. 4-6.

Length 1.25-1.30mm; width 0.75mm.

Holotype (male) and allotype (female): ‘Ft. Clayton C.Z. 1.-21.45” ‘Pres. by K. E. Frick

Collector.”” Deposited in the collection of the California Academy of Sciences, San Fran-

cisco.

Paratypes: 1d and 12, same data as the holotype. Deposited in the same collection as

the holotype, and in the Canadian National Collection, Ottawa.

Etymology: The specific name is the Latin adjective tenuis, -e (fine); it refers to the fine

The Pan-Pacific Entomologist 53:142-144. April 1977.

VOL. 53, NO. 2, APRIL 1977 143

=

=

ie

oO

6 = A

Figs. 1-3. Cryptopleurum impressum. Fig. 1. Tegmen with parameres. Fig. 2. Median lobe.

=ig. 3. Genital segment. Figs. 4-6. Cryptopleurum tenue. Fig. 4. Tegmen with parameres.

“ig. 5. Median lobe. Fig. 6. Genital segment.

144 THE PAN-PACIFIC ENTOMOLOGIST

elytral sculpture and to the relatively narrow and small body shape.

Distribution: The species is at present known only from the Panama Canal Zone. It is

probably more widely distributed in Central America.

Bionomics: Nothing is known about the habits of this species.

The species belongs to the impressum group (Smetana, in press);

however, it seems to be rather isolated there due to the fact that the

6th and 7th elytral striae are only vaguely developed and inconspi-

cuous. The species can be distinguished very easily from all New

World species of this genus by the very fine and superficial elytral

sculpture, and by the indistinct elytral striae 6 and 7 in particular.

It cannot be confused with any other New World species.

The 2 New World species of Cryptop/eurum, occurring south of the

United States border, can be distinguished as follows:

1 (2) Elytral striae 6 and 7 distinct. Punctation of elevated middle

portion of metasternum moderately coarse and dense.

Aedoeagus and genital segment as in Figs. 1-3. Length 1.50-

1.60mm. Mexico and Southern Arizona........... C. impressum

2 (1) Elytral striae 6 and 7 indistinct, 6th stria rudimentary, formed_

only by few punctures. Punctation of elevated middle portion of ©

metasternum fine and sparse. Aedoeagus and genital segment

as in Figs. 4-6. Length 1.25-1.30mm. Panama Canal Zone.......

Literature Cited

Smetana, A. Revision of the subfamily Sphaeridiinae of America north of Mexico

(Coleoptera: Hydrophilidae). Mem. Ent. Soc. Canada, in press.

SCIENTIFIC NOTE

Prey Specificity in Clypeadon (Hymenoptera: Sphecidae).—In 1962 (Behaviour, 19:

239-260) | reported that each species of C/ypeadon appears to prey upon only one species

of harvester ant, sometimes rejecting other species in the close vicinity. The following

new records may be of interest. In Larimer Co., Colorado, C. /aticinctus (Cresson) appears

to prey only on worker Pogonomyrmex occidentalis Cresson (as it does elsewhere), hardly

a surprising fact since these are the only members of their respective genera occurring

in north central Colorado. At Hasty, Bent Co., in southeastern Colorado, | have taken

C. dreisbachi Bohart capturing worker P. rugosus Emery at their nest entrances, thus con-

firming a similar record from Zacatecas, Mexico. At Tornillo, El Paso Co., Texas, in May

1974, | took a female C. utahensis (Baker) taking workers at a nest entrance of P.

californicus (Buckley). This wasp had previously been reported preying upon P. barbatus

(Smith) in California. The Texas record is from the eastern extremity of the range of both

wasp and prey, and it is possible that in this area C. utahensis replaces the usual predators

on P. californicus (e.g. Listropygia bechteli Bohart). | am indebted to Dr. R. M. Bohart for

identifying the Clypeadon and to Dr. A. C. Cole for identifying the Pogonomyrmex. —

HOWARD E. EVANS, Department of Zoology and Entomology, Colorado State University, Fort

Collins, CO 80523.

The Pan-Pacific Entomologist 53:144. April 1977.

Elevation of Loxaulus brunneus variety atrior (Kinsey)

to full species status

(Hymenoptera: Cynipidae)

D. Charles Dailey

Sierra College, 5000 Rocklin Road, Rocklin, CA 95677

Loxaulus brunneus variety atrior (Kinsey, 1922) was originally des-

cribed in the genus Compsodryoxenus Ashmead as a host restricted

form on Quercus (Lepidobalanus) lobata Nee, from Byron, Contra Costa

County, California. Weld did not believe Loxaul/us brunneus (Ashmead,

1896), a species on Quercus (Protobalanus) chrysolepis Leibmann,

would occur on Q. /obata; so he listed L. brunneus var. atrior as a

separate entity in the Catalog of the Hymenoptera of North America

North of Mexico (R. J. Lyon — personal communication). Of the 501

species names of cynipid gallmakers occuring on oaks listed in the

catalog and both supplements (Weld, 1951, 1958, 1967), this is the

only one listed as occurring on more than one subgenus of oaks. This

deviation from such a distinct biological pattern suggested the

separate identity of the two forms. The following indicates it is speci-

fically distinct from L. brunneus (Ashmead) on Q. chrysolepis.

Comparison of the primary types showed variety atrior can be dis-

tinguished from L. brunneus (Ashmead) by the ventrally diverging

inner -ocular margins; rounded occiput; head greater than 1.25 times

as wide as high; anteriorly incomplete sculptured notaulicies, which

are narrowly separated posteriorly; posteriorly curved rear margin of

mesoscutum; weakly ridged, round to oval foveae; conspicuously

branched medical propodeal carina; and ventral spine 1.5 to 2.0 times

as long as high. Loxau/us brunneus has parallel inner-ocular margins;

flattened occiput; head less than 1.25 times as wide as high; com-

plete non-sculptured notaulicies, separated posteriorly by almost

the width of one of the strongly ridged transverse foveae; slightly

anteriorly curved rear margin of mesoscutum between notaulicies;

numerous propodeal carinae; and ventral spine 2.0 to 2.5 times as

long as high.

Gall biology — The original records relating to the identity of the

gall of L. brunneus (Ashm.) are highly confused. No attempt to resolve

this problem is presented, as these species can be readily distin-

guished by using only morphological differences.

Systematics — The original identification label of Kinsey’s type

specimen reads, ‘‘Comps. atrior HOLOTYPE.” As with much of

Kinsey’s work, the imprecisely used term ‘‘variety’” could have been

listed as “host restricted ecological subspecies.”’

Type designation and deposition — The original description of

C. brunneus var. atrior mentions the designation of ‘‘Holotype fe-

males” (plural). There are no records of more than one Holotype

specimen other than in the original publication (M. Favereau —

The Pan-Pacific Entomologist 53:145-146. April 1977.

146 THE PAN-PACIFIC ENTOMOLOGIST

personal communication). Assuming the listed designation was

a typographical and proof reading error, the existing single specimen

in the American Museum of Natural History labeled Holotype is

accepted as the primary type, and the name atrior raised to full

species status.

Loxaulus atrior (Kinsey), new combination

Distribution — Recorded from Byron, Contra Costa Co., Kinsey;

Davis (12 specimens) and 5 mi. West of Madison, (6 specimens) Yolo

Co., D. C. Dailey, Q. /obata; Marysville, Yuba Co., D. C. Dailey, Q.

lobata, (2 specimens), Folsom Lake, Placer Co., D. C. Dailey, Q. /Jobata

(1 specimen); Pentz, Butte Co., Calif. H. H. Kieffer, Quercus douglasii

(new host record, if accurate) IV-16, 1928 (1 specimen). L. atrior prob-

ably occurs throughout the range of Quercus /obata Nee and possibly

Quercus douglassii Hooker and Arnott.

Acknowledgments

Loan of the Holotype of L. atrior by M. Favereau of the American

Museum of Natural History and L. brunneus by Dr. Paul D. Hurd, Ur.,

of the U.S. National Museum and use of Weld’s notes and personal

notes by Mr. Robert J. Lyon of the Los Angeles City College are

greatly appreciated.

Literature Cited

Ashmead, W. H. 1896. Descriptions of New Cynipidous galls and gall wasps in the

United States National Museum. Proc. U.S. Natl. Mus. 19:129.

Kinsey,A.G. 1922. New and described Cynipidae. Ind. Univ. Studies 53:39.

Weld, L. H. 1951. Cynipoidea, in Hymenoptera of North America North of Mexico,

Muesebeck, Krombein, and Townes. U.S.D.A., pp. 594-654. 1958. Ibid. Supp. |.

1967. Ibid. Supp. Il.

Editorial Notice

The editors are attempting to put the Pan-Pacific Entomologist

back on schedule, however we are having some problems. The

typeface we now use takes approximately 20% less space than the

former style. This, coupled with several other factors, means we are

in desperate need of manuscripts. We always have need for short

notes of less than a full printed page, including book reviews of

appropriate subjects.

A List and New Distributional Records of Pacific Coast Odonata

Dennis R. Paulson

Washington State Museum, University of Washington, Seattle, 93103

and

Rosser W. Garrison

Division of Entomology and Parasitology, University of California, Berkeley, 94720

In the course of considerable field work and examination of insect

collections in the Pacific states (Washington, Oregon and California),

we have accumulated records of anumber of species of Odonata that

extend their known ranges. Species lists with little detailed informa-

tion on distribution were published for California (Smith and

Pritchard, 1956) and Washington (Paulson, 1970). Virtually nothing

has been written about the Odonata of Oregon since the excellent

papers by Kennedy (1915, 1917), which remain the basic references

for these states. The following list includes only those records that

are the first fora state or that substantially extend the known limits of

distribution of a species. Some species were included on the

California list by Smith and Pritchard (1956) on the basis of specimens

in the University of California, Berkeley, and California Academy of

Sciences collections rather than published records (R. F. Smith, in /itt.

to Paulson, 22 November 1974), and we present herein distributional

data for these species. In addition, we question the presence of

several species in California.

We appreciate greatly the courtesies of the staff of the museums

visited: Paul Arnaud and Carolyn Mullinex at the California Academy

of Sciences (CAS); Jerry Powell and John Chemsak at the California

Insect Survey, University of California, Berkeley (CIS); and Charles

Hogue, Julian Donahue and Roy Snelling at the Los Angeles County

Museum of Natural History (LACM). Specimens in a few other collec-

-tions are cited, and the remainder are in our own collections (DRP,

RWG). We have had considerable assistance in the field from the

following persons: Mary Lynn Erckmann, Charles Turner, Elisabeth

Schaublin, Eileen O’Connor, Paulette Bierzychudek and Cindy Power

(DRP); and Jo Allyn Garrison (RWG). All of them made time in the field

more pleasant as well as augmenting our collections.

CALOPTERYGIDAE

Hetaerina americana (Fabricius). CALIFORNIA, Siskiyou Co., Klamath River, 4.4 mi. W

Cal. 263 on Cal. 96, 7 July 1974, 20 39; Shasta River, 4.3 mi. N Yreka, 16 August 1973, 4d

19; 7 July 1974, 4d 39 (all DRP). Previous northernmost record Butte County (Kennedy,

1917).

The Pan-Pacific Entomologist 53:147-160. April 1977.

148 THE PAN-PACIFIC ENTOMOLOGIST

LESTIDAE

Archilestes grandis (Rambur). CALIFORNIA, north to Ventura Co., Sespe Creek, 4.0

mi. N Fillmore, 4 October 1965, 16 (DRP); 24 October 1965, 12 (DRP); Riverside Co., Indio,

5 October 1918, 1d (CAS); and Tuolumne Co., near Groveland, 14 August 1954, 2d teneral

(CIS). Listed from California by Smith and Pritchard (1956) but no previous records other

than those before the confusion between A. grandis and A. californica was terminated

(Kennedy, 1915). The record from Tuolumne County, in the foothills of the Sierras, is con-

siderably northwest of a line drawn through the other northwesternmost records: Wash-

ington County, Utah (Brown, 1934) and SOUTH DAKOTA, Custer Co., stream 1 mi. W west

entrance to Wind Cave National Park on U. S. 385, 4200’, 26 August 1969, 106 (DRP).

Lestes congener Hagen. CALIFORNIA, San Diego Co., Cuyamaca Reservoir, 4600’, 21

August 1974, 11¢ (RWG); 24 August 1974, 2d (DRP). Previous southernmost record Santa

Clara County (Kennedy, 1917).

Lestes dryas Kirby. CALIFORNIA, San Bernardino Co., San Bernardino Mountains,

South Fork, 6200’, 30 June 1907, 1¢ (LACM); Barton Flats, 26 June 1936, 19; 1 July 1936, 19

(both CAS). Previous southernmost record Tuolumne County (Kennedy, 1917).

COENAGRIONIDAE

Amphiagrion abbreviatum (Selys). CALIFORNIA, San Diego Co., Laguna Mountains, 18

~ June 1938 (LACM). Previous southernmost record Fresno County (Kennedy, 1917). The

species has not been recorded from Mexico, but there are numerous specimens from Baja

California (DRP, CAS, CIS).

Anomalagrion hastatum (Say). CALIFORNIA, Imperial Co., canal 8 mi. E Holtville, 1

September 1967, 12; ditches near Imperial Dam, 23 August 1974, 4d 59 (both DRP). Known

from the United States as far west as Reeves County, Texas (Johnson, 1972) and in

western Mexico as far north as 15.6 mi. WNW Guasave turnoff on Mex. 15, SINALOA (DRP)

but not from Sonora or Baja California. Thus this is a considerable northwestward range

extension.

Argia agrioides Calvert. CALIFORNIA, many specimens and localities north to Siskiyou

Co., Shasta River, 4.3 mi. N Yreka, 16 August 1973, 2¢ (DRP). This species, described from

Baja California (Calvert, 1895), has long been confused with A. nahuana. Gloyd (1958)

adequately differentiated the two species but stated that agrioides was confined to Baja

California. Subsequently she has examined many California specimens (determinations

in LACM and CAS) and confirmed specimens taken by Garrison in Arizona (in litt. to

Garrison, 13 February 1977).

Argia emma Kennedy. OREGON, specimens from Douglas, Jackson, Josephine and

Lake Counties (DRP). Previously recorded from Washington and California (Kennedy,

1915, 1917).

Argia lugens (Hagen). OREGON, Josephine Co., Rough and Ready Creek, 5 mi. S Cave

Junction, 13 August 1971, 1¢ (DRP). Previous northernmost record Butte County,

California (Kennedy, 1917). ‘

Argia nahuana Calvert. CALIFORNIA, Siskiyou Co., Shasta River, 4.3 mi. N Yreka, 16

August 1973, 3d (DRP). Previous northernmost record difficult to determine, as Kennedy

(1917) confused this species and A. agrioides, but his northernmost record of the

combined species was from Butte County. Paulson has taken agrioides at five localities

and nahuana at 14 localities in California with much overlap in distribution and co-

existence at three sites.

Enallagma anna Williamson. OREGON, Lake Co., 1.0 mi. N Summer Lake, 15 August

1974, 1d; Paisley, 15 August 1974, 2d; CALIFORNIA, Lassen Co., 15.0 mi. N Susanville, 12

September 1967, 1d; Long Valley Creek, 7.3 mi. SE Doyle, 24 August 1973, 9d 79 (all DRP);

Plumas Co., Bucks Lake, 1 July 1949, 16 12 (CIS); Inyo Co., Big Pine, 13 August 1955, 1d;

4.6 mi. S Big Pine, 21 June 1973, 11¢ 59 (all DRP). Westernmost records for this species,

which was recorded from California by Smith and Pritchard (1956) with no further locality

records.

VOL. 53, NO. 2, APRIL 1977 149

Enallagma basidens Calvert. CALIFORNIA, Imperial Co., ditches near Imperial Dam, 23

August 1974, 5d 12 (DRP). Originally described from Texas (Calvert, 1902), this species

has undergone a dramatic range extension during the past 40 years (Montgomery, 1966),

extending its distribution widely into the eastern states. It has been recorded previously

as far west as Arizona (Johnson, 1972), and its spread toward the west has doubtless been

possible by the increase in irrigation.

Enallagma civile (Hagen). CALIFORNIA, many localities through the southern part of

the state north to Butte Co., Bidwell City Park by Big Chico Creek, Chico, 14 June 1975, 3d

(RWG); Solano Co., 5.0 mi. W Vacaville, 18 October 1975, 14d 82 (RWG); and SanFrancisco

Co., San Francisco, 27 September 1963, 2d (CAS). Listed by Smith and Pritchard (1956),

but we have been unable to find a specific locality record in the literature. As it is now

common in areas at which Kennedy (1917) collected but did not find it, it must be spread-

ing into and within the state. It is one of the most characteristic species of disturbed

habitats — irrigation ditches, reservoirs, cattle tanks — in the Southwest and should

continue to increase with human modification of the entire area.

Enallagma clausum Morse. OREGON, Lake Co., Lake Abert, 15 August 1974, 2d 29;

CALIFORNIA, several localities west to Siskiyou Co., Tule Lake National Wildlife Refuge,

8 July 1974, 2d; and south to Inyo Co., Little Lake, 20 June 1973, 2306 69 (all DRP). These

are westernmost records for this species, which was known from Yakima County,

Washington (Kennedy, 1915) and California (Smith and Pritchard, 1956) with no specific

locality records.

Ischnura barberi Currie. CALIFORNIA, numerous localities north to San Bernardino

Co., Saratoga Springs, 10-12 July 1953, 1¢6 (LACM); and west to Orange Co., Newport Bay,

5-7 September 1971, 9d 89 (DRP). Included in the key to California /schnura by Smith and

Pritchard (1956) but California not indicated in their account of its distribution.

Ischnura denticollis (Burmeister). OREGON, Lake Co., 1.0 mi. N Summer Lake, 15

August 1974, 4d (DRP). Previous northernmost record Butte County, California (Kennedy,

1917).

Ischnura perparva Hagen. CALIFORNIA, San Diego Co., San Diego, 10 August 1916, 19

(CAS). Previous southernmost record Tulare County (Kennedy, 1917).

Nehalennia irene (Hagen). CALIFORNIA, Lassen Co., Silver Lake, 26 July 1973, 69

(CAS). Closest known records Chelan County, Washington (Paulson, 1970) and Weston

County, Wyoming (Bick and Hornuff, 1972). We consider the record from Santa Clara

County, California (Seemann, 1927) very unlikely on ecological grounds.

Telebasis salva (Hagen). CALIFORNIA, Butte Co., Bidwell City Park by Big Chico Creek,

Chico, 14 June 1975, 10 (RWG). Previous northernmost record Yuba County (Kennedy,

1917). Amphiagrion abbreviatum was also taken at the same locality, the first recorded

instance of sympatry between these red species. T. sa/va occurs at low elevations in the

Coast Range and Sierra Nevada foothills, while A. abbreviatum normally occurs at higher

elevations in the Sierra Nevada and outlying southern California ranges. Chico, at 200’

elevation, is considerably lower than Kennedy’s (1917) record from Coulterville (1740’).

Zoniagrion exclamationis (Selys). CALIFORNIA, Humboldt Co., Orick, 4 June 1931, 19.

(CAS). Previous northernmost record Butte County (Kennedy, 1917).

PETALURIDAE

Tanypteryx hageni (Selys). CALIFORNIA, Napa Co., Samuel Spring, 13 May 1956, 1d

(Univ. California, Davis); and above Pope Creek, 1 mi. W Lake Berryessa (near Samuel

Spring), 30 May 1967, 36 (R. W. Cruden). These are the southernmost localities in the

Coast Range for this locally distributed species.

AESHNIDAE

Aeshna canadensis Walker. WASHINGTON, Mason Co., Long Marsh, 5.2 mi. SW and 3.7

150 THE PAN-PACIFIC ENTOMOLOGIST

mi. N Belfair, 24 August 1971, 1d; also several localities in Snohomish County (all DRP).

Known from only the Spokane area in the state previously (Paulson, 1970), the present

records the first from west of the Cascades in the Pacific states. Also recorded from

southwestern British Columbia (Walker, 1958).

Aeshna constricta Say. WASHINGTON, Clark Co., pond just E Woodland, 10 Bidet

1974, 12; also from Douglas, Klickitat and Yakima Counties (all DRP). Known previously

from only the Spokane area in this state (Paulson, 1970); the Woodland record is the first

from west of the Cascades. Unlike other northern Aeshna that occur in western Washing-

ton, this one is lacking from southwestern British Columbia (Walker, 1958), and the

Columbia River rather than the Fraser River has probably provided it a route through the

Cascades.

Aeshna eremita Scudder. WASHINGTON, Snohomish Co., near Monroe, 17 September

1974, 2d (DRP). Only previous record in Pacific states from Pend Oreille County (Paulson,

1970). As in A. canadensis, this species also occurs in southwestern British Columbia

(Walker, 1958) but was otherwise unknown west of the Cascades.

. Aeshna juncea (Linnaeus). OREGON, Clackamas Co., Still Creek Campground, 1 mi. SE

Government Camp, 3800’, 26 July 1973, 10 (RWG). Previous southernmost record King

County, Washington (Paulson, 1970). Calvert’s (1895) record from California probably

refers to some other species of this group, perhaps A. interrupta.

Aeshna palmata Hagen. CALIFORNIA, San Diego Co., Indian Canyon, 20 July 1948, 1d

CIS). Previous southernmost record Santa Clara County (Kennedy, 1917). This record is

surprising, as pa/mata is otherwise known from ponds at higher elevations, desert water-

courses east of the Sierras and streams in the northern part of the state, and there has

been considerable collecting in southern California with no additional records. Probably

all extralimital records of well-known species based on single specimens should be

treated with caution when assessing the distributional patterns of these species.

Aeshna tuberculifera Walker. WASHINGTON, Stevens Co., Hande Creek, 1.0 mi. S and

0.6 mi. W Middleport, 22 July 1973, 1d; Twin Lake, SW of Middleport, 24 July 1973, 19

(both DRP). Previously known in the Pacific states from a single locality in western

Washington (Paulson, 1970), this species was not expected east of the Cascades as it has

not been reported from between the Great Lakes and Vancouver Island (Walker, 1958). The

present records indicate the hiatus in its distribution may be more apparent than real.

Aeshna umbrosa Walker. CALIFORNIA, Sonoma Co., Annapolis, 23 September 1962, 1d

(CAS); and Tulare Co., Sequoia National Park, near Tharp’s Log, 1.5 mi. E Giant Forest,

6700’, 27 July 1972, 1d (RWG). Previous southernmost record Tehama County (Roback

and Westfall, 1967). The first locality furnishes the only known instance of sympatry

between this species and A. walkeri, although their ranges overlap to some extent.

Likewise, it is the only record of umbrosa away from the northeastern part of the state. See

comments under A. pa/mata.

Aeshna walkeri Kennedy. OREGON, Lake Co., Crooked Creek, 5.0 mi. S Valley Falls, 15

August 1974, 9d 12 (DRP). Previous northernmost record Santa Clara County, California

(Kennedy, 1917). The published records indicate a Coast-Range distribution, but in

addition the species is locally common along the west side of the Sierras, whence we

have specimens from Amador, Calaveras, El Dorado, Mariposa, Shasta and Tuolumne

Counties. Kennedy (1917:587) listed A. pa/mata from the ‘‘colder streams of the Sierras,”

but we have found only wal/keri on streams on the west slope. Both species were taken

together on 16 August 1973 at the Shasta River, 4.3 mi. N Yreka, Siskiyou County (DRP).

Anax walsinghami McLachlan. CALIFORNIA, Shasta Co., Cayton, 19 July 1918, 1d

(CAS). This is another surprising peripheral record, as the species is characteristic of the

Sonoran region, the previous northernmost record being from Orange County (Seemann,

1927). Other species in the CAS taken at Cayton include Amphiagrion abbreviatum, Argia

vivida*, Ischnura cervula*, Aeshna multicolor*, Ophiogomphus morrisoni, Cordulegaster

dorsalis, Lepthemis collocata*, Leucorrhinia intacta and Sympetrum corruptum*, a mixture of

mostly wide-ranging species. Those marked with an asterisk are known to occur with

walsinghami farther south. Some support for the record is furnished by the occurrence of

this species at an intervening locality, Del Puerto Canyon in Stanislaus County (RWG).

VOL. 53, NO. 2, APRIL 1977 151

GOMPHIDAE

Erpetogomphus compositus Hagen. WASHINGTON, Grant Co., Crab Creek, 7 mi. NW

Othello, 3-6 August 1972, 2d; Benton Co., Yakima River, 9 mi. N Benton City, 16-17 August

1971, 1d 12 (all DRP). Listed for Oregon with no locality by Calvert (1905), the northern-

most specific locality Butte County, California (Kennedy, 1917).

Gomphus olivaceus Selys. WASHINGTON, localities in Grant, Yakima and Benton

Counties, from 10 July to 17 August (all DRP). Recorded from British Columbia (Walker,

1958) and California (Kennedy, 1917), this species had not been taken definitely in

Washington at the time of compilation of the state list (Paulson, 1970). Kennedy (1917:

554) noted that exuviae reported by Needham (1904) from Seattle, King County, as G.

sobrinus ( = G. kurilis) are actually olivaceus, and Needham’s illustration matches olivaceus

better than kurilis, but we consider this record unlikely enough on ecological grounds to

suggest that it not be given credence until confirmed.

Gomphus plagiatus Selys. CALIFORNIA, Imperial Co., Winterhaven, 5 September 1965,

3d; 1 September 1967, 2d 19; canal 8 mi. E Holtville, 5 September 1965, 1d, 1 September

1967, 16 22 (all DRP). This species, recorded west to Texas by Needham and Westfall

(1955), occurs west through southern New Mexico and Arizona into the southeastern

corner of California and in northern Mexico at NUEVO LEON, Rio Las Lajas at Mex. 40, 2

mi. N China, 24 June 1965, 19 (DRP). California specimens are very much paler, both in

intensity of color and extent of markings, than those from Florida, and central Texas

specimens are intermediate in color between the two extremes. Material from California

has the anterior pale areas of the thorax broadly confluent with the pale collar and keys to

G. olivaceus in Needham and Westfall (1955). Western specimens also differ in size from

Florida ones. Hind-wing lengths in millimeters of mature males are as follows, with the

mean, range and number of specimens measured in order for each population sample:

Florida — 35.3, 35-36 (7); Texas — 32.8, 32-34 (4); California — 32.6, 31-34 (7).

Ophiogomphus bison Selys. OREGON, Douglas Co., 10 mi. S Roseburg, 22 June 1960,

12 (CAS). Previous northernmost record Butte County, California (Kennedy, 1917).

Ophiogomphus severus Hagen. OREGON, Lake Co., Crooked Creek, 5.0 mi. S Valley

Falls, 4500’, 15 August 1974, 12 (DRP); CALIFORNIA, Modoc Co., Warner Mountains, 6 mi.

E Fandango Pass, 20 July 1974, 12 (RWG). Previous southernmost record Umatilla

’ County, Oregon (Kennedy, 1915).

Progomphus borealis McLachlan. CALIFORNIA, Napa Co., Pope Creek by Pope Creek

Rd., 3.8 mi. W jct. Berryessa-Knoxville Rd., W of Lake Berryessa, 20 April 1975, 1 larva

(RWG); Inyo Co., just S of Lower Haiwee Reservoir, 18 July 1961, 16 (DRP); end of Five

Bridges Rd., 3 mi. N Bishop, 17 July 1973, 1¢ (RWG). These records are the farthest north

in the Coast Range and the first east of the Sierra Nevada, respectively. Kennedy (1917)

suggested that it did not occur immediately east of the Sierras, and La Rivers (1940, 1941)

did not record it from Nevada. The original record from Oregon (Selys, 1873) has not been

confirmed, but its occurrence in that state is likely, as it is common at Chico, Butte

County, California (Kennedy, 1917; RWG).

CORDULEGASTRIDAE

Cordulegaster dorsalis Hagen. WASHINGTON, Klickitat Co., Brooks Memorial State

Park, 11 August 1974, 1d 12; Asotin Co., canyon above Grande Ronde River, 2.5 mi. W

Wash. 129, 28 August 1975, 1d (all DRP). No definite previous locality record in Wash-

ington (Paulson, 1970) but to be expected widely in light of its occurrence in southern

British Columbia (Walker, 1958) and western Montana (Bick and Hornuff, 1974).

CORDULIIDAE

Epitheca canis McLachlan. WASHINGTON, King Co., 1 mi. S Duvall, 31 May 1969, 4d 3°

(DRP). This species was not known from any definite locality in this state, although

152 THE PAN-PACIFIC ENTOMOLOGIST

recorded from “Washington Territory” by Muttkowski (1910).

Somatochlora albicincta (Burmeister), OREGON, Klamath Co., Winema National

Forest, Johnson Meadow on Forest Rd. 2519, 23 July 1973, 1d; Lane Co., Campers Lake,

by Ore. 242, 5 mi. SW McKenzie Pass, 4700’, 24 July 1973, 9¢' (both RWG); CALIFORNIA,

Shasta Co., Summit Lake, Lassen Park, August 1952, 19 (CAS); Plumas Co., 0.5 mi. W

Bucks Summit, 4 mi. SW Meadow Valley, 5000’, 20 July 1973, 16 (RWG). Previously

unknown south of the Washington Cascades.

Somatochlora walshi Scudder. WASHINGTON, Kittitas Co., lake at Hyak, 2500’, 12

August 1975, 1d (DRP). Previously known only from Stevens County in northeastern

Washington (Paulson, 1970) but expected in the Cascades from its occurrence in south-

western British Columbia (Walker, 1958).

LIBELLULIDAE

Leucorrhinia glacialis Hagen. WASHINGTON, Pend Oreille Co., Frater Lake, 6 mi. SW

Tiger, 3400’, 22 July 1973, 7d 22; Okanogan Co., Summit Lake on Mt. Hull, 4300’, 2 July

1976, 1d; Kittitas Co., lake at Hyak, 2500’, 12 August 1975, 8c (all DRP). These records,

from the northeast corner, Okanogan highlands and central Cascades, respectively, are

the first for the state, as the previous record (Paulson, 1970) was based on the misidenti-

fication of a female L. intacta. Thus L. glacialis is not known from the western lowlands as

indicated in that paper. It occurs in the Sierra Nevada (Kennedy, 1917) but is not yet

recorded from Oregon.

Libellula croceipennis Selys. CALIFORNIA, Inyo Co., Antelope Spring, N of Cal. 168, 6

July 1974, 12 (RWG); Riverside Co., Riverside, 6 September 1938, 1¢ (LACM); Whitewater,

9 September 1935, 12 (LACM); San Diego Co., Vallecito, 16 September 1945, 106 19

(LACM); Indian Canyon, 26 July 1948, 3d (CIS). Previously known north to Baja California

and Oklahoma but not from Arizona or New Mexico (Needham and Westfall, 1955), it has

subsequently been taken in Maricopa, Pima and Yavapai Counties in Arizona (RWG). The

Inyo County record is well north of the other known localities.

Libellula julia Uhler. CALIFORNIA, Siskiyou Co., Castle Lake, 6000’, 5 August 1953, 1d;

- Tehama Co., Wilson Lake, 26 June 1960, 10 (both CAS). Previously unrecorded south of

Washington.

Libellula luctuosa Burmeister. CALIFORNIA, numerous localities north to Siskiyou

County and west to Napa County, 8 June to 18 August (RWG, DRP, CAS, CIS, LACM).

Surprisingly, this species has not been recorded from the state before although listed by

Smith and Pritchard (1956). Certainly it was not observed by Kennedy (1917) in his travels

through central California. This indicates relatively recent invasion of the state, although

the presence of avery distinctive population of this species centered around the Colorado

River of California and Arizona (Garrison, 1976) may indicate long occupancy of that area.

Libellula nodisticta Hagen. OREGON, Lake Co., Lake Abert, NE side, 16 July 1974, 1d

(RWG). Previous northernmost record Butte County, California (Kennedy, 1917).

Libellula saturata Uhler. OREGON, Josephine Co., Williams Creek, 0.6 mi. NE Provolt

on Ore. 238, 6 July 1974, 1d; pond at Provolt, 6 July 1974, 2d (all DRP). Previous northern-

most record Tehama County, California (Roback and Westfall, 1967).

Macrodiplax balteata (Hagen). CALIFORNIA, Riverside Co., Salton Sea State Park near

Mecca, 23 July 1961, 1¢ (Michigan State University); Whitewater River near Salton Sea, 22

August 1974, 12 (DRP). The published distribution of this species along the Atlantic and

Gulf coasts from North Carolina to Florida and in the West Indies (Needham and Westfall,

1955) and Venezuela (R&acenis, 1953) furnishes no hint that it would occur in California.

However, we have specimens from North Beach, Mazatlan, SINALOA, 31 August 1965

(DRP); San Felipe, BAJA CALIFORNIA, 31 October 1965 (DRP); and Graham, Maricopa and

Yuma Counties, ARIZONA (RWG), indicating a much wider distribution. In addition, DRP

saw a male at an aguada 8.8 mi. N Muna, YUCATAN, on 8 July 1965, indicating a probable

distribution around the Gulf and Caribbean side of Middle America. M. ba/teataisprimarily

coastal and occurs at brackish ponds, which are not.much visited by collectors of

Odonata.

VOL. 53, NO. 2, APRIL 1977 153

Orthemis ferruginea (Fabricius), CALIFORNIA, numerous localities north to Los

Angeles and Riverside Counties, 18 April to 13 November (DRP, CAS, LACM). This is

another of the species listed by Smith and Pritchard (1956) from California with no actual

localities. It has been recorded as far north as Washington County, Utah (Musser, 1962),

and there are two males from Indian Springs, Clark County, NEVADA, collected 25-26 May

1940 (CAS, CIS), afirst record from that state.

Paltothemis lineatipes Karsch. CALIFORNIA, Madera Co., Big Sandy Flat, 15 July 1946,

1d (CAS). In addition DRP saw a male and a female on 18 August 1974 at a locality still

farther north, Calaveritas Creek, 3.1 mi. SE San Andreas, Calaveras County. Previous

northernmost record San Bernardino County (Needham, 1904).

Pantala flavescens (Fabricius). CALIFORNIA, Inyo Co., Little Lake, 3100’, 11 September

1967, 16 (DRP). Also specimens from Los Angeles, Riverside and:Imperial Counties (DRP,

LACM). This species has been listed for California from at least the time of Muttkowski

(1910), but we can find no specific locality records in the literature.

Pantala hymenaea (Say). WASHINGTON, Benton Co., Yakima River, 9 mi. N Benton

City, 12 July 1971, 12; 16 August 1971, 12 (both DRP). In addition DRP has seen

individuals at the following localities: WASHINGTON, Snohomish Co., slough 1 mi. E

Cathcart, 10 August 1971; OREGON, Marion Co., north side of Santiam River at I-5, 13

June 1973; Josephine Co., pond at Provolt, 6 July 1974. Previous northernmost record

Tuolumne County, California (Ahrens, 1938). This species is locally abundant in California

and probably breeds in Washington, as the 16 August specimen listed above was quite

young. It has probably increased greatly in recent years with irrigation of the Central

Valley of California, as Kennedy (1917) listed it from only a single locality in central

California, where it was uncommon.

Perithemis intensa Kirby. CALIFORNIA, Los Angeles Co., Wilson’s Lake, Pasadena, 21

July 1900, 12; Riverside Co., Blythe, 30 July 1935, 12, 12 August 1938, 1d (all LACM). Only

previous record from Californiain Imperial County (Dunkle, 1975).

Sympetrum illotum (Hagen). WASHINGTON, Klickitat Co., pond 10.2 mi. S Brooks

Memorial State Park on U.S. 97, 11 August 1974, 3d (DRP); Yakima Co., 6.8 mi. NW White

Swan, 3 July 1972, 1¢ (Washington State University). Previously known in Washington

from only west of the Cascades (Paulson, 1970). A male from IDAHO, Shoshone Co.,

Wallace, 9 July 1938 (LACM) furnishes a first state record and considerable eastward

range extension. Northern Idaho has much in common biogeographically with western

Washington, and this record further supports the association.

Sympetrum internum Montgomery. WASHINGTON, Snohomish Co., Cedar Ponds Lake,

700’, 17 September 1974, 1d; Whatcom Co., Silver Lake, 3 mi. N Maple Falls, 750’, 26

September 1974, 1d (both DRP). Previously unknown west of the Cascades in Washington

(Paulson, 1970) but recorded from southwestern British Columbia (Whitehouse, 1941).

Tramea onusta Hagen. CALIFORNIA, Inyo Co., Little Lake, 3100’, 20 June 1973, 1d

(DRP). Previous northernmost record Los Angeles County (Calvert, 1906).

Deletions from California List

The following species have been reported from California in the

literature but have not been convincingly documented, although a

few of them may yet be found within its boundaries. Most are listed

for the state by Smith and Pritchard (1956). No specimens from

California of any of these species have been found in several major

collections (University of Florida, Minter Westfall; and University of

Michigan, Leonora Gloyd, both in /itt. to Paulson; as well as the CAS,

CIS and LACM).

154 _ THE PAN-PACIFIC ENTOMOLOGIST

CALOPTERYGIDAE

Calopteryx maculata (Beauvois). First listed for the state by Calvert (1895) on the basis

of aCAS specimen from “California.” The species otherwise is known from eastern North

America west to Manitoba and Texas with an isolated population in Montana and is

unlikely to occur naturally in California (Johnson, 1974).

LESTIDAE

Lestes alacer Hagen. We have found no other reference to this species in California and

are unable to trace the source of Smith and Pritchard’s (1956) listing of it. It is a Middle

American species, known from Texas, New Mexico and Arizona in the United States

(Needham and Heywood, 1929).

COENAGRIONIDAE

Ischnura ramburi (Selys). Agrion defixum Hagen (1861), described from “northern

California,’ was declared a synonym of /. ramburi by Calvert (1895), and this has been the

basis of the subsequent listing of this species from California by other authors

(Muttkowski, 1910; Smith and Pritchard, 1956). The species occurs in southern Baja

California (Calvert, 1895) and in Sonora (DRP) but is not known from the United States

west of Texas other than Hagen’s (1861) record. It seems highly unlikely to occur in

northern Californiaif in the state at all.

AESHNIDAE

Aeshna constricta Say. This species, listed from California and Baja California by

Calvert (1895, 1905), was thought to occur widely in the West until Walker’s (1972) revision

of the genus, in which it was made clear that western records referred to A. pa/mata and A.

umbrosa. Subsequently, Kennedy (1917) described A. wa/keri, which is the species that

occurs in Baja California (CAS), and Paulson (1970) listed the first records of the redefined

constricta from the Pacific states. Although Needham and Westfall (1955) and Smith and

Pritchard (1956) continued to list it from California, we can find no definite records.

Aeshna verticalis Hagen. This species was listed from California by Calvert (1895)

before Walker (1912) showed that western records of it should be attributed to A.

interrupta. Ahrens (1938) again recorded it from California, this time from two specimens

from Yosemite National Park. DRP has examined one of these (Dog Lake, 22 July 1936) in

the collection of the park and found it to be A. interrupta. Thus there is no evidence that

the eastern A. verticalis occurs west of Minnesota and lowa (Needham and Westfall, 1955).

Gynacantha nervosa Rambur. This species has been listed from California by most

faunal list compilers since Calvert (1905) cited a record from “California.” G. nervosa is a

tropical dragonfly, not Known from elsewhere along the Mexico-United States border nor

even from Baja California, and there is no typical breeding habitat (forest swamp) for it in

California. If the specimens listed by Calvert were really from this state they could have

been wanderers or wind-blown individuals from much farther south, but we would prefer

to delete it from the state list until further evidence of its occurrence is at hand.

Triacanthagyna trifida (Rambur). This species parallels the preceding one in its history

in California; although Smith and Pritchard (1956) did not actually list it, they illustrated

its wings and presumably meant to do so. Since Calvert’s (1905) original record,

Williamson (1923) showed that T. trifida itself is restricted to the West Indies and south-

eastern United States, the similar 7. caribbea, T. ditzleri and T. satyrus occurring on the

Middle American mainland, whence any California individuals should originate. All of

these species are unlikely to occurin California, based on the argument presented under

G. nervosa.

VOL. 53, NO. 2, APRIL 1977 155

GOMPHIDAE

Progomphus obscurus (Rambur). Although the California species of Progomphus was

correctly listed as P. borealis by Smith and Pritchard (1956), other authors as recently as

Needham and Westfall (1955) continued to list P. obscurus from the state. The latter is an

eastern species, and there is no recent record of it from west of Texas. Probably the

Oregon, Wyoming and Baja California records in Needham and Westfall (1955) all refer to

borealis.

MACROMIIDAE

Macromia pacifica Hagen. This species was recorded from California by Williamson

(1900), with no further data. The record was questioned by Needham and Westfall (1955)

and Smith and Pritchard (1956), and the species should be deleted from the state list until

further evidence is forthcoming. It has been recorded from Nevada (La Rivers, 1941) but is

otherwise unknown west of Kansas and Texas (Needham and Westfall, 1955). Macromia

are notoriously difficult to determine, and Cruden (1969) has questioned the identification

of specimens of Cordulegaster from Nevada by La Rivers in the same paper. Macromia

magnifica, which occurs widely in the West, is quite variable (Paulson, ms.).

LIBELLULIDAE

Dythemis velox Hagen. This species was listed from California by Muttkowski (1910).

Ris (1916) subsequently showed that the name velox as previously used was a composite

of four species, all of which occur at least as far north as northern Mexico. No member of

this genus has been taken subsequently in California, although D. nigrescens and D.

sterilis occur in Baja California. Until further evidence is at hand, no species of Dythemis

should be considered a member of the odonate fauna of the state.

Erythrodiplax funerea (Hagen). This species is the only one on the “deletion list”

actually represented by a specimen with specific locality data, in this case ‘‘Lagomito”

(probably Lagunita), Stanford University Campus, Santa Clara County (Calvert, 1906). We

suggest the record is in error, as the nearest known populations occur in southernArizona

and Sonora, Mexico (Needham and Westfall, 1955). As discussed under G. nervosa, it is

possible this individual was a wanderer or windblown, but there has been no further

evidence to indicate E. funerea as a resident of the state.

Lepthemis simplicicollis (Say). Both Needham and Westfall (1955) and Smith and

Pritchard (1956) list this species from California, probably based on the listing of the

“subspecies” collocata from the state by various authors (e.g., Muttkowski, 1910). Gloyd

(1958) showed that col/locata and simplicicollis are distinct species, occurring sympatrically

in west Texas and easily distinguishable in the hand. There is no evidence that simpli-

cicollis occurs west of the Great Plains (Paulson, 1970). Walker and Corbet (1975) persisted

in using the name simpliicicollis for British Columbia populations, although they

acknowledged the question as open. Subsequently, Robert Cannings has examined all of

the specimens of this genus collected in British Columbia and determined them as

' collocata (in litt., 12 July 1976).

Pseudoleon superbus (Hagen). First listed by Muttkowski (1910) for California, there is

no further evidence of its occurrence. The confusion about this species, as well as others

previously discussed, may have come about by the labelling of specimens from Baja

California as merely ‘‘California.”’

Discussion

Table 1 lists all the species of Odonata believed by us to occur in

the three Pacific states. Names of species, where they differ from

156 THE PAN-PACIFIC ENTOMOLOGIST

TABLE 1

List of the Odonata of the Pacific States

Wash. Ore. Calif.

CALOPTERYGIDAE

Calopteryx aequabilis x X xX

Hetaerina americana X

LESTIDAE

Archilestes californica xX x

A. grandis xX

Lestes congener x x x

L. disjunctus x X x

L. dryas x x x

L. stultus X

L. unguiculatus x x x

COENAGRIONIDAE

Amphiagrion abbreviatum x x 4

Anomalagrion hastatum x

Argia agrioides x

A. alberta xX

A. emma x X x

A. hinei ; x

A. lugens x x

A. moesta xX

A. nahuana X

A. sedula xX

A. vivida xX xX X

Coenagrion resolutum x xX x

Enallagma anna x x

E. basidens X

E. boreale x x x

E. carunculatum x 4 X

E. civile x

E. clausum x x x

E. cyathigerum xX xX x

E. ebrium x

E. praevarum X

Ischnura barberi x

/, cervula xX x 4

!. denticollis X X

I. erratica X x

!. gemina x

|, perparva x x x

Nehalennia irene xX X

Telebasis salva xX

Zoniagrion exclamationis x

PETALURIDAE

Tanypteryx hageni

AESHNIDAE

Aeshna californica

. canadensis

. constricta

. eremita

. interrupta

juncea

. multicolor

. palmata

. tuberculifera

. umbrosa

. Walkeri x

Anax junius

A. walsinghami

x<

x<

<

x «KK KK KK KK OK

x x KK

>S>PPBBRBRDBD

=<

x<

<x KK XK

VOL. 53, NO. 2, APRIL 1977

List of the Odonata of the Pacific States

GOMPHIDAE

Erpetogomphus compositus

E. lampropeltis

Gomphus graslinellus

G. intricatus

G. kurilis

G. olivaceus

G. plagiatus

Octogomphus specularis

Ophiogomphus bison

O. morrisoni

O. occidentis

O. severus

Progomphus borealis

CORDULEGASTRIDAE

Cordulegaster dorsalis

MACROMIIDAE

Macromia magnifica

CORDULIIDAE

Cordulia shurtleffi

Epitheca canis

E. spinigera

Somatochlora albicincta

S. minor

S. semicircularis

S. walshi

LIBELLULIDAE

Brachymesia furcata

Brechmorhoga mendax

Lepthemis collocata

Leucorrhinia glacialis

L. hudsonica

L. intacta

L. proxima

Libellula comanche

. composita

. croceipennis

. forensis

. julia

. luctuosa

lydia

. nodisticta

. pulchella

. quadrimaculata

. Saturata

. subornata

Macrodiplax balteata

Orthemis ferruginea

Pachydiplax longipennis

Paltothemis lineatipes

Pantala flavescens

P. hymenaea

Perithemis intensa

Sympetrum corruptum

S. costiferum

S. danae

Pr rrrrrreee

TABLE 1 (cont’d.)

Wash.

x x KK XK

Pa

x Kx KKK XK

x KK KK OK

bad

157

Calif.

«x x

<< «KKK KK KK

x<

x KK XK x<

x

x «KK KK XK

x «x «~ «KK KK KKK KK KK KK KK KK KK

158 THE PAN-PACIFIC ENTOMOLOGIST

TABLE 1 (cont’d.)

List of the Odonata of the Pacific States

Wash. Ore. Calif.

S. illotum X xX X

S. internum X X X

S. madidum xX xX X

S. obtrusum X X X

S. occidentale x X xX

S. pallipes x X xX

S. vicinum xX

Tramea lacerata x xX

T. onusta xX

TOTAL SPECIES 70 62 102

common usage in the literature, are based on information to be dis-

cussed elsewhere (Paulson, ms.). Oregon remains relatively terra

incognita, as indicated by the nine species recorded from Washington

and California but not in between. Most of these gaps (all but

Nehalennia irene) have been filled in by the work of Schuh (1936) in his

unpublished thesis on the Odonata of Oregon, but we have left this

work entirely uncited pending examination of his specimens.

That this paper adds to the knowledge of the distribution of 55 of

the 113 species (49%) of Odonata that occur in these states is

indicative of the primitive state of distributional knowledge for these

states; the same is true for most of the West. However, it is clear that

not only were/are there gaps in distribution to be filled in by further

collecting, but also some species are actively extending their ranges.

The only differences we have been able to document are those

between the time of Kennedy (1915, 1917) and our own observations,

but some of these are marked. The following species have certainly

spread in the Pacific states since Kennedy’s time: Enallagma civile,

Libellula luctuosa, Pantala hymenaea and Tramea lacerata. The last

species, not discussed above, was not reported from Washington by

Kennedy (1915) but now occurs in areas he studied intensively in

Yakima County and is abundant in the Columbia Basin (Paulson,

1970, and further observations). All of these changes are probably

consequences of human alteration of the landscape, in particular

damming and irrigating. These activities furnish standing water in

great quantities, and some odonates, in particular the opportunistic

Pantala and Tramea, are rapid colonizers.

A group of species first reported from southeastern California

herein is more difficult to analyze. This group includes Anomalagrion

hastatum, Enallagma basidens, Gomphus plagiatus and Macrodiplax

balteata. All these species are associated with irrigation water, and

VOL. 53, NO. 2, APRIL 1977 159

they may be present because of the extensive modification of the

Colorado River valley. Two alternative hypotheses are less satis-

factory to us: that the species formerly occurred at the Colorado River

itself or that the new records are attributable to insufficient

collecting in the past.

Many species of odonates have become rare or extinct locally in

the Pacific states because of human activities: damming or polluting

rivers, draining marshes, introduction of predators such as trout or

foulers such as carp. Happily, we can find no evidence of any severe

contractions of ranges of any species in this region, although one

species, /schnura gemina of the San Francisco Bay area, has remained

poorly known and may be on the verge of extinction.

We have concentrated considerable effort along the northern

border of Washington and the southern and eastern borders of

California, where we would expect peripheral species to occur. Based

on their known distributions very few additional species might occur

in these border areas. Otherwise, we believe the present list is

reasonably complete and is a good indication of the diversity of the

odonate fauna in this region.

Literature Cited

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States. (Odonata). Entomol. News, 49: 9-16.

Bick, G. H., and L. E. Hornuff. 1972. Odonata collected in Wyoming, South Dakota, and

Nebraska. Proc. Entomol. Soc. Wash., 74: 1-8.

Bick, G. H., and L. E. Hornuff. 1974. New records of Odonata from Montana and

Colorado. Proc. Entomol. Soc. Wash., 76: 90-93.

Brown, C. J. D. 1934. A preliminary list of Utah Odonata. Occ. Pap. Mus. Zool. Univ.

Mich., No. 291: 1-17.

Calvert, P. P. 1895. The Odonata of Baja California, Mexico. Proc. Calif. Acad. Sci. (2),

4: 463-558.

Calvert, P. P. 1901-1908. Biologia Centrali-Americana. Vol. 50: Neuroptera. Porter &

Dulau & Co., 420 pp.

Cruden, R. W. 1969. A new species of Cordulegaster from the Great Basin region of the

United States (Odonata: Cordulegasteridae). Pan-Pacific Entomol. 45: 126-132.

Dunkle, S. W. 1975. New records of North American anisopterous dragonflies. Fla.

Entomol., 58: 117-119.

Garrison, R. W. 1976. Multivariate analysis of geographic variation in Libellula luctuosa

Burmeister (Odonata: Libellulidae). Pan-Pacific Entomol., 52: 181-203.

Gloyd, L. K. 1958. The dragonfly fauna of the Big Bend region of Trans-Pecos Texas.

Occ. Pap. Mus. Zool. Univ. Mich., No. 593: 1-23.

Hagen, H. A. 1861. Synopsis of the Neuroptera of North America. Smithsonian Misc.

Coll., 347 pp.

Johnson, C. 1972. The damselflies (Zygoptera) of Texas. Bull. Fla. State Mus., Biol.

Sci., 16: 55-128.

Johnson, C. 1974. Taxonomic keys and distributional patterns for Nearctic species of

Calopteryx damselflies. Fla. Entomol., 57: 231-248.

Kennedy, C. H. 1915. Notes on the life history and ecology of the dragonflies (Odonata)

of Washington and Oregon. Proc. U.S. Nat. Mus., 49: 259-345.

Kennedy, C. H. 1917. Notes on the life history and ecology of the dragonflies (Odonata)

of central California and Nevada. Proc. U.S. Nat. Mus., 52: 483-635.

160 THE PAN-PACIFIC ENTOMOLOGIST

La Rivers, |. 1940. A preliminary synopsis of the dragonflies of Nevada. Pan-Pacific

Entomol., 16: 111-123.

La Rivers, |. 1941. Additions to the list of Nevada dragonflies. Entomol. News, 52: 126-

130, 155-157.

Montgomery, B. E. 1966. Distribution of Odonata, pp. 348-349, in Chandler, L., The

origin and composition of the insect fauna, Chapt. 20, Natural Features of

Indiana, Indiana Acad. Sci.

Musser, R. J. 1962. Dragonfly nymphs of Utah (Odonata: Anisoptera). Univ. Utah Biol.

Ser., 12: 1-66.

Muttkowski, R. A. 1910. Catalogue of the Odonata of North America. Bull. Publ. Mus.

Milwaukee, 1: 1-207.

Needham, J. G. 1904. New dragon-fly nymphs in the United States National Museum.

Proc. U.S. Nat. Mus., 27: 685-720.

Needham, J. G. and H. B. Heywood. 1929. A handbook of the dragonflies of North

America. Charles C. Thomas, 378 pp.

Needham, J. G. and M. J. Westfall, Jr. 1955. A manual of the dragonflies of North

America (Anisoptera). Univ. California Press, 615 pp.

Paulson, D.R. 1970. A list of the Odonata of Washington with additions to and deletions

from the state list. Pan-Pacific Entomol., 46: 194-198.

Racenis, J. 1953. Contribucion al estudio de los Odonata de Venezuela. An. Univ. Centr.

Venez., 35: 31-96.

Ris, F. 1909-1919. Libellulines. Cat. Systematique et Descriptif des Coll. Zool. Edm. de

Selys Longchamps, Fasc. 9-16, pp. 1-1278.

Roback, S. S., and M. J. Westfall, Jr. 1967. New records of Odonata nymphs from the

United States and Canada with water quality data. Trans. Amer. Entomol. Soc.,

93: 101-124.

Schuh, J. 1936. A contribution to the knowledge of the Odonata of Oregon. Unpubl.

M. S. thesis, Oregon State University.

Seemann, T. M. 1927. Dragonflies, mayflies and stoneflies of southern California. J.

Entomol. Zool. Pomona Coll., 19: 1-69.

Selys Longchamps, E. de. 1873. Troisiemes additions au Synopsis des Gomphines.

Bull. Acad. Roy. Belg. (2) 35: 732-774.

Smith, R. F., and A. E. Pritchard. 1956. Odonata, pp. 106-153 in Usinger, R. L., ed., Aquatic

Insects of California. Univ. California Press, 508 pp. ,

Walker, E. M. 1912. The North American dragonflies of the genus Aeshna. Univ.

Toronto Stud., Biol. Ser., No. 11: 1-213.

Walker, E. M. 1958. The Odonata of Canada and Alaska. Vol. 2. Univ. Toronto Press, 318

Pp.

Walker, E. M. and P. S. Corbet. 1975. The Odonata of Canada and Alaska. Vol. 3. Univ.

Toronto Press, 307 pp.

Whitehouse, F. C. 1941. British Columbia dragonflies (Odonata), with notes on distri-

bution and habits. Amer. Midland Nat., 26: 488-557.

Williamson, E. B. 1900. The dragonflies of Indiana. Indiana Dept. Geol. & Nat. Res.,

24th Ann. Report: 229-333.

Williamson, E. B. 1923. Notes on American species of Triacanthagyna and Gynacantha.

Misc. Publ. Mus. Zool. Univ. Mich., No. 9: 1-80.

January issue mailed June 23,1977

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